Eyes are
organs
In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a fu ...
of the
visual system
The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the a ...
. They provide living organisms with
vision
Vision, Visions, or The Vision may refer to:
Perception Optical perception
* Visual perception, the sense of sight
* Visual system, the physical mechanism of eyesight
* Computer vision, a field dealing with how computers can be made to gain un ...
, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect
light
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 tera ...
and convert it into electro-chemical impulses in
neurons
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. N ...
(neurones). In higher organisms, the eye is a complex
optical
Optics is the branch of physics that studies the behaviour and properties of light, including its interactions with matter and the construction of instruments that use or detect it. Optics usually describes the behaviour of visible, ultraviole ...
system which collects light from the surrounding environment, regulates its intensity through a
diaphragm
Diaphragm may refer to:
Anatomy
* Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen
* Pelvic diaphragm or pelvic floor, a pelvic structure
* Urogenital diaphragm or triangular ligament, a pelvic structure
Other
* Diap ...
,
focuses it through an adjustable assembly of
lenses
A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
to form an
image
An image is a visual representation of something. It can be two-dimensional, three-dimensional, or somehow otherwise feed into the visual system to convey information. An image can be an artifact, such as a photograph or other two-dimensiona ...
, converts this image into a set of electrical signals, and transmits these signals to the
brain
A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
through complex neural pathways that connect the eye via the
optic nerve
In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual system, visual information from the retina to the brain. In humans, the optic nerve i ...
to the
visual cortex
The visual cortex of the brain is the area of the cerebral cortex that processes visual information. It is located in the occipital lobe. Sensory input originating from the eyes travels through the lateral geniculate nucleus in the thalamus and ...
and other areas of the brain. Eyes with resolving power have come in ten fundamentally different forms, and 96% of
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
species possess a complex optical system.
Image-resolving eyes are present in
molluscs
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
,
chordates
A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...
and
arthropods
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
.
The most simple eyes, pit eyes, are eye-spots which may be set into a pit to reduce the angle of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light.
From more complex eyes, retinal
photosensitive ganglion cell
Intrinsically photosensitive retinal ganglion cells (ipRGCs), also called photosensitive retinal ganglion cells (pRGC), or melanopsin-containing retinal ganglion cells (mRGCs), are a type of neuron in the retina of the mammalian eye. The presence ...
s send signals along the
retinohypothalamic tract
In neuroanatomy, the retinohypothalamic tract (RHT) is a photic neural input pathway involved in the circadian rhythms of mammals. The origin of the retinohypothalamic tract is the intrinsically photosensitive retinal ganglion cells (ipRGC), whi ...
to the
suprachiasmatic nuclei
The suprachiasmatic nucleus or nuclei (SCN) is a tiny region of the brain in the hypothalamus, situated directly above the optic chiasm. It is responsible for controlling circadian rhythms. The neuronal and hormonal activities it generates regula ...
to effect circadian adjustment and to the
pretectal area
In neuroanatomy, the pretectal area, or pretectum, is a midbrain structure composed of seven nuclei and comprises part of the subcortical visual system. Through reciprocal bilateral projections from the retina, it is involved primarily in mediati ...
to control the
pupillary light reflex
The pupillary light reflex (PLR) or photopupillary reflex is a reflex that controls the diameter of the pupil, in response to the intensity (luminance) of light that falls on the retinal ganglion cells of the retina in the back of the eye, there ...
.
Overview
![Bison bonasus right eye close-up](https://upload.wikimedia.org/wikipedia/commons/c/c4/Bison_bonasus_right_eye_close-up.jpg)
Complex eyes distinguish shapes and
colour
Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associ ...
s. The
visual
The visual system comprises the sensory organ (the eye) and parts of the central nervous system (the retina containing photoreceptor cells, the optic nerve, the optic tract and the visual cortex) which gives organisms the sense of sight (the ...
fields of many organisms, especially predators, involve large areas of
binocular vision
In biology, binocular vision is a type of vision in which an animal has two eyes capable of facing the same direction to perceive a single three-dimensional image of its surroundings. Binocular vision does not typically refer to vision where an ...
for
depth perception
Depth perception is the ability to perceive distance to objects in the world using the visual system and visual perception. It is a major factor in perceiving the world in three dimensions. Depth perception happens primarily due to stereopsis an ...
. In other organisms, particularly prey animals, eyes are located to maximise the field of view, such as in
rabbit
Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s and
horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
s, which have
monocular vision In human species
Monocular vision vision is known as seeing and using only one eye in the human species. Depth perception in monocular vision is reduced compared to binocular vision, but still is active primarily due to accommodation of the eye a ...
.
The first proto-eyes evolved among animals about the time of the
Cambrian explosion
The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
. The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in six of the ~35 main
phyla Phyla, the plural of ''phylum'', may refer to:
* Phylum, a biological taxon between Kingdom and Class
* by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another
Phyl ...
.
