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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 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 ...
such as occurs 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. In this sense "simple eye" is distinct from a multi-lensed "
compound eye 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 disti ...
", and is not necessarily at all simple in the usual sense of the word. The structure of an animal's eye is determined by the environment in which it lives, and the behavioural tasks it must fulfill to survive.
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 differ widely in the habitats in which they live, as well as their visual requirements for finding food or conspecifics, and avoiding predators. Consequently, an enormous variety of eye types are found in arthropods. They possess a wide variety of novel solutions to overcome visual problems or limitations. Use of the term ''simple eye'' is flexible, and must be interpreted in proper context; for example, the eyes of humans and of other large animals such as most cephalopods, are ''
camera 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'' and in some usages are classed as "simple" because a single lens collects and focuses light onto 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 ...
(analogous to
film A film also called a movie, motion picture, moving picture, picture, photoplay or (slang) flick is a work of visual art that simulates experiences and otherwise communicates ideas, stories, perceptions, feelings, beauty, or atmosphere ...
). By other criteria, the presence of a complex retina distinguishes the vertebrate camera eye from the simple stemma or ommatidium. Additionally not all ocelli (''
ocellus 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 ...
'' in singular) and ommatidia of invertebrates have simple photoreceptors; many, including the ommatidia of most insects and the central eyes of
Solifugae Solifugae is an order of animals in the class Arachnida known variously as camel spiders, wind scorpions, sun spiders, or solifuges. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are ...
have various forms of '' retinula'', and the
Salticidae Jumping spiders are a group of spiders that constitute the 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 species. Jumping spi ...
and some other predatory
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s with seemingly simple eyes, emulate retinal vision in various ways. Many
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s have unambiguously compound eyes consisting of multiple lenses (up to tens of thousands), but achieve an effect similar to that of a camera eye, in that each ommatidium lens focuses light onto a number of neighbouring retinulae.


Ocelli or eye spots

Some jellyfish, sea stars, flatworms, and ribbonworms have the simplest "eyes" – ''pigment spot ocelli'' – which have randomly distributed pigment, and which have no other structure (such as a
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 ...
, or
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''), ...
). The apparent "eye color" in these animals is red or black. However,
box jellyfish 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 '' ...
have more complex eyes, including those of which have distinct retina, lens, and cornea. Many snails and slugs (gastropod mollusks) also have ocelli, either at the tips, or at the bases, of their tentacles. However, some other gastropods, such as the
Strombidae Strombidae, commonly known as the true conchs, is a taxonomic family of medium-sized to very large sea snails in the superfamily Stromboidea, and the Epifamily Neostromboidae. The term true conchs, being a common name, does not have an exact me ...
, have much more sophisticated eyes. Giant clams (
Tridacna ''Tridacna'' is a genus of large saltwater clams, marine bivalve molluscs in the subfamily Tridacninae, the giant clams. They have heavy shells, fluted with 4 to 6 folds. The mantle is brightly coloured. They inhabit shallow waters of coral ...
) have ocelli that allow light to penetrate their mantles.


Simple eyes in arthropods


Spider eyes

Spiders do not have compound eyes, but instead have several pairs of simple eyes with each pair adapted for a specific task or tasks. The principal and secondary eyes in spiders are arranged in four, or occasionally fewer, pairs. Only the principal eyes have moveable retinas. The secondary eyes have a reflector at the back of the eyes. The light-sensitive part of the receptor cells is next to this, so they get direct and reflected light. In hunting or jumping spiders, for example, a forward-facing pair possesses the best resolution (and even telescopic components) to see the (often small) prey at a large distance. Night-hunting spiders' eyes are very sensitive in low light levels with a large aperture, f/0.58.


