Evolution of the eye
The evolution of the eye is the origin and development with diversification by natural selection over geological time of organs of photosensitivity and vision in living organisms. Many scientists have found the evolution of the eye attractive to study because the eye distinctively exemplifies an analogous organ found in many animal forms. Simple light detection is found in bacteria, single-celled organisms, plants and animals. Complex, image-forming eyes have evolved independently several times.
Diverse eyes are known from the Burgess Shale of the Middle Cambrian, and from the slightly older Emu Bay Shale.
Eyes vary in their visual acuity, the range of wavelengths they can detect, their sensitivity in no light, their ability to detect motion or to resolve objects, and whether they can discriminate colours.
History of research
In 1802, philosopher William Paley called it a miracle of "design." In 1859, Charles Darwin himself wrote in his Origin of Species, that the evolution of the eye by natural selection seemed at first glance "absurd in the highest possible degree".However, he went on that despite the difficulty in imagining it, its evolution was perfectly feasible:
... if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist, each grade being useful to its possessor, as is certainly the case; if further, the eye ever varies and the variations be inherited, as is likewise certainly the case and if such variations should be useful to any animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection,
though insuperable by our imagination, should not be considered as subversive of the theory.
He suggested a stepwise evolution from "an optic nerve merely coated with pigment, and without any other mechanism" to "a moderately high stage of perfection", and gave examples of existing intermediate. Current research is investigating the genetic mechanisms underlying eye development and evolution.
Rate of evolution
The first possible fossils of eyes found to date are from the Ediacaran period, while the oldest certain fossilized eye is from a Schmidtiellus reetae fossil from 530 mya, collected in Saviranna in northern Estonia. The structure is similar to the compound eyes of modern-day dragonflies and bees, but with ommatidia spaced further apart, and without a lens. The lower Cambrian had a burst of apparently rapid evolution, called the "Cambrian explosion". One of the many hypotheses for "causes" of the Cambrian explosion is the "Light Switch" theory of Andrew Parker: it holds that the evolution of advanced eyes started an arms race that accelerated evolution. Before the Cambrian explosion, animals may have sensed light, but did not use it for fast locomotion or navigation by vision.The rate of eye evolution is difficult to estimate because the fossil record, particularly of the lower Cambrian, is poor. How fast a circular patch of photoreceptor cells can evolve into a fully functional vertebrate eye has been estimated based on rates of mutation, relative advantage to the organism, and natural selection. However, the time needed for each state was consistently overestimated and the generation time was set to one year, which is common in small animals. Even with these pessimistic values, the vertebrate eye could still evolve from a patch of photoreceptor cells in less than 364,000 years.
Origins of the eye
Whether the eye evolved once or many times depends on the definition of an eye. All eyed animals share much of the genetic machinery for eye development. This suggests that the ancestor of eyed animals had some form of light-sensitive machinery – even if it was not a dedicated optical organ. However, even photoreceptor cells may have evolved more than once from molecularly similar chemoreceptor cells. Probably, photoreceptor cells existed long before the Cambrian explosion. Higher-level similarities – such as the use of the protein crystallin in the independently derived cephalopod and vertebrate lenses – reflect the co-option of a more fundamental protein to a new function within the eye.A shared trait common to all light-sensitive organs are opsins. Opsins belong to a family of photo-sensitive proteins and fall into nine groups, which already existed in the urbilaterian, the last common ancestor of all bilaterally symmetrical animals. Additionally, the genetic toolkit for positioning eyes is shared by all animals: The PAX6 gene controls where eyes develop in animals ranging from octopuses to mice and fruit flies. Such high-level genes are, by implication, much older than many of the structures that they control today; they must originally have served a different purpose, before they were co-opted for eye development.
Eyes and other sensory organs probably evolved before the brain: There is no need for an information-processing organ before there is information to process. A living example are cubozoan jellyfish that possess eyes comparable to vertebrate and cephalopod camera eyes despite lacking a brain.
Stages of evolution
The earliest predecessors of the eye were photoreceptor proteins that sense light, found even in unicellular organisms, called "eyespots". Eyespots can sense only 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, 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 types of eyes, one possessed by the protostomes, the other by the deuterostomes.