[ In most ]vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s and some mollusc
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
s, the eye allows light to enter and project onto a light-sensitive layer of cells
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
known as the retina
The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
. The cone cell
Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...
s (for colour) and the rod cell
Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in per ...
s (for low-light contrasts) in the retina detect and convert light into neural signals which are transmitted to the brain
A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
via the optic nerve
In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual system, visual information from the retina to the brain. In humans, the optic nerve i ...
to produce vision. Such eyes are typically spheroid, filled with the transparent
Transparency, transparence or transparent most often refer to:
* Transparency (optics), the physical property of allowing the transmission of light through a material
They may also refer to:
Literal uses
* Transparency (photography), a still, ...
gel-like vitreous humour
The vitreous body (''vitreous'' meaning "glass-like"; , ) is the clear gel that fills the space between the lens and the retina of the eyeball (the vitreous chamber) in humans and other vertebrates. It is often referred to as the vitreous humor ...
, possess a focusing lens
A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
, and often an iris
Iris most often refers to:
*Iris (anatomy), part of the eye
*Iris (mythology), a Greek goddess
* ''Iris'' (plant), a genus of flowering plants
*Iris (color), an ambiguous color term
Iris or IRIS may also refer to:
Arts and media
Fictional enti ...
. Muscles around the iris change the size of the pupil
The pupil is a black hole located in the center of the iris of the eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company. It appears black ...
, regulating the amount of light that enters the eye and reducing aberrations when there is enough light. The eyes of most cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, fish
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
, amphibian
Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s and snake
Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s have fixed lens shapes, and focusing is achieved by telescoping the lens in a similar manner to that of a camera
A camera is an Optics, optical instrument that can capture an image. Most cameras can capture 2D images, with some more advanced models being able to capture 3D images. At a basic level, most cameras consist of sealed boxes (the camera body), ...
.
The compound eyes of the arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s are composed of many simple facets which, depending on anatomical detail, may give either a single pixelated image or multiple images per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors arranged hexagonally, which can give a full 360° field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera
The Strepsiptera are an order of insects with eleven extant families that include about 600 described species. They are endoparasites in other insects, such as bees, wasps, leafhoppers, silverfish, and cockroaches. Females of most species never ...
, have compound eyes of only a few facets, each with a retina capable of creating an image. With each eye producing a different image, a fused, high-resolution image is produced in the brain.
Possessing detailed hyperspectral
Hyperspectral imaging collects and processes information from across the electromagnetic spectrum. The goal of hyperspectral imaging is to obtain the spectrum for each pixel in the image of a scene, with the purpose of finding objects, identifyi ...
colour vision, the Mantis shrimp
Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in lengt ...
has the world's most complex colour vision system. Trilobite
Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s, now extinct, had unique compound eyes. Clear calcite
Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...
crystals formed the lenses of their eyes. They differ in this from most other arthropods, which have soft eyes. The number of lenses in such an eye varied widely; some trilobites had only one while others had thousands of lenses per eye.
In contrast to compound eyes, simple eyes have a single lens. Jumping spider
Jumping spiders are a group of spiders that constitute the family (biology), family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all sp ...
s have one pair of large simple eyes with a narrow field of view
The field of view (FoV) is the extent of the observable world that is seen at any given moment. In the case of optical instruments or sensors it is a solid angle through which a detector is sensitive to electromagnetic radiation.
Humans a ...
, augmented by an array of smaller eyes for peripheral vision
Peripheral vision, or ''indirect vision'', is vision as it occurs outside the point of fixation, i.e. away from the center of gaze or, when viewed at large angles, in (or out of) the "corner of one's eye". The vast majority of the area in the ...
. Some insect larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
e, like caterpillar
Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths).
As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
s, have a type of simple eye (stemmata
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
) which usually provides only a rough image, but (as in sawfly
Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
larvae) can possess resolving powers of 4 degrees of arc, be polarization-sensitive, and capable of increasing its absolute sensitivity at night by a factor of 1,000 or more. Ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
, some of the simplest eyes, are found in animals such as some of the snail
A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...
s. They have photosensitive Photosensitivity is the amount to which an object reacts upon receiving photons, especially visible light. In medicine, the term is principally used for abnormal reactions of the skin, and two types are distinguished, photoallergy and phototoxicity. ...
cells but no lens or other means of projecting an image onto those cells. They can distinguish between light and dark but no more, enabling them to avoid direct sunlight
Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when t ...
.
In organisms dwelling near deep-sea vents
A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspot ...
, compound eyes are adapted to see the infra-red light
Infrared (IR), sometimes called infrared light, is electromagnetic radiation (EMR) with wavelengths longer than those of visible light. It is therefore invisible to the human eye. IR is generally understood to encompass wavelengths from around ...
produced by the hot vents; enabling the creatures to avoid being boiled alive.[
]
Types
There are ten different eye layouts—indeed every technological method of capturing an optical image commonly used by human beings, with the exceptions of zoom
Zoom may refer to:
Technology Computing
* Zoom (software), videoconferencing application
* Page zooming, the ability to magnify or shrink a portion of a page on a computer display
* Zooming user interface, a graphical interface allowing for image ...
and Fresnel lens
A Fresnel lens ( ; ; or ) is a type of composite compact lens developed by the French physicist Augustin-Jean Fresnel (1788–1827) for use in lighthouses. It has been called "the invention that saved a million ships."
