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Putting a date on the first appearance of eyes depends on what one means by eye. If the term refers to a multicellular organ, even if it has just a few cells, then by definition, eyes could not form before there were multicellular animals. But many protists (animal-like, plantlike, or fungus-like unicellular organisms that require a water-based environment) can detect light by using aggregations of pigment molecules, and they use this information to modify their metabolic activity or motility (the ability to move spontaneously and independently). One of the familiar living examples, probably known to anyone who has taken a biology class, is the aquatic protozoan Euglena, which has an eyespot near its motile flagellum (hairlike structure). Some living protists are very like their ancestral forms embedded in ancient sedimentary rocks, and this similarity suggests that the ability to detect light and modify behavior in response to light has been around for a very long time. Animals arose from one of such unicellular creatures, perhaps from one already specialized for a primitive kind of vision. An eye is a collection of cells that are specialized for light detection through the presence of photosensitive pigment as well as a means of restricting the direction of incoming light that will strike the photosensitive cells. This definition says nothing about image formation, lenses, eye movements, or any of the other features we associate with our own eyes, but it does recognize the simplest form of functional and anatomical specialization namely, detection of light. Everything else can be built up from this simple beginning, and some animals appear to have had eyes almost from the beginning of the animal kingdom. Animals were scarce 600 million years ago in the geological era called the Precambrian. There are very few fossil remains from that time (though more keep turning up), and most evidence of the presence of animals is indirect, such as small tunnels in rock that could be ancient worm burrowings. But just 50 million years or so later, fossilized bits and pieces of animals abound, suggesting that a great burst of evolutionary creativity occurred in the 50-million-year interval. This surge of new life, marked by an abundance of animals, is called the Cambrian explosion. The first direct evidence for the early origin of eyes comes from fossils that are about 530 million years old, a time shortly after the Cambrian explosion; they were found on a mountainside in British Columbia in a deposit known as the Burgess Shale. The Burgess Shale fossils are extraordinarily important because among them are remains of soft-bodied creatures, many of them lacking shells and other hard parts that fossilize easily. Consequently, their preservation is little short of miraculous (as are the delicate methods used to reconstruct three-dimensional structure from these flattened fossils), and they are one of the few known repositories of early soft-bodied animals. Not all of the Burgess animals had eyes. However, some did. (Gross features location, size, and hemispheric shape are responsible for the designation of some structures as eyes). The reconstructed eyes of these Burgess animals look superficially like eyes of some living crustaceans, particularly those of shrimp and crabs whose eyes are mounted on stalks that improve the range of vision by raising the eyes above the surface of the head. The eyes of some Burgess organisms sat on stalks; those of others were on or a part of the body surface. One animal, Opabinia, had five eyes: two lateral pairs and a single medial eye; at least one of the lateral pairs had stalks that could have been movable. And some trilobite-like animals in the Burgess Shale had faceted eyes much like those of later fossil trilobites. Although the presence of eyes on some of the Burgess animals indicates that eyes have been around for a very long time, it is unlikely that these were the first eyes; they seem much too large and (potentially) well developed to be brand new inventions. The best we can do is put the origin of eyes somewhere between the beginning of the Cambrian explosion, about 600 million years ago, and the death of the Burgess animals, some 530 million years ago.