The functional unit of the eye is the photoreceptor cell, which contains the opsin proteins and responds to light by initiating a nerve impulse. The light sensitive opsins are borne on a hairy layer, to maximise the surface area. The nature of these "hairs" differs, with two basic forms underlying photoreceptor structure: microvilli and cilia. In the eyes of protostomes, they are microvilli: extensions or protrusions of the cellular membrane. But in the eyes of deuterostomes, they are derived from cilia, which are separate structures. However, outside the eyes an organism may use the other type of photoreceptor cells, for instance the clamworm Platynereis dumerilii uses microvilliar cells in the eyes but has additionally deep brain ciliary photoreceptor cells. The actual derivation may be more complicated, as some microvilli contain traces of cilia – but other observations appear to support a fundamental difference between protostomes and deuterostomes. These considerations centre on the response of the cells to light – some use sodium to cause the electric signal that will form a nerve impulse, and others use potassium; further, protostomes on the whole construct a signal by allowing more sodium to pass through their cell walls, whereas deuterostomes allow less through.
This suggests that when the two lineages diverged in the Precambrian, they had only very primitive light receptors, which developed into more complex eyes independently.
Early eyes
The basic light-processing unit of eyes is the photoreceptor cell, a specialized cell containing two types of molecules bound to each other and located in a membrane: the opsin, a light-sensitive protein; and a chromophore, the pigment that absorbs light. Groups of such cells are termed "eyespots", and have evolved independently somewhere between 40 and 65 times. These eyespots permit animals to gain only a basic sense of the direction and intensity of light, but not enough to discriminate an object from its surroundings.Developing an optical system that can discriminate the direction of light to within a few degrees is apparently much more difficult, and only six of the thirty-some phyla possess such a system. However, these phyla account for 96% of living species.
These complex optical systems started out as the multicellular eyepatch gradually depressed into a cup, which first granted the ability to discriminate brightness in directions, then in finer and finer directions as the pit deepened. While flat eyepatches were ineffective at determining the direction of light, as a beam of light would activate exactly the same patch of photo-sensitive cells regardless of its direction, the "cup" shape of the pit eyes allowed limited directional differentiation by changing which cells the lights would hit depending upon the light's angle. Pit eyes, which had arisen by the Cambrian period, were seen in ancient snails, and are found in some snails and other invertebrates living today, such as planaria. Planaria can slightly differentiate the direction and intensity of light because of their cup-shaped, heavily pigmented retina cells, which shield the light-sensitive cells from exposure in all directions except for the single opening for the light. However, this proto-eye is still much more useful for detecting the absence or presence of light than its direction; this gradually changes as the eye's pit deepens and the number of photoreceptive cells grows, allowing for increasingly precise visual information.
When a photon is absorbed by the chromophore, a chemical reaction causes the photon's energy to be transduced into electrical energy and relayed, in higher animals, to the nervous system. These photoreceptor cells form part of the retina, a thin layer of cells that relays visual information, including the light and day-length information needed by the circadian rhythm system, to the brain. However, some jellyfish, such as Cladonema, have elaborate eyes but no brain. Their eyes transmit a message directly to the muscles without the intermediate processing provided by a brain.
During the Cambrian explosion, the development of the eye accelerated rapidly, with radical improvements in image-processing and detection of light direction.
File:Nautilus pompilius.jpg|thumb|The nautilus eye functions similarly to a pinhole camera.
After the photosensitive cell region invaginated, there came a point when reducing the width of the light opening became more efficient at increasing visual resolution than continued deepening of the cup. By reducing the size of the opening, organisms achieved true imaging, allowing for fine directional sensing and even some shape-sensing. Eyes of this nature are currently found in the nautilus. Lacking a cornea or lens, they provide poor resolution and dim imaging, but are still, for the purpose of vision, a major improvement over the early eyepatches.
Overgrowths of transparent cells prevented contamination and parasitic infestation. The chamber contents, now segregated, could slowly specialize into a transparent humour, for optimizations such as colour filtering, higher refractive index, blocking of ultraviolet radiation, or the ability to operate in and out of water. The layer may, in certain classes, be related to the moulting of the organism's shell or skin. An example of this can be observed in Onychophorans where the cuticula of the shell continues to the cornea. The cornea is composed of either one or two cuticular layers depending on how recently the animal has moulted. Along with the lens and two humors, the cornea is responsible for converging light and aiding the focusing of it on the back of the retina. The cornea protects the eyeball while at the same time accounting for approximately 2/3 of the eye's total refractive power.
It is likely that a key reason eyes specialize in detecting a specific, narrow range of wavelengths on the electromagnetic spectrum—the visible spectrum—is that the earliest species to develop photosensitivity were aquatic, and water filters out electromagnetic radiation except for a range of wavelengths, the shorter of which we refer to as blue, through to longer wavelengths we identify as red. This same light-filtering property of water also influenced the photosensitivity of plants.