Experimental evidence for the gaze-signaling hypothesis: White sclera enhances the visibility of eye gaze direction in humans and chimpanzees

Experimental evidence for the gaze-signaling hypothesis: White sclera enhances the visibility of eye gaze direction in humans and chimpanzees. Fumihiro Kano, Yuri Kawaguchi, Hanling Yeow. bioRxiv, Sep 21 2021. https://doi.org/10.1101/2021.09.21.461201

Abstract: Hallmark social activities of humans, such as cooperation and cultural learning, involve eye-gaze signaling through joint attentional interaction and ostensive communication. The gaze-signaling and related cooperative-eye hypotheses posit that humans evolved unique external eye morphology, including exposed white sclera (the white of the eye), to enhance the visibility of eye-gaze for conspecifics. However, experimental evidence is still lacking. This study tested the ability of human and chimpanzee participants to detect the eye-gaze directions of human and chimpanzee images in computerized tasks. We varied the level of brightness and size in the stimulus images to examine the robustness of the eye-gaze directional signal against visually challenging conditions. We found that both humans and chimpanzees detected gaze directions of the human eye better than that of the chimpanzee eye, particularly when eye stimuli were darker and smaller. Also, participants of both species detected gaze direction of the chimpanzee eye better when its color was inverted compared to when its color was normal; namely, when the chimpanzee eye has artificial white sclera. White sclera thus enhances the visibility of eye-gaze direction even across species, particularly in visually challenging conditions. Our findings supported but also critically updated the central premises of the gaze-signaling hypothesis.

Final version: What is unique about the human eye? Comparative image analysis on the external eye morphology of human and nonhuman great apes. Fumihiro Kano et al. Evolution and Human Behavior, December 29 2021. https://doi.org/10.1016/j.evolhumbehav.2021.12.004

Abstract: 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 certain features of human eyes are not necessarily unique among great ape species. Accordingly, there is currently a heated debate over whether external eye features of humans are distinct from those of other apes and how such distinguishable features contribute to the visibility of eye-gaze direction. The present 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 great apes, a key difference was that humans have uniformly white sclera which creates clear visibility of both the 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 noise, such as shading 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 unique external eye morphology among other great apes, which ensures the robustness of eye-gaze signals in various visual conditions. Our results support and also critically update the central premises of the gaze-signaling hypothesis.

Keywords: Eye colorCommunicationComparative analysisHuman evolutionGreat apeScleraGaze detection