Perception of Fourier and non- Fourier motion by larval zebrafish

articles Zebrafish larvae innately begin responding to moving stimuli shortly after hatching. In their optomotor response, which is elicited by large moving stimuli presented from below or the side 1,2 , larvae swim in the direction of perceived motion. The distance they travel in a given time indicates the effectiveness of the stimulus. By observing the response of many larvae to computer-animated displays, we could determine which attributes of a moving stimulus the zebrafish visual system detects. If luminance-defined features drift smoothly or jump in space, they can produce strong sensations of motion. A number of proposed models explain how motion information can be extracted. In a simple model, a point-to-point comparison is made between the luminance pattern and a spatially displaced copy of the pattern that was seen a short time before 3 . The displacement that gives the best fit tells the brain the direction and speed of movement. A more complex strategy is to look at the Fourier motion energy in the visual scene Although there is evidence that humans can use both feature matching and motion energy to detect movement 7 , they may also sense motion when presented with stimuli in which only secondorder features such as contrast, texture or flicker are moving Here we find that the fish larvae detect moving features of visu- A moving grating elicits innate optomotor behavior in zebrafish larvae; they swim in the direction of perceived motion. We took advantage of this behavior, using computer-animated displays, to determine what attributes of motion are extracted by the fish visual system. As in humans, first-order (luminance-defined or Fourier) signals dominated motion perception in fish; edges or other features had little or no effect when presented with these signals. Humans can see complex movements that lack first-order cues, an ability that is usually ascribed to higher-level processing in the visual cortex. Here we show that second-order (non-Fourier) motion displays induced optomotor behavior in zebrafish larvae, which do not have a cortex. We suggest that second-order motion is extracted early in the lower vertebrate visual pathway. al stimuli in a way that is qualitatively similar to humans: both firstorder and second-order cues drive their behavioral response. Our demonstration of second-order motion detection in fish challenges the idea that higher-level, cortical mechanisms are necessary to explain this capacity of the visual system. RESULTS Optomotor responses to Fourier motion The assay used to measure optomotor responses is similar to the one described previously 2 (Methods). Movies showing drifting gratings evoke strong optomotor responses in almost all fish in a clutch. Fish do not respond to a moving grating with a stripe width narrower than approximately 9°, which is slightly less than the predicted resolution limit of the larval cone mosaic, 6°at this age In the following experiments, responses were normalized to the effect of a designated strong stimulus, a 100% contrast square wave subtending 100°of visual angle per cycle and moving at 1 Hz for 30 seconds Although the fish seemed to follow a motion signal in the movies, it was possible that they were tracking features such as light or dark regions or edges that were being displaced. We did an experiment to show that the optomotor response is truly a response to motion. A motion display was shown of a sine wave grating tha

Effects of luminance and contrast on direction of ambiguous apparent motion

A study is reported of the role of luminance and contrast in resolving ambiguous apparent motion (AM). Different results were obtained for the short-range (SR) and the long-range (LR) motion-detecting processes. For short-range jumps (7.5 min arc), the direction of ambiguous AM depended on brightness polarity, with AM only from white to white and from black to black. But for larger jumps, or when an interstimulus interval (ISI) was introduced, AM was less dependent on polarity, with white often jumping to black and black jumping to white. Two potential AMs were pitted against each other, one carried by a light stimulus and the other by a dark stimulus. The stimulus whose luminance differed most from the uniform surround captured the AM. Visual response to luminance was linear, not logarithmic. When the stimulus was modified to give continuous AM in one direction it was followed by a negative aftereffect of motion only when the spatial displacement was 1 min arc. A larger displacement (10 min arc) gave good AM but no motion aftereffect. Thus only short-range motion adapts motion-sensitive channels

Luminance processing in apparent motion, Vernier offset and stereoscopic depth

We obtained (apparently) linear responses to luminance from three special displays of apparent motion, Vernier offset and stereoscopic depth. In our motion stimulus a dark and a light bar exchanged luminances repetitively on a grey surround. Motion was attributed to the bar that differed more from the surround, that is, on a dark surround the light bar appeared to jump, and on a light surround the dark bar appeared to jump. The apparent motion disappeared when the luminance of the surround lay halfway between that of the bars - on a linear, not a logarithmic scale. Similar results were obtained for special Vernier offset and stereo stimuli. These results cannot be explained if all luminances are processed within the same luminance pathway and that pathway transforms input luminance using non-linear compression. However, the apparent linearity of our results could arise from opposite and equal non-linearities cancelling out within separate ON- and OFF-spatial luminance pathways. A second set of experiments presented one bar separately into each eye on different surrounds (dichoptic presentation of competing apparent motion signals) or manipulated the display spatially so that different surrounds were associated with different bars (binocular presentation of competing Vernier targets). Results showed that apparent motion and Vernier signals of equal Weber contrast (normalisation of linear difference to surround luminance) evoked equal-motion and equal Vernier offset strengths. Given that motion and Vernier strength followed Weber's law, we infer that the ON- and OFF-pathways transform luminance non-linearly. Our third experiment presents an example of a brightness bisection task in which we were able to influence the bisection steps, to follow either a linear or non-linear series. The benefits of parsing the visual scene so that visual information is processed within two opposite luminance pathways is discussed. Copyright (C) 1999 Elsevier Science Ltd