A simple eye or ocellus (sometimes called a pigment pit [1] [2] ) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish them from "compound eyes", which have muliple lenses. They are not necessarily simple in the sense of being uncomplicated or basic.
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. Arthropods 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 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 most large animals are camera eyes and are sometimes considered "simple" because a single lens collects and focuses an entire image onto the retina (analogous to a camera). By other criteria, the presence of a complex retina distinguishes the vertebrate camera eye from the simple stemma or ommatidia which make up compound eyes. Additionally, not all invertebrate ocelli and ommatidium have simple photoreceptors. Many have various forms of retinula (a retina-like cluster of photoreceptor cells), including the ommatidia of most insects and the central eyes of camel spiders. Jumping spiders and some other predatory spiders with seemingly simple eyes also emulate retinal vision in various ways. Many insects 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.
Some jellyfish, sea stars, flatworms, and ribbonworms [3] have the simplest "eyes" – pigment spot ocelli – which have randomly distributed pigment, and which have no other structure (such as a cornea, or lens). The apparent "eye color" in these animals is red or black. [4] Certain groups such as box jellyfish have more complex eyes, including some with a distinct retina, lens, and cornea. [5]
Many snails and slugs also have ocelli, either at the tips or bases of their tentacles. [6] Some other gastropods, such as the Strombidae, have much more sophisticated eyes. Giant clams have ocelli that allow light to penetrate their mantles. [7]
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 some telescopic ability) to help spot prey from a distance. Nocturnal spiders' eyes are very sensitive in low light levels and are large to capture more light, equivalent to f/0.58 in the rufous net-casting spider. [8]
The term "ocellus" (plural ocelli) is derived from the Latin oculus (eye), and literally means "little eye". In insects, two distinct ocellus types exist: [9] dorsal (top-most) ocelli, and lateral ocelli (often referred to as ocelli and stemmata, respectively), most insects have dorsal ocelli while stemmata are found in the larvae of some insect orders. Despite the shared name, they are structurally and functionally very different. Simple eyes of other animals may also be referred to as ocelli, but again the structure and anatomy of these eyes is quite distinct from those of insect dorsal ocelli.
Dorsal ocelli are light-sensitive organs found on the dorsal surface or frontal surface of the head of many insects, including Hymenoptera (bees, ants, wasps, sawflies), Diptera (flies), Odonata (dragonflies, damselflies), Orthoptera (grasshoppers, locusts) and Mantodea (mantises). These ocelli coexist with 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, [10] wasps, dragonflies and locusts) where they are typically found as a triplet. Two ocelli are directed to either side of the head, while a central (median) ocellus is directed forwards. In some terrestrial insects (e.g. some ants and cockroaches), the median ocellus is absent. The sideways-facing ocelli can be called "lateral ocelli", referring to their direction and position in the triplet, however this is not to be confused with the stemmata of some insect larvae, which are also known as lateral ocelli.
A dorsal ocellus consists of a lens element (cornea) and a layer of photoreceptors (rod cells). The ocellar lens may be strongly curved or flat. The photoreceptor layer may also be separated from the lens by a clear vitreous humour. The number of photoreceptors also varies widely, but may number in the hundreds or thousands for well-developed ocelli. In bees, locusts, and dragonflies, the lens is strongly curved; while in cockroaches it is flat. Locusts possess vitreous humour while blowflies and dragonflies do not.
Two somewhat unusual features of ocelli are particularly notable and generally common between insect orders.
These two factors have led to the conclusion that, with some exceptions in predatry insects, the ocelli are incapable of perceiving proper images 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 (amplification of photoreceptor signals), 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. [11]
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. Locusts [12] and dragonflies [13] in tethered flight have been observed to try and "correct" their flight posture based on changes in light. 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" similar to camera eyes, as the ocellar lens forms an image within, or close to, the photoreceptor layer. [14] [15] In dragonflies it has been demonstrated that the receptive fields of both the photoreceptors [16] and the second-order neurons [17] can be quite restricted. Further research has demonstrated these eyes not only resolve spatial details of the world, but also perceive motion. [18] 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 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. 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. [19]
Stemmata (singular stemma) are a class of simple eyes. Many kinds of holometabolous 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 fleas, springtails, and Thysanura. Some other Arthropoda, such as some Myriapoda, rarely have any eyes other than stemmata at any stage of their lives (exceptions include the large and well-developed compound eyes of the house centipedes, Scutigera [20] ).