The design allows the c ...
es, occur in nature.[ Eye types can be categorised into "simple eyes", with one concave photoreceptive surface, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface.][ Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution—the physics of ]compound eyes
A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eye
Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conve ...
s, so are better suited to dark-dwelling creatures.[ Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or ]rhabdomeric
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
. These two groups are not monophyletic; the cnidaria
Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter.
Their distinguishing feature is cnidocytes, specialized cells that th ...
also possess cilliated cells,
and some gastropods
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, as well as some annelid
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s possess both.
Some organisms have photosensitive Photosensitivity is the amount to which an object reacts upon receiving photons, especially visible light. In medicine, the term is principally used for abnormal reactions of the skin, and two types are distinguished, photoallergy and phototoxicity. ...
cells that do nothing but detect whether the surroundings are light or dark
Darkness, the direct opposite of lightness, is defined as a lack of illumination, an absence of visible light, or a surface that absorbs light, such as black or brown.
Human vision is unable to distinguish colors in conditions of very low lum ...
, which is sufficient for the entrainment
Entrainment may refer to:
* Air entrainment, the intentional creation of tiny air bubbles in concrete
* Brainwave entrainment, the practice of entraining one's brainwaves to a desired frequency
* Entrainment (biomusicology), the synchronization o ...
of circadian rhythm
A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., Endogeny (biology), endogeno ...
s. These are not considered eyes because they lack enough structure to be considered an organ, and do not produce an image.
Non-compound eyes
Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s, cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, annelid
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s, crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s and cubozoa
Box jellyfish (class Cubozoa) are cnidarian invertebrates distinguished by their box-like (i.e. cube-shaped) body. Some species of box jellyfish produce potent venom delivered by contact with their tentacles. Stings from some species, including '' ...
.
Pit eyes
Pit eyes, also known as stemma
Stemma (plural stemmata) may refer to:
* In stemmatics, an approach to textual criticism, a stemma or stemma codicum is a diagram showing the relationships of the various versions of a text to earlier versions or manuscripts
* Tree-like diagrams ...
, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light.[ Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eyes". They are small, comprising up to about 100 cells covering about 100 µm.][ The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.][
]Pit vipers
The Crotalinae, commonly known as pit vipers,Mehrtens JM (1987). ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . crotaline snakes (from grc, κρόταλον ''krotalon'' castanet), or pit adders, are a subfa ...
have developed pits that function as eyes by sensing thermal infra-red radiation, in addition to their optical wavelength eyes like those of other vertebrates (see infrared sensing in snakes
The ability to sense infrared thermal radiation evolved independently in two different groups of snakes, one consisting of the families Boidae (boas) and Pythonidae (pythons), the other of the family Crotalinae (pit vipers). What is commonly calle ...
). However, pit organs are fitted with receptors rather different from photoreceptors, namely a specific transient receptor potential channel
Transient receptor potential channels (TRP channels) are a group of ion channels located mostly on the plasma membrane of numerous animal cell types. Most of these are grouped into two broad groups: Group 1 includes TRPC ( "C" for canonical), TR ...
(TRP channels) called TRPV1
The transient receptor potential cation channel subfamily V member 1 (TrpV1), also known as the capsaicin receptor and the vanilloid receptor 1, is a protein that, in humans, is encoded by the ''TRPV1'' gene. It was the first isolated member of th ...
. The main difference is that photoreceptors are G-protein coupled receptors
G protein-coupled receptors (GPCRs), also known as seven-(pass)-transmembrane domain receptors, 7TM receptors, heptahelical receptors, serpentine receptors, and G protein-linked receptors (GPLR), form a large group of protein family, evolution ...
but TRP are ion channel
Ion channels are pore-forming membrane proteins that allow ions to pass through the channel pore. Their functions include establishing a resting membrane potential, shaping action potentials and other electrical signals by gating the flow of io ...
s.
Spherical lens eye
The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index
In optics, the refractive index (or refraction index) of an optical medium is a dimensionless number that gives the indication of the light bending ability of that medium.
The refractive index determines how much the path of light is bent, or ...
to form a lens, which may greatly reduce the blur radius encountered—hence increasing the resolution obtainable.[ The most basic form, seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina.][ This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing ]spherical aberration
In optics, spherical aberration (SA) is a type of optical aberration, aberration found in optical systems that have elements with spherical surfaces. Lens (optics), Lenses and curved mirrors are prime examples, because this shape is easier to man ...
.[ Such a non-homogeneous lens is necessary for the focal length to drop from about 4 times the lens radius, to 2.5 radii.][
Heterogeneous eyes have evolved at least nine times: four or more times in ]gastropods
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, once in the copepod
Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s, once in the annelid
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s, once in the cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s,[ and once in the ]chiton
Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized.
