Eye movements of vertebrates and their relation to eye form and function.

The types of eye movements shown by all vertebrates originated in the earliest fishes. These consisted of compensatory movements, both vestibular and visual, to prevent image motion, and saccades to relocate gaze. All vertebrates fixate food items with their heads to enable ingestion, but from teleosts onwards some species also use eye movements to target particular objects, especially food. Eye movement use is related to the resolution distribution in the retina, with eyes that contain foveas, or areas of high ganglion cell density, being more likely to make targeting eye movements, not seen in animals with more uniform retinas. Birds, in particular, tend mainly to use head movements when shifting gaze. Many birds also make translatory head saccades (head bobbing) when walking. It is common for animals to use both eyes when locating food items ahead, but the use of binocular disparity for distance judgment is rare, and has only been demonstrated in toads, owls, cats and primates. Smooth tracking with eyes alone is probably confined to primates. The extent of synchrony and directional symmetry in the movements of the two eyes varies greatly, from complete independence in the sandlance and chameleon, to perfect coordination in primates.