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 and especially those of Tenthredinidae, 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. [21] They may be optimized for light sensitivity, as opposed to detailed image formation. [22] In the pupal stage, such stemmata develop into fully fledged compound eyes. One feature offering a clue to their ontogenetic 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. [9]
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. [23] In Drosophila, the rhodopsin Rh2 is only expressed in simple eyes. [24]
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. [25]
Epidermal growth factor receptor (Egfr) promotes the expression of orthodenticle and possibly eyes absent (Eya) and as such is essential for simple eye formation. [25]
The retina 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 processes that image within the retina and sends nerve impulses along the optic nerve to the visual cortex to create visual perception. The retina serves a function which is in many ways analogous to that of the film or image sensor in a camera.
Arachnids are arthropods in the class Arachnida of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.
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.
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 spiders have some of the best vision among arthropods and use it in courtship, hunting, and navigation. Although they normally move unobtrusively and fairly slowly, most species are capable of very agile jumps, notably when hunting, but sometimes in response to sudden threats or crossing long gaps. Both their book lungs and tracheal system are well-developed, and they use both systems. Jumping spiders are generally recognized by their eye pattern. All jumping spiders have four pairs of eyes, with the anterior median pair being particularly large.
The visual system is the physiological basis of visual perception. The system detects, transduces and interprets information concerning light within the visible range to construct an image and build a mental model of the surrounding environment. The visual system is associated with the eye and functionally divided into the optical system and the neural system.
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 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. 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.
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.
Rhopalia from Ancient Greek ῥόπαλον (rhópalon) 'club' are small sensory structures of certain Scyphozoan and Cubozoan species.
The supraesophageal ganglion is the first part of the arthropod, especially insect, central nervous system. It receives and processes information from the first, second, and third metameres. The supraesophageal ganglion lies dorsal to the esophagus and consists of three parts, each a pair of ganglia that may be more or less pronounced, reduced, or fused depending on the genus:
A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhythmicity and hormone production for thermoregulation. The hole in the head which contains the eye is known as a pineal foramen or parietal foramen, since it is often enclosed by the parietal bones.
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.
Misumena vatia is a species of crab spider with a holarctic distribution. In North America, it is called the goldenrod crab spider or flower (crab) spider, as it is commonly found hunting in goldenrod sprays and milkweed plants. They are called crab spiders because of their unique ability to walk sideways as well as forwards and backwards. Both males and females of this species progress through several molts before reaching their adult sizes, though females must molt more to reach their larger size. Females can grow up to 10 mm (0.39 in) while males are quite small, reaching 5 mm (0.20 in) at most. Misumena vatia are usually yellow or white or a pattern of these two colors. They may also present with pale green or pink instead of yellow, again, in a pattern with white. They have the ability to change between these colors based on their surroundings through the molting process. They have a complex visual system, with eight eyes, that they rely on for prey capture and for their color-changing abilities. Sometimes, if Misumena vatia consumes colored prey, the spider itself will take on that color.
Photoreceptor can refer to:
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.
Orthodenticle (otd) is a homeobox gene found in Drosophila that regulates the development of anterior patterning, with particular involvement in the central nervous system function and eye development. It is located on the X chromosome. The gene is an ortholog of the human OTX1/OTX2 gene.
Thermonectus marmoratus is a relatively colorful North American species of diving beetle known by the common names sunburst diving beetle and yellow-spotted diving beetle. The behavior of this diving beetle has been compared to a scuba diver, since it carries with it a bubble of air as it dives down into the water. Its aquatic larval stage was the first ever recorded use of bifocals in the animal world. The beetle uses in its principal eyes two retinas and two distinct focal planes that are substantially separated, in the manner of bifocals to switch their vision from up-close to distance, for easy and efficient capture of their prey.
Insects are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body, three pairs of jointed legs, compound eyes, and a pair of antennae. Insects are the most diverse group of animals, with more than a million described species; they represent more than half of all animal species.
Vision is an important sensory system for most species of fish. Fish eyes are similar to the eyes of terrestrial vertebrates like birds and mammals, but have a more spherical lens. Birds and mammals normally adjust focus by changing the shape of their lens, but fish normally adjust focus by moving the lens closer to or further from the retina. Fish retinas generally have both rod cells and cone cells, and most species have colour vision. Some fish can see ultraviolet and some are sensitive to polarised light.
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.
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.
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