They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
s, which have aragonite
Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including prec ...
lenses. No extant aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".[
This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimise the effect of eye motion while the animal moves, most such eyes have stabilising eye muscles.][
The ]ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
of insects bear a simple lens, but their focal point usually lies behind the retina; consequently, those can't form a sharp image. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain.[ Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity.]
This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).[
]
Multiple lenses
Some marine organisms bear more than one lens; for instance the copepod
Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
''Pontella
''Pontella'' is a marine copepod genus in the family Pontellidae. It is an organism that bears three lenses in the eye. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed ...
'' has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, ''Copilia
''Copilia'' is a genus of copepods in the family Sapphirinidae. The eyes in members of this genus have two lenses, arranged like those in a telescope.
Species
The genus ''Copilia'' contains the following species:
* '' Copilia asiaticus'' ( ...
'', has two lenses in each eye, arranged like those in a telescope.[ Such arrangements are rare and poorly understood, but represent an alternative construction.
Multiple lenses are seen in some hunters such as eagles and jumping spiders, which have a refractive cornea: these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.][
]
Refractive cornea
In the eyes of most mammals, birds
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air.[ In general, the lens is not spherical. Spherical lenses produce spherical aberration. In refractive corneas, the lens tissue is corrected with inhomogeneous lens material (see ]Luneburg lens
A Luneburg lens (original German ''Lüneburg lens'', sometimes incorrectly spelled ''Luneberg lens'') is a spherically symmetric gradient-index lens. A typical Luneburg lens's refractive index ''n'' decreases radially from the center to the outer ...
), or with an aspheric shape.[ Flattening the lens has a disadvantage; the quality of vision is diminished away from the main line of focus. Thus, animals that have evolved with a wide field-of-view often have eyes that make use of an inhomogeneous lens.][
As mentioned above, a refractive cornea is only useful out of water. In water, there is little difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures that have returned to the water—penguins and seals, for example—lose their highly curved cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strongly focusing cornea.][
]
Reflector eyes
An alternative to a lens is to line the inside of the eye with "mirrors", and reflect the image to focus at a central point.[ The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.][
Many small organisms such as ]rotifer
The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals.
They were first described by Rev. John H ...
s, copepods and flatworm
The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...
s use such organs, but these are too small to produce usable images.[ Some larger organisms, such as ]scallop
Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families ...
s, also use reflector eyes. The scallop '' Pecten'' has up to 100 millimetre-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.[
There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of ]guanine
Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine (uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is called ...
crystal
A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macros ...
s.
Compound eyes
A compound eye may consist of thousands of individual photoreceptor units or ommatidia (ommatidium
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
, singular). The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarisation of light. Because the individual lenses are so small, the effects of diffraction
Diffraction is defined as the interference or bending of waves around the corners of an obstacle or through an aperture into the region of geometrical shadow of the obstacle/aperture. The diffracting object or aperture effectively becomes a s ...
impose a limit on the possible resolution that can be obtained (assuming that they do not function as phased array
In antenna theory, a phased array usually means an electronically scanned array, a computer-controlled array of antennas which creates a beam of radio waves that can be electronically steered to point in different directions without moving th ...
s). This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require very large compound eyes, around in radius.
Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, annelids and some bivalved molluscs. Compound eyes in arthropods grow at their margins by the addition of new ommatidia.[
]
Apposition eyes
Apposition eyes are the most common form of eyes and are presumably the ancestral form of compound eyes. They are found in all arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
groups, although they may have evolved more than once within this phylum.[ Some ]annelids
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
and bivalves
Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
also have apposition eyes. They are also possessed by ''Limulus
''Limulus'' is a genus of horseshoe crab, with one extant species, the Atlantic horseshoe crab (''Limulus polyphemus''). One fossil species is currently assigned to the genus though several other species have been named, which have since been as ...
'', the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point.[ (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)
Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information. The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ]ommatidium
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
.
Superposition eyes
The second type is named the superposition eye. The superposition eye is divided into three types:
* refracting,
* reflecting and
* parabolic superposition
The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This type of compound eye, for which a minimal size exists below which effective superposition cannot occur, is normally found in nocturnal insects, because it can create images up to 1000 times brighter than equivalent apposition eyes, though at the cost of reduced resolution. In the parabolic superposition compound eye type, seen in arthropods such as mayflies
Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the orde ...
, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp
Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are refer ...
, prawn
Prawn is a common name for small aquatic crustaceans with an exoskeleton and ten legs (which is a member of the order decapoda), some of which can be eaten.