Dorsal ocelli

The term "ocellus" (plural ocelli) is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
' (eye), and literally means "little eye". In insects, two distinct ocellus types exist:
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
ocelli (or simply "ocelli"), found in most insects, and lateral ocelli (or
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 are found in the larvae of some insect orders. They are structurally and functionally very different. Simple eyes of other animals, e.g.
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 ...
ns, may also be referred to as ocelli, but again the structure and anatomy of these eyes is quite distinct from those of the dorsal ocelli of insects. Dorsal ocelli are light-sensitive organs found on the dorsal (top-most) surface or frontal surface of the head of many insects, e.g.
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
( bees,
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s,
sawflies 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 ...
), Diptera (flies),
Odonata Odonata is an order of flying insects that includes the dragonflies and damselflies. Members of the group first appeared during the Triassic, though members of their total group, Odonatoptera, first appeared in Late Carboniferous. The two com ...
(
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 ...
, damselflies), Orthoptera ( grasshoppers,
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
s) and
Mantodea 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 ...
(mantises). The ocelli coexist with the compound eyes; thus, most insects possess two anatomically separate and functionally different visual pathways. The number, forms, and functions of the dorsal ocelli vary markedly throughout insect orders. They tend to be larger and more strongly expressed in flying insects (particularly bees, wasps, dragonflies and locusts), where they are typically found as a triplet. Two lateral ocelli are directed to the left and right of the head, respectively, while a central (median) ocellus is directed frontally. In some terrestrial insects (e.g. some ants and cockroaches), only two lateral ocelli are present: the median ocellus is absent. The unfortunately labelled "lateral ocelli" here refers to the sideways-facing position of the ocelli, which are of the dorsal type. They should not be confused with the lateral ocelli of some insect larvae (see
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 ...
). A dorsal ocellus consists of a lens element (
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 a layer of photoreceptors (
rod cells 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 p ...
). The ocellar lens may be strongly curved (e.g. bees, locusts, dragonflies) or flat (e.g. cockroaches). The photoreceptor layer may (e.g. locusts) or may not (e.g. blowflies, dragonflies) be separated from the lens by a clear zone (
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 ...
). The number of photoreceptors also varies widely, but may number in the hundreds or thousands for well-developed ocelli. Two somewhat unusual features of the ocelli are particularly notable and generally well conserved between insect orders. #The refractive power of the lens is not typically sufficient to form an image on the photoreceptor layer. #Dorsal ocelli ubiquitously have massive convergence ratios from first-order (photoreceptor) to second-order neurons. These two factors have led to the conclusion that the dorsal ocelli are incapable of perceiving form, and are thus solely suitable for light-metering functions. Given the large aperture and low ''f''-number of the lens, as well as high convergence ratios and synaptic gains, the ocelli are generally considered to be far more sensitive to light than the compound eyes. Additionally, given the relatively simple neural arrangement of the eye (small number of synapses between detector and effector), as well as the extremely large diameter of some ocellar interneurons (often the largest diameter neurons in the animal's nervous system), the ocelli are typically considered to be "faster" than the compound eyes. One common theory of ocellar function in flying insects holds that they are used to assist in maintaining flight stability. Given their underfocused nature, wide fields of view, and high light-collecting ability, the ocelli are superbly adapted for measuring changes in the perceived brightness of the external world as an insect rolls or pitches around its body axis during flight. Corrective flight responses to light have been demonstrated in locusts and dragonflies in tethered flight. Other theories of ocellar function have ranged from roles as light adaptors or global excitatory organs to polarization sensors and circadian entrainers. Recent studies have shown the ocelli of some insects (most notably the dragonfly, but also some wasps) are capable of form vision, as the ocellar lens forms an image within, or close to, the photoreceptor layer. In dragonflies it has been demonstrated that the receptive fields of both the photoreceptors and the second-order neurons can be quite restricted. Further research has demonstrated these eyes not only resolve spatial details of the world, but also perceive motion. Second-order neurons in the dragonfly median ocellus respond more strongly to upwards-moving bars and gratings than to downwards-moving bars and gratings, but this effect is only present when
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 ...
light is used in the stimulus; when ultraviolet light is absent, no directional response is observed. Dragonfly ocelli are especially highly developed and specialised visual organs, which may support the exceptional acrobatic abilities of these animals. Research on the ocelli is of high interest to designers of small
unmanned aerial vehicles An unmanned aerial vehicle (UAV), commonly known as a drone, is an aircraft without any human pilot, crew, or passengers on board. UAVs are a component of an unmanned aircraft system (UAS), which includes adding a ground-based controlle ...
. Designers of these craft face many of the same challenges that insects face in maintaining stability in a three-dimensional world. Engineers are increasingly taking inspiration from insects to overcome these challenges.


Stemmata

Stemmata (singular stemma) are a class of simple eyes. Many kinds of
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
larvae bear no other form of eyes until they enter their final stage of growth. Adults of several orders of hexapods also have stemmata, and never develop compound eyes at all. Examples include
flea Flea, the common name for the order Siphonaptera, includes 2,500 species of small flightless insects that live as external parasites of mammals and birds. Fleas live by ingesting the blood of their hosts. Adult fleas grow to about long, a ...
s,
springtail Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
s, and
Thysanura Thysanura is the now deprecated name of what was, for over a century, recognised as an order in the class Insecta. The two constituent groups within the former order, the Archaeognatha (jumping bristletails) and the Zygentoma (silverfish and fireb ...
. Some other
Arthropoda 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 ...
, such as some
Myriapoda Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
, rarely have any eyes other than stemmata at any stage of their lives (exceptions include the large and well-developed compound eyes of
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 compo ...
). Behind each lens of a typical, functional stemma, lies a single cluster of photoreceptor cells, termed a retinula. The lens is biconvex, and the body of the stemma has a vitreous or crystalline core. Although stemmata are simple eyes, some kinds, such as those of the larvae of
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
and especially those of
Tenthredinidae Tenthredinidae is the largest family of sawflies, with well over 7,500 species worldwide, divided into 430 genera. Larvae are herbivores and typically feed on the foliage of trees and shrubs, with occasional exceptions that are leaf miners, stem ...
, a family of sawflies, are only "simple" in that they represent immature or embryonic forms of the compound eyes of the adult. They can possess a considerable degree of acuity and sensitivity, and can detect polarized light. In the
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
l stage, such stemmata develop into fully fledged compound eyes. One feature offering a clue to their
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the st ...
role is their lateral position on the head; ocelli, that in other ways resemble stemmata, tend to be borne in sites median to the compound eyes, or nearly so. Among some researchers, this distinction has led to the use of the term "lateral ocelli" for stemmata.


Genetic controls

A number of genetic pathways are responsible for the occurrence and positioning of the ocelli. The gene orthodenticle is allelic to ocelliless, a mutation that stops ocelli from being produced. In ''Drosophila'', the rhodopsin Rh2 is only expressed in simple eyes. While (in ''Drosophila'' at least) the genes eyeless and dachshund are both expressed in the compound eye but not the simple eye, no reported 'developmental' genes are uniquely expressed in the simple eye. Epidermal growth factor receptor (''Egfr'') promotes the expression of orthodenticle nd possibly eyes absent (''Eya'') and as such is essential for simple eye formation.


See also

* Arthropod eye
* Evolution of the eye * Eyespot apparatus * Mollusc eye * Parietal eye * Ocelloid


References


Further reading

*


External links

* {{DEFAULTSORT:Simple Eyes In Arthropods Eye Invertebrates