Compound eye

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Compound eye of a house centipede 20140421 Scutigera coleoptrata left eye.jpg
Compound eye of a house centipede
Compound eye of a dragonfly DragonFly macrogiants E.jpg
Compound eye of a dragonfly

A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, [1] which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distinguish brightness and color. The image perceived by this arthropod eye is a combination of inputs from the numerous ommatidia, which are oriented to point in slightly different directions. Compared with single-aperture eyes, compound eyes have poor image resolution; however, they possess a very large view angle and the ability to detect fast movement and, in some cases, the polarization of light. [2] Because a compound eye is made up of a collection of ommatidia, each with its own lens, light will enter each ommatidium instead of using a single entrance point. The individual light receptors behind each lens are then turned on and off due to a series of changes in the light intensity during movement or when an object is moving, creating a flicker-effect known as the flicker frequency, which is the rate at which the ommatidia are turned on and off– this facilitates faster reaction to movement; honey bees respond in 0.01s compared with 0.05s for humans. [3]

Contents

Types

Head of a mantisfly showing a compound eye Mantis Fly - Genus Plega.jpg
Head of a mantisfly showing a compound eye
Drawing from Robert Hooke's Micrographia of the compound eye of a grey drone fly Compoundeye.png
Drawing from Robert Hooke's Micrographia of the compound eye of a grey drone fly

Compound eyes are typically classified as either apposition eyes, which form multiple inverted images, or superposition eyes, which form a single erect image. [4]

Apposition eyes

Apposition eyes can be divided into two groups. 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 mantis shrimp is the most advanced example of an animal with this type of eye. In the other kind of apposition eye, found in the Strepsiptera, each lens forms an image, and the images are combined in the brain. [5] This is called the schizochroal compound eye or the neural superposition eye (which, despite its name, is a form of the apposition eye).

Superposition eyes

The superposition eye is divided into three subtypes; the refracting, the reflecting, and the parabolic superposition eye. 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 kind is used mostly by nocturnal insects. In the parabolic superposition eye, seen in arthropods such as mayflies, 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, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also have a transparent gap but use corner mirrors instead of lenses.

Other

Compound eyes of a paper wasp. Paper Wasp Head.jpg
Compound eyes of a paper wasp.

Good fliers like flies or honey bees, or prey-catching insects like praying mantises or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eye is flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution.

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 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 specialized retina. The resulting eye is a mixture of a simple eye within a compound eye.

Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organized in a way that resembles a true compound eye.

Asymmetries in compound eyes may be associated with asymmetries in behaviour. For example, Temnothorax albipennis ant scouts show behavioural lateralization when exploring unknown nest sites, showing a population-level bias to prefer left turns. One possible reason for this is that its environment is partly maze-like and consistently turning in one direction is a good way to search and exit mazes without getting lost. [6] This turning bias is correlated with slight asymmetries in the ants' compound eyes (differential ommatidia count). [7]

Fossilized compound eyes from the Florissant Formation Fossilized compound eyes.jpg
Fossilized compound eyes from the Florissant Formation

The body of Ophiomastix wendtii , a type of brittle star, was previously thought to be covered with ommatidia, turning its whole skin into a compound eye, but this has since been found to be erroneous; the system does not rely on lenses or image formation. [8]

See also

Cultural references

"Dragonfly eyes" (Chinese: 蜻蜓眼 qingting yan] is a term for knobbly multi-coloured glass beads made in Western and Eastern Asia 2000–2500 years ago. [9] Owing to the multiple views and stimuli, compound eyes or dragonfly eyes have become a feature in art, film and literature, particularly in the 2010s. For example:

Related Research Articles

<span class="mw-page-title-main">Optics</span> Branch of physics that studies light

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, ultraviolet, and infrared light. Light is a type of electromagnetic radiation, and other forms of electromagnetic radiation such as X-rays, microwaves, and radio waves exhibit similar properties.

<span class="mw-page-title-main">Strepsiptera</span> Order of insects

The Strepsiptera are an order of insects with eleven extant families that include about 600 described species. They are endoparasites of other insects, such as bees, wasps, leafhoppers, silverfish, and cockroaches. Females of most species never emerge from the host after entering its body, finally dying inside it. The early-stage larvae do emerge because they must find an unoccupied living host, and the short-lived males must emerge to seek a receptive female in her host. They are believed to be most closely related to beetles, from which they diverged 300–350 million years ago, but do not appear in the fossil record until the mid-Cretaceous around 100 million years ago.