The term "prawn"Mortenson, Philip B (2010''This is not a weasel: a close look at nature ...
s, crayfish
Crayfish are freshwater crustaceans belonging to the clade Astacidea, which also contains lobsters. In some locations, they are also known as crawfish, craydids, crawdaddies, crawdads, freshwater lobsters, mountain lobsters, rock lobsters, mu ...
and lobster
Lobsters are a family (biology), family (Nephropidae, Synonym (taxonomy), synonym Homaridae) of marine crustaceans. They have long bodies with muscular tails and live in crevices or burrows on the sea floor. Three of their five pairs of legs ...
s are alone in having reflecting superposition eyes, which also have a transparent gap but use corner mirror
A mirror or looking glass is an object that Reflection (physics), reflects an image. Light that bounces off a mirror will show an image of whatever is in front of it, when focused through the lens of the eye or a camera. Mirrors reverse the ...
s instead of lenses.
Parabolic superposition
This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.[
]
Other
Another kind of compound eye, found in males of Order Strepsiptera
The Strepsiptera are an order of insects with eleven extant families that include about 600 described species. They are endoparasites in other insects, such as bees, wasps, leafhoppers, silverfish, and cockroaches. Females of most species never ...
, employs a series of simple eyes—eyes having one opening that provides light for an entire image-forming retina. Several of these ''eyelets'' together form the strepsipteran compound eye, which is similar to the 'schizochroal' compound eyes of some trilobites
Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
. Because each eyelet is a simple eye, it produces an inverted image; those images are combined in the brain to form one unified image. Because the aperture of an eyelet is larger than the facets of a compound eye, this arrangement allows vision under low light levels.[
Good fliers such as flies or honey bees, or prey-catching insects such as ]praying mantis
Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
or dragonflies
A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, have specialised zones of ommatidia
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
organised into a fovea
Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure.
Human anatomy
*Fovea centralis of the retina
* Fovea buccalis or Dimple
* Fovea of the femoral head
* Trochlear fovea of the fr ...
area which gives acute vision. In the acute zone, the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution. The black spot that can be seen on the compound eyes of such insects, which always seems to look directly at the observer, is called a pseudopupil
In the compound eye of invertebrates such as insects and crustaceans, the pseudopupil appears as a dark spot which moves across the eye as the animal is rotated. This occurs because the ommatidia that one observes "head-on" (along their optica ...
. This occurs because the ommatidia
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
which one observes "head-on" (along their optical axes) absorb the incident light
In optics a ray is an idealized geometrical model of light, obtained by choosing a curve that is perpendicular to the ''wavefronts'' of the actual light, and that points in the direction of energy flow. Rays are used to model the propagation o ...
, while those to one side reflect it.
There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid
Mysida is an order of small, shrimp-like crustaceans in the malacostracan superorder Peracarida. Their common name opossum shrimps stems from the presence of a brood pouch or "marsupium" in females. The fact that the larvae are reared in thi ...
shrimp, ''Dioptromysis paucispinosa''. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialised retina. The resulting eye is a mixture of a simple eye within a compound eye.
Another version is a compound eye often referred to as "pseudofaceted", as seen in ''Scutigera
''Scutigera'' is a centipede genus in the scutigeromorph (house centipede) family Scutigeridae, a group of centipedes with long limbs and true compound eyes (which were once thought to be secondary, re-evolved "pseudofacetted eyes"). It compose ...
''. This type of eye consists of a cluster of numerous ommatidia
The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ou ...
on each side of the head, organised in a way that resembles a true compound eye.
The body of ''Ophiocoma wendtii
''Ophiomastix wendtii'', also known by its common name, the red ophiocoma, and formerly as ''Ophiocoma wendtii'', is a species of brittle stars that inhabits coral reefs from Bermuda to Brazil, primarily in the Caribbean sea. club-like spines al ...
'', a type of brittle star
Brittle stars, serpent stars, or ophiuroids (; ; referring to the serpent-like arms of the brittle star) are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomo ...
, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chiton
Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized.
They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
s. The tube feet of sea urchins contain photoreceptor proteins, which together act as a compound eye; they lack screening pigments, but can detect the directionality of light by the shadow cast by its opaque body.
Nutrients
The ciliary body is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. The inner layer is transparent and covers the vitreous body, and is continuous from the neural tissue of the retina. The outer layer is highly pigmented, continuous with the retinal pigment epithelium, and constitutes the cells of the dilator muscle.
The vitreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the eye and the retina lining the back of the eye. It is produced by certain retinal cells. It is of rather similar composition to the cornea, but contains very few cells (mostly phagocytes which remove unwanted cellular debris in the visual field, as well as the hyalocytes of Balazs of the surface of the vitreous, which reprocess the hyaluronic acid), no blood vessels, and 98–99% of its volume is water (as opposed to 75% in the cornea) with salts, sugars, vitrosin (a type of collagen), a network of collagen type II fibres with the mucopolysaccharide hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye.
Evolution
Photoreception is phylogenetically
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
very old, with various theories of phylogenesis. The common origin (monophyly
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
) of all animal eyes is now widely accepted as fact. This is based upon the shared genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 650-600 million years ago, and the PAX6
Paired box protein Pax-6, also known as aniridia type II protein (AN2) or oculorhombin, is a protein that in humans is encoded by the ''PAX6'' gene.
