Eye evolution is driven by the evolution of visually guided behavior.

Accumulation of gradually more demanding behaviors have

continuously increased the performance requirements on the
photoreceptor organs. Starting with nondirectional photoreception, I
argue for an evolutionary sequence continuing with directional
photoreception, low-resolution vision, and finally, high-resolution
vision. Calculations of the physical requirements for these four
sensory tasks show that they correlate with major innovations in eye
evolution and thus work as a relevant classification for a functional
analysis of eye evolution. Together with existing molecular and
morphological data, the functional analysis suggests that urbilateria
had a simple set of rhabdomeric and ciliary receptors used for
directional photoreception, and that organ duplications, positional
shifts and functional shifts account for the diverse patterns of eyes and
photoreceptors seen in extant animals. The analysis also suggests that
directional photoreception evolved independently at least twice
before the last common ancestor of bilateria and proceeded several
times independently to true vision in different bilaterian and cnidarian
groups. This scenario is compatible with Pax-gene expression in eye
development in the different animal groups. The whole process from
the first opsin to high-resolution vision took about 170 million years
and was largely completed by the onset of the Cambrian, about 530
million years ago. Evolution from shadow detectors to multiple
directional photoreceptors has further led to secondary cases of eye
evolution in bivalves, fan worms, and chitons.