A common classification of the Lepidoptera involves their differentiation into butterflies and moths. Butterflies are a natural monophyletic group, often given the suborder Rhopalocera, which includes Papilionoidea, Hesperiidae (skippers), and Hedylidae. In this taxonomic scheme, moths belong to the suborder Heterocera. Other taxonomic schemes have been proposed, the most common putting the butterflies into the suborder Ditrysia and then the "superfamily" Papilionoidea and ignoring a classification for moths.

<span class="mw-page-title-main">Eye</span> Organ that detects light and converts it into electro-chemical impulses in neurons

An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system.

<span class="mw-page-title-main">Prism (optics)</span> Transparent optical element with flat, polished surfaces that refract light

An optical prism is a transparent optical element with flat, polished surfaces that are designed to refract light. At least one surface must be angled — elements with two parallel surfaces are not prisms. The most familiar type of optical prism is the triangular prism, which has a triangular base and rectangular sides. Not all optical prisms are geometric prisms, and not all geometric prisms would count as an optical prism. Prisms can be made from any material that is transparent to the wavelengths for which they are designed. Typical materials include glass, acrylic and fluorite.

<span class="mw-page-title-main">Mantis shrimp</span> Order of crustaceans

Mantis shrimp 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 10 cm (3.9 in) in length, while a few can reach up to 38 cm (15 in). A mantis shrimp's carapace covers only the rear part of the head and the first four segments of the thorax. Varieties range in colour from shades of brown to vivid colours, with more than 520 species of mantis shrimp known. They are among the most important predators in many shallow, tropical and subtropical marine habitats. However, despite being common, they are poorly understood, as many species spend most of their lives sheltering in burrows and holes.

<span class="mw-page-title-main">Ommatidium</span> Component of compound eyes of arthropods

The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia. An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The outer part of the ommatidium is overlaid with a transparent cornea. Each ommatidium is innervated by one axon bundle and provides the brain with one picture element. The brain forms an image from these independent picture elements. The number of ommatidia in the eye depends upon the type of arthropod and range from as low as 5 as in the Antarctic isopod Glyptonotus antarcticus, or a handful in the primitive Zygentoma, to around 30,000 in larger Anisoptera dragonflies and some Sphingidae moths.

<span class="mw-page-title-main">Refractive error</span> Problem with focusing light accurately on the retina due to the shape of the eye

Refractive error is a problem with focusing light accurately on the retina due to the shape of the eye and/or cornea. The most common types of refractive error are near-sightedness, far-sightedness, astigmatism, and presbyopia. Near-sightedness results in far away objects being blurry, far-sightedness and presbyopia result in close objects being blurry, and astigmatism causes objects to appear stretched out or blurry. Other symptoms may include double vision, headaches, and eye strain.

<span class="mw-page-title-main">Eye examination</span> Series of tests assessing vision and pertaining to the eyes

An eye examination is a series of tests performed to assess vision and ability to focus on and discern objects. It also includes other tests and examinations pertaining to the eyes. Eye examinations are primarily performed by an optometrist, ophthalmologist, or an orthoptist. Health care professionals often recommend that all people should have periodic and thorough eye examinations as part of routine primary care, especially since many eye diseases are asymptomatic.

<span class="mw-page-title-main">Simple eye in invertebrates</span> Simple eye without retina

A simple eye 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-lensed "compound eye", and is not necessarily at all simple in the usual sense of the word.

<span class="mw-page-title-main">Evolution of the eye</span> Origins and diversification of the organs of sight through geologic time

Many scientists have found the evolution of the eye attractive to study because the eye distinctively exemplifies an analogous organ found in many animal forms. Simple light detection is found in bacteria, single-celled organisms, plants and animals. Complex, image-forming eyes have evolved independently several times.

<span class="mw-page-title-main">Pseudopupil</span> Dark spot on the compound eyes of some invertebrates

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" absorb the incident light, while those to one side reflect it. The pseudopupil therefore reveals which ommatidia are aligned with the axis along which the observer is viewing.

<span class="mw-page-title-main">Holochroal eye</span>

Holochroal eyes are compound eyes with many tiny lenses. They are the oldest and most common type of trilobite eye, and found in all orders of trilobite from the Cambrian to the Permian periods. Lenses covered a curved, kidney-shaped visual surface in a hexagonal close packing system, with a single corneal membrane covering all lenses. Unlike in schizochroal eyes, adjacent lenses were in direct contact with one another. Lens shape generally depended on cuticle thickness. The lenses of trilobites with thin cuticles were thin and biconvex, whereas those with thick cuticles had thick lenses, which in extreme cases, could be thick columns with the outer surface flattened and the inner surface hemispherical. Regardless of lens thickness, however, the point at which light was focused was roughly the same distance below the lens.