Function
PAX6 is a member of the Pax gene family which is responsible for carrying the geneti ...
gene is considered a key factor in this. The majority of the advancements in early eyes are believed to have taken only a few million years to develop, since the first predator to gain true imaging would have touched off an "arms race" among all species that did not flee the photopic environment. Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel (except those of groups, such as the vertebrates, that were only forced into the photopic environment at a late stage).
Eyes in various animals show adaptation to their requirements. For example, the eye of a bird of prey
Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predators h ...
has much greater visual acuity than a human eye
The human eye is a sensory organ, part of the sensory nervous system, that reacts to visible light and allows humans to use visual information for various purposes including seeing things, keeping balance, and maintaining circadian rhythm.
...
, and in some cases can detect ultraviolet
Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
radiation. The different forms of eye in, for example, vertebrates and molluscs are examples of parallel evolution
Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
, despite their distant common ancestry. Phenotypic convergence of the geometry of cephalopod and most vertebrate eyes creates the impression that the vertebrate eye evolved from an imaging cephalopod eye
Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions.Budelmann BU. "Cephalopod sense organs, nerves and the brain: Adaptations for high performance and life style." Marine and Freshwater Behavi ...
, but this is not the case, as the reversed roles of their respective ciliary and rhabdomeric opsin classes and different lens crystallins show.
The very earliest "eyes", called eye-spots, were simple patches of photoreceptor protein
Photoreceptor proteins are light-sensitive proteins involved in the sensing and response to light in a variety of organisms. Some examples are rhodopsin in the photoreceptor cells of the vertebrate retina, phytochrome in plants, and bacteriorhodo ...
in unicellular animals. In multicellular beings, multicellular eyespots evolved, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the light source.[
Through gradual change, the eye-spots of species living in well-lit environments depressed into a shallow "cup" shape. The ability to slightly discriminate directional brightness was achieved by using the angle at which the light hit certain cells to identify the source. The pit deepened over time, the opening diminished in size, and the number of photoreceptor cells increased, forming an effective ]pinhole camera
A pinhole camera is a simple camera without a lens but with a tiny aperture (the so-called ''pinhole'')—effectively a light-proof box with a small hole in one side. Light from a scene passes through the aperture and projects an inverted image o ...
that was capable of dimly distinguishing shapes. However, the ancestors of modern hagfish
Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
, thought to be the protovertebrate, were evidently pushed to very deep, dark waters, where they were less vulnerable to sighted predators, and where it is advantageous to have a convex eye-spot, which gathers more light than a flat or concave one. This would have led to a somewhat different evolutionary trajectory for the vertebrate eye than for other animal eyes.
The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialise into a transparent humour that optimised colour filtering, blocked harmful radiation, improved the eye's refractive index
In optics, the refractive index (or refraction index) of an optical medium is a dimensionless number that gives the indication of the light bending ability of that medium.
The refractive index determines how much the path of light is bent, or ...
, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent crystallin
In anatomy, a crystallin is a water-soluble structural protein found in the lens and the cornea of the eye accounting for the transparency of the structure. It has also been identified in other places such as the heart, and in aggressive breast c ...
protein.[Fernald, Russell D. (2001)]
The Evolution of Eyes: Where Do Lenses Come From?
''Karger Gazette'' 64: "The Eye in Focus".
The gap between tissue layers naturally formed a biconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical power ...
and iris
Iris most often refers to:
*Iris (anatomy), part of the eye
*Iris (mythology), a Greek goddess
* ''Iris'' (plant), a genus of flowering plants
*Iris (color), an ambiguous color term
Iris or IRIS may also refer to:
Arts and media
Fictional enti ...
. Separation of the forward layer again formed a humour, the aqueous humour
The aqueous humour is a transparent water-like fluid similar to plasma, but containing low protein concentrations. It is secreted from the ciliary body, a structure supporting the lens of the eyeball. It fills both the anterior and the posterio ...
. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes.
Relationship to life requirements
Eyes are generally adapted to the environment and life requirements of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
n plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre.
Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition.[ Optical superposition eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia.][ Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example, if the animal is on a slope.]
An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey. In deep water organisms, it may not be the centre of the eye that is enlarged. The hyperiid
The Hyperiidea are a suborder of amphipods, small aquatic crustaceans. Unlike the other suborders of Amphipoda, hyperiids are exclusively marine and do not occur in fresh water. Hyperiids are distinguished by their large eyes and planktonic habi ...
amphipod
Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from and are mostly detritivores or scavengers. There are more than 9,900 amphipod species so far descr ...
s are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey—or predators—against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether.[ In the giant Antarctic isopod Glyptonotus a small ventral compound eye is physically completely separated from the much larger dorsal compound eye.] Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy.[
Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop.][ On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be captured.][
It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function.][
Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head.]
