Bottlenose Dolphin (<I>Tursiops truncatus</I>) Double-Slit Pupil Asymmetries Enhance Vision

Abstract

Geometries of the iris, retinal cell distributions, and the optical characteristics of the lens and cornea have evolved to optimize the visual adaptations of the bottlenose dolphin (Tursiops truncatus) to the oceanic environment. Under high ambient light conditions, the operculum of the iris shields the lens and forms two asymmetrical slit pupils. Under these conditions, light entering the eye is channeled and focused onto the two areas of the retina having a finer retinal mosaic of ganglion cells (typically associated with higher image resolution). The paths of light determined by tracing rays in the reverse direction through these pupils coincide with a dolphin’s behaviorally observed preferred viewing directions. These rays aid in determining the interdependence between the graininess of the retinal mosaic and resolution spot sizes in the object space. For oblique forward and downward viewing directions in air, the larger temporal pupil admits light which passes through the weakly refractive margin of a bifocal lens, counterbalancing the optically strong cornea in air. In water, light passing through the optically strong lens core is focused from a wide lateral and downward field-of-vision. Although other explanations for comparable aerial and underwater vision remain plausible, a dolphin eye model incorporating a bifocal lens offers an explanation consistent with ophthalmoscopic refractive state measurements. The model is also consistent with visual acuity study results conducted in air and in water under both high and low ambient light levels. From insight gained after applying a common data analysis technique to visual acuity studies conducted by other researchers and tracing oblique rays through the asymmetric double-slit pupils, a re-examination of explanatory hypotheses for the paradoxical observations of comparable aerial and underwater vision is presented. Based in part on these findings and supportive evidence from dolphin vision researchers, the unique distinguishing characteristics of dolphin vision are summarized.