<span class="mw-page-title-main">Arthropod eye</span> Visual organs possessed by arthropods

Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum. Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, 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.

A compound refractive lens (CRL) is a series of individual lenses arranged in a linear array in order to achieve focusing of X-rays in the energy range of 5–40 keV. They are an alternative to the KB mirror.

<i>Temnothorax albipennis</i> Species of ant

Temnothorax albipennis, the rock ant is a species of small ant in the subfamily Myrmicinae. It occurs in Europe and builds simple nests in rock crevices.

<i>Megalopta</i> Genus of bees

Megalopta is a widespread neotropical genus of bees in the tribe Augochlorini in family Halictidae, known as the sweat bees. They are the largest of the five nocturnal genera in Augochlorini. Most have pale integumentary pigmentation, and all have large ocelli, most likely a feature of their nocturnal behavior. They live in tropical Central America and the entirety of South America. The subgenus Noctoraptor is cleptoparasitic. They are not known from the fossil record.

<i>Gonodactylus chiragra</i> Species of crustacean

Gonodactylus chiragra is a medium to large mantis shrimp that is distributed widely throughout the West Indo-Pacific.

<span class="mw-page-title-main">Spider vision</span> Eyes of spiders

The eyes of spiders vary significantly in their structure, arrangement, and function. They usually have eight, each being a simple eye with a single lens rather than multiple units as in the compound eyes of insects. The specific arrangement and structure of the eyes is one of the features used in the identification and classification of different species, genera, and families. Most haplogynes have six eyes, although some have eight (Plectreuridae), four or even two. In some cave species, there are no eyes at all. Sometimes one pair of eyes is better developed than the rest. Several families of hunting spiders, such as jumping spiders and wolf spiders, have fair to excellent vision. The main pair of eyes in jumping spiders even sees in colour.

References

  1. "Senses. Insect eyes" . Insects and Spiders of the World. Volume 8: Scorpion fly - Stinkbug. New York: Marshall Cavendish. 2003. p.  459. ISBN   978-0761473428.
  2. Völkel, R.; Eisner, M.; Weible, K.J. (June 2003). "Miniaturized imaging systems" (PDF). Microelectronic Engineering. 67–68 (8): 461–472. doi:10.1016/S0167-9317(03)00102-3. Archived from the original (PDF) on 2008-10-01.
  3. Biologically Inspired Computer Vision: Fundamentals and Applications
  4. Gaten, Edward (1998). "Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea)". Contributions to Zoology. 67 (4): 223–236. doi: 10.1163/18759866-06704001 .
  5. Buschbeck, Elke K. (1 July 2005). "The compound lens eye of Strepsiptera: morphological development of larvae and pupae". Arthropod Structure & Development. 34 (3): 315–326. doi:10.1016/j.asd.2005.04.002. ISSN   1467-8039 . Retrieved 3 July 2022.
  6. Hunt ER, et al. (2014). "Ants show a leftward turning bias when exploring unknown nest sites". Biology Letters . 10 (12): 20140945. doi:10.1098/rsbl.2014.0945. PMC   4298197 . PMID   25540159.
  7. Hunt ER, et al. (11 April 2018). "Asymmetric ommatidia count and behavioural lateralization in the ant Temnothorax albipennis". Scientific Reports . 8 (1): 5825. Bibcode:2018NatSR...8.5825H. doi:10.1038/s41598-018-23652-4. PMC   5895843 . PMID   29643429.
  8. Sumner-Rooney L, Rahman IA, Sigwart JD, Ullrich-Lüter E (January 2018). "Whole-body photoreceptor networks are independent of 'lenses' in brittle stars". Proceedings. Biological Sciences. 285 (1871): 20172590. doi:10.1098/rspb.2017.2590. PMC   5805950 . PMID   29367398.
  9. https://www.cambridge.org/core/journals/microscopy-and-microanalysis/article/abs/nondestructive-analysis-of-dragonfly-eye-beads-from-the-warring-states-period-excavated-from-a-chu-tomb-at-the-shenmingpu-site-henan-province-china/E2FCF854D5324115F503E1643C33BDBD DOI: https://doi.org/10.1017/S1431927612014201