Physiology
Visual acuity
Visual acuity
Visual acuity (VA) commonly refers to the clarity of vision, but technically rates an examinee's ability to recognize small details with precision. Visual acuity is dependent on optical and neural factors, i.e. (1) the sharpness of the retinal ...
, or resolving power, is "the ability to distinguish fine detail" and is the property of cone cells
Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cones ...
. It is often measured in ''cycles per degree
Degree may refer to:
As a unit of measurement
* Degree (angle), a unit of angle measurement
** Degree of geographical latitude
** Degree of geographical longitude
* Degree symbol (°), a notation used in science, engineering, and mathematics
...
'' (CPD), which measures an angular resolution
Angular resolution describes the ability of any image-forming device such as an optical or radio telescope, a microscope, a camera, or an eye, to distinguish small details of an object, thereby making it a major determinant of image resolution. ...
, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white/black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white/black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a grey block, is then the measurement of visual acuity of the eye.
For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD (1.2 arcminute
A minute of arc, arcminute (arcmin), arc minute, or minute arc, denoted by the symbol , is a unit of angular measurement equal to of one degree. Since one degree is of a turn (or complete rotation), one minute of arc is of a turn. The na ...
per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD. A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central fovea
Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure.
Human anatomy
*Fovea centralis of the retina
* Fovea buccalis or Dimple
* Fovea of the femoral head
* Trochlear fovea of the fr ...
region.
Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair. A resolution of 2 arcminutes per line pair, equivalent to a 1 arcminute gap in an optotype
__NOTOC__
An eye chart, or optotype, is a chart used to subjectively measure visual acuity. Eye charts are often used by health care professionals, such as optometrists, physicians or nurses, to screen persons for vision impairment. Ophthalmologis ...
, corresponds to 20/20 (normal vision
Visual acuity (VA) commonly refers to the clarity of visual perception, vision, but technically rates an examinee's ability to recognize small details with precision. Visual acuity is dependent on optical and neural factors, i.e. (1) the sharpn ...
) in humans.
However, in the compound eye, the resolution is related to the size of individual ommatidia and the distance between neighbouring ommatidia. Physically these cannot be reduced in size to achieve the acuity seen with single lensed eyes as in mammals. Compound eyes have a much lower acuity than vertebrate eyes.
Colour perception
"Colour vision is the faculty of the organism to distinguish lights of different spectral qualities." All organisms are restricted to a small range of electromagnetic spectrum; this varies from creature to creature, but is mainly between wavelengths of 400 and 700 nm.
This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range.
The most sensitive pigment, rhodopsin
Rhodopsin, also known as visual purple, is a protein encoded by the RHO gene and a G-protein-coupled receptor (GPCR). It is the opsin of the rod cells in the retina and a light-sensitive receptor protein that triggers visual phototransduction ...
, has a peak response at 500 nm.[ Small changes to the genes coding for this protein can tweak the peak response by a few nm;][ pigments in the lens can also filter incoming light, changing the peak response.][ Many organisms are unable to discriminate between colours, seeing instead in shades of grey; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of ]cone cell
Cone cells, or cones, are photoreceptor cells in the retinas of vertebrate eyes including the human eye. They respond differently to light of different wavelengths, and the combination of their responses is responsible for color vision. Cone ...
s, from which the more sensitive rod cell
Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in per ...
s evolved. Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioural tests can this be deduced.[
Most organisms with colour vision can detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light—this precludes the possibility of any UV light being detected, as it does not even reach the retina.][
]
Rods and cones
The retina contains two major types of light-sensitive photoreceptor cell
A photoreceptor cell is a specialized type of neuroepithelial cell found in the retina that is capable of visual phototransduction. The great biological importance of photoreceptors is that they convert light (visible electromagnetic radiatio ...
s used for vision: the rods and the cones
A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex.
A cone is formed by a set of line segments, half-lines, or lines conn ...
.
Rods cannot distinguish colours, but are responsible for low-light (scotopic
In the study of human visual perception, scotopic vision (or scotopia) is the vision of the eye under low-light conditions. The term comes from Greek ''skotos'', meaning "darkness", and ''-opia'', meaning "a condition of sight". In the human eye, ...
) monochrome (black-and-white
Black-and-white (B&W or B/W) images combine black and white in a continuous spectrum, producing a range of shades of grey.
Media
The history of various visual media began with black and white, and as technology improved, altered to color. ...
) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher (photopic
Photopic vision is the vision of the eye under well-lit conditions (luminance levels from 10 to 108 cd/m2). In humans and many other animals, photopic vision allows color perception, mediated by cone cells, and a significantly higher visua ...
) intensities. Rods are distributed throughout the retina but there are none at the fovea
Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure.
Human anatomy
*Fovea centralis of the retina
* Fovea buccalis or Dimple
* Fovea of the femoral head
* Trochlear fovea of the fr ...
and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina.
Cones are responsible for colour vision
Color vision, a feature of visual perception, is an ability to perceive differences between light composed of different wavelengths (i.e., different spectral power distributions) independently of light intensity. Color perception is a part of t ...
