The evolution of the eye has been a subject of significant study, as a distinctive example of a analogous organ present in a wide variety of taxa. Complex, image-forming eyes evolved independently some 50 to 100 times. Complex eyes appear to have first evolved within a few million years, in the rapid burst of evolution known as the Cambrian explosion. There is no evidence of eyes before the Cambrian, but a wide range of diversity is evident in the Middle Cambrian Burgess shale, and the slightly older Emu Bay Shale. Eyes show a wide range of adaptations to meet the requirements of the organisms which bear them. Eyes vary in their visual acuity, the range of wavelengths they can detect, their sensitivity in low light levels, their ability to detect motion or resolve objects, and whether they can discriminate colours.
Rate of evolution
The first fossils of eyes that have been found to date are from the lower Cambrian period (about 540 million years ago). This period saw a burst of apparently rapid evolution, dubbed the "Cambrian explosion". One of the many hypotheses for "causes" of this diversification, the "Light Switch" theory of Andrew Parker, holds that the evolution of eyes initiated an arms race that led to a rapid spate of evolution. Earlier than this, organisms may have had use for light sensitivity, but not for fast locomotion and navigation by vision.
It is difficult to estimate the rate of eye evolution because the fossil record, particularly of the Early Cambrian, is poor. The evolution of a circular patch of photoreceptor cells into a fully functional vertebrate eye has been approximated based on rates of mutation, relative advantage to the organism, and natural selection. Based on pessimistic calculations that consistently overestimate the time required for each stage and a generation time of one year, which is common in small animals, it has been proposed that it would take less than 364,000 years for the vertebrate eye to evolve from a patch of photoreceptors.
Stages of eye evolution
The earliest predecessors of the eye were photoreceptor proteins that sense light, found even in unicellular organisms, called "eyespots". Eyespots can only sense ambient brightness: they can distinguish light from dark, sufficient for photoperiodism and daily synchronization of circadian rhythms. They are insufficient for vision, as they cannot distinguish shapes or determine the direction light is coming from. Eyespots are found in nearly all major animal groups, and are common among unicellular organisms, including euglena. The euglena's eyespot, called a stigma, is located at its anterior end. It is a small splotch of red pigment which shades a collection of light sensitive crystals. Together with the leading flagellum, the eyespot allows the organism to move in response to light, often toward the light to assist in photosynthesis,[19] and to predict day and night, the primary function of circadian rhythms. Visual pigments are located in the brains of more complex organisms, and are thought to have a role in synchronising spawning with lunar cycles. By detecting the subtle changes in night-time illumination, organisms could synchronise the release of sperm and eggs to maximise the probability of fertilisation
Vision itself relies on a basic biochemistry which is common to all eyes. However, how this biochemical toolkit is used to interpret an organism's environment varies widely: eyes have a wide range of structures and forms, all of which have evolved quite late relative to the underlying proteins and molecules.
At a cellular level, there appear to be two main "designs" of eyes, one possessed by the protostomes (molluscs, annelid worms and arthropods), the other by the deuterostomes (chordates and echinoderms).