Elaborate pupils in skates may help camouflage the eye

The little skate Leucoraja erinacea has elaborately shaped pupils, whose characteristics and functions have not been studied extensively. It has been suggested that such pupil shapes may camouflage the eye; yet, no experimental evidence has been presented to support this claim. Skates are bottom-dwellers that often bury into the substrate with their eyes protruding. If these pupils serve any camouflage function, we expect there to be a pupillary response related to the spatial frequency (\"graininess\") of the background against which the eye is viewed. Here, we tested whether skate pupils dilate or constrict in response to background spatial frequency. We placed skates on background substrates with different spatial frequencies and recorded pupillary responses at three light intensities. In experiment 1, the skates pupillary response to three artificial checkerboards of different spatial frequencies was recorded. Skates responded to changing light intensity with pupil dilation/constriction; yet, their pupils did not change in response to spatial frequency. In experiment 2, in which skates could bury into three natural substrates with different spatial frequencies, such that their eyes protruded above the substrate, the pupils showed a subtle but statistically significant response to changes in substrate spatial frequency. Given the same light intensity, the smaller the spatial frequency of the natural substrate, the more constricted the pupil. While light intensity is the primary factor determining pupil dilation, these experiments are the first to show that pupils also change in response to background spatial frequency, which suggests that the pupil may aid in camouflaging the eye.