. They require brighter light to function than rods require. In humans, there are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colours). The colour seen is the combined effect of stimuli
A stimulus is something that causes a physiological response. It may refer to:
*Stimulation
**Stimulus (physiology), something external that influences an activity
**Stimulus (psychology), a concept in behaviorism and perception
*Stimulus (economi ...
to, and responses from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on the fovea, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibres of the optic nerve
In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual system, visual information from the retina to the brain. In humans, the optic nerve i ...
. When rods and cones are stimulated by light, they connect through adjoining cells within the retina to send an electrical signal to the optic nerve fibres. The optic nerves send off impulses through these fibres to the brain.[
]
Pigmentation
The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related—although problems of convergence do exist.[
]Opsins
Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most p ...
are the pigments involved in photoreception. Other pigments, such as melanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
, are used to shield the photoreceptor cells from light leaking in from the sides.
The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since.[
There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells.] The eyes of vertebrates usually contain ciliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric cells in the eye with r-opsins. However, some ''ganglion'' cells of vertebrates express r-opsins, suggesting that their ancestors
An ancestor, also known as a forefather, fore-elder or a forebear, is a parent or (recursively) the parent of an antecedent (i.e., a grandparent, great-grandparent, great-great-grandparent and so forth). ''Ancestor'' is "any person from whom ...
used this pigment in vision, and that remnants survive in the eyes.[ Likewise, c-opsins have been found to be expressed in the ''brain'' of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently resorbed into the brain on metamorphosis to the adult form.][ C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use r-opsins.][ ]Cnidaria
Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter.
Their distinguishing feature is cnidocytes, specialized cells that th ...
, which are an outgroup to the taxa mentioned above, express c-opsins—but r-opsins are yet to be found in this group.[ Incidentally, the melanin produced in the cnidaria is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment.][
]
Additional images
File:Three Main Layers of the Eye.png, The structures of the eye labelled
File:Three Internal chambers of the Eye.svg, Another view of the eye and the structures of the eye labelled
See also
* Adaptation (eye)
In visual physiology, adaptation is the ability of the retina of the eye to adjust to various levels of light. Natural night vision, or scotopic vision, is the ability to see under low-light conditions. In humans, rod cells are exclusively resp ...
(night vision)
* Emission theory (vision)
Emission theory or extramission theory (variants: extromission) or extromissionism is the proposal that visual perception is accomplished by eye beams emitted by the eyes. This theory has been replaced by ''intromission theory'' (or '' intromissi ...
* Eye color
Eye color is a polygenic phenotypic character determined by two distinct factors: the pigmentation of the eye's iris and the frequency-dependence of the scattering of light by the turbid medium in the stroma of the iris.
In humans, the pi ...
* Eye development
Eye formation in the human embryo begins at approximately three weeks into embryonic development and continues through the tenth week.Ort, D., David, H."Development of the Eye" Retrieved 22 April 2015. Cells from both the mesodermal and the ecto ...
* Eye disease
This is a partial list of human eye diseases and disorders.
The World Health Organization publishes a classification of known diseases and injuries, the International Statistical Classification of Diseases and Related Health Problems, or ICD-10. ...
* Eye injury
Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conve ...
* Eye movement
Eye movement includes the voluntary or involuntary movement of the eyes. Eye movements are used by a number of organisms (e.g. primates, rodents, flies, birds, fish, cats, crabs, octopus) to fixate, inspect and track visual objects of interests. ...
* Eyelid
An eyelid is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. The human eyel ...
* Nictitating membrane
The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. All ...
* Ophthalmology
Ophthalmology ( ) is a surgical subspecialty within medicine that deals with the diagnosis and treatment of eye disorders.
An ophthalmologist is a physician who undergoes subspecialty training in medical and surgical eye care. Following a medic ...
* Orbit (anatomy)
In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
* Simple eye in invertebrates
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
* ''Tapetum lucidum
The ''tapetum lucidum'' ( ; ; ) is a layer of tissue in the eye of many vertebrates and some other animals. Lying immediately behind the retina, it is a retroreflector. It reflects visible light back through the retina, increasing the light a ...
''
* Tears
Tears are a clear liquid secreted by the lacrimal glands (tear gland) found in the eyes of all land mammals. Tears are made up of water, electrolytes, proteins, lipids, and mucins that form layers on the surface of eyes. The different types of ...
Notes
References
Citations
Bibliography
*
Further reading
*
External links
Evolution of the eye
Anatomy of the eye – flash animated interactive.
(Adobe Flash
Adobe Flash (formerly Macromedia Flash and FutureSplash) is a multimedia Computing platform, software platform used for production of Flash animation, animations, rich web applications, application software, desktop applications, mobile apps, mo ...
)
Webvision. The organisation of the retina and visual system.
An in-depth treatment of retinal function, open to all but geared most towards graduate students.
{{Authority control
Sensory organs
Visual system