What is unique about the human eye? Comparative image analysis on the external eye morphology of human and nonhuman great apes

The gaze-signaling hypothesis and the related cooperative-eye hypothesis posit that humans have evolved special external eye morphology, including exposed white sclera (the white of the eye), to enhance the visibility of eye-gaze direction and thereby facilitate conspecific communication through joint-attentional interaction and ostensive communication. However, recent quantitative studies questioned these hypotheses based on new findings that humans are not necessarily unique in certain eye features compared to other great ape species. Therefore, there is currently a heated debate on whether external eye features of humans are distinguished from those of other apes and how such distinguished features contribute to the visibility of eye-gaze direction. This study leveraged updated image analysis techniques to test the uniqueness of human eye features in facial images of great apes. Although many eye features were similar between humans and other species, a key difference was that humans have uniformly white sclera which creates clear visibility of both eye outline and iris –the two essential features contributing to the visibility of eye-gaze direction. We then tested the robustness of the visibility of these features against visual noises such as darkening and distancing and found that both eye features remain detectable in the human eye, while eye outline becomes barely detectable in other species under these visually challenging conditions. Overall, we identified that humans have distinguished external eye morphology among other great apes, which ensures robustness of eye-gaze signal against various visual conditions. Our results support and also critically update the central premises of the gaze-signaling hypothesis. ### Competing Interest Statement The authors have declared no competing interest.