The compound eyes of arthropods like insects, crustaceans and millipedes [1] are composed of units called ommatidia (sg.: ommatidium). 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 (usually consisting of 6–9 axons, depending on the number of rhabdomeres) [2] : 162 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 , [3] or a handful in the primitive Zygentoma, to around 30,000 in larger Anisoptera dragonflies and some Sphingidae moths. [4]
Ommatidia are typically hexagonal in cross-section and approximately ten times longer than wide. The diameter is largest at the surface, tapering toward the inner end. At the outer surface, there is a cornea, below which is a pseudocone that further focuses the light. The cornea and pseudocone form the outer ten percent of the length of the ommatidium.
The specific organ of ommatidia, or eye units, vary between different organisms. The butterfly compound eye consists of multiple ommatidia, each of which consist of nine photoreceptor cells (numbered from R1–R9), and primary and secondary pigment cells. [5] Nymphalid butterflies have the simplest eye ommatidium structure, consisting of eight photoreceptor cells (R1–R8) and a tiny R9 cell organized into a different tier. [5] These "R cells" tightly pack the ommatidium. The portion of the R cells at the central axis of the ommatidium collectively form a light guide, a transparent tube, called the rhabdom.
Although composed of over 16,000 cells, [6] the Drosophila compound eye is a simple repetitive pattern of 700 to 750 ommatidia, [7] initiated in the larval eye imaginal disc. Each ommatidium consists of 14 neighboring cells: 8 photoreceptor neurons in the core, 4 non-neuronal cone cells and 2 primary pigment cells. [6] A hexagonal lattice of pigment cells insulates the ommatidial core from neighboring ommatidia to optimize coverage of the visual field, which therefore affects the acuity of Drosophila vision. [7]
In true flies, the rhabdom has separated into seven independent rhabdomeres (there are actually eight, but the two central rhabdomeres responsible for color vision sit one atop the other), such that a small inverted 7-pixel image is formed in each ommatidium. The angle between adjacent rhabdomeres within a single ommatidium (the acceptance angle) is similar to the angle between adjacent ommatidia (the inter-ommatidial angle), giving the eye a continuous field of view with areas of overlap between neighboring ommatidia; [2] : 161–2 the advantage of this arrangement is that the same visual axis is sampled from a larger area of the eye, increasing overall sensitivity by a factor of seven, without increasing the size of the eye or reducing its acuity. Achieving this has also required the rewiring of the eye such that the axon bundles are twisted through 180 degrees (re-inverted), and each rhabdomere is united with those from the six adjacent ommatidia that share the same visual axis. Thus, at the level of the lamina – the first optical processing center of the insect brain – the signals are input in exactly the same manner as in the case of a normal apposition compound eye, but the image is enhanced. This visual arrangement is known as neural superposition. [2] : 163–4
Since an image from the compound eye is created from the independent picture elements produced by ommatidia, it is important for the ommatidia to react only to that part of the scene directly in front of them. To prevent light entering at an angle from being detected by the ommatidium it entered, or by any of the neighboring ommatidia, six pigment cells are present. The pigment cells line the outside of each ommatidium. Each pigment cell is situated at the apex of the hexagons and thus lines the outside of three ommatidia. Light entering at an angle passes through the thin cross-section of the photoreceptor cell, with only a tiny chance of exciting it, and is absorbed by the pigment cell, before it can enter a neighboring ommatidium. In many species, in low-light situations, the pigment is withdrawn, so that light entering the eye might be detected by any of several ommatidia. This enhances light detection but lowers resolution.
The size of the ommatidia varies according to species, but ranges from 5 to 50 micrometres. The rhabdoms within them may cross-section at least as small as 1.x micrometres, the category of "small" being assigned in some cross-species studies to those under 2 micrometers. [8] A microlens array can be seen as an elementary, biomimetic analogy of ommatidia.
Retinal cell fate determination relies on positional cell–cell signaling that activates signal transduction pathways, rather than cell lineage. Cell–cell signal that is released from R8 photoreceptors (already differentiated retinal cells) of each ommatidium is received by neighboring progenitor retinal cells, stimulating their incorporation into developing ommatidia. The undifferentiated retinal cells select their appropriate cell fates based on their position with their differentiated neighbors. The local signal, Growth Factor Spitz, activates the epidermal growth factor receptor (EGFR) signal transduction pathway, and initiates a cascade of events that will result in transcription of genes involved in cell fate determination. [9] This process leads to the induction of cell fates, starting from the R8 photoreceptor neurons and progresses to the sequential recruitment of neighboring undifferentiated cells. The first seven neighboring cells receive R8 signaling to differentiate as photoreceptor neurons, followed by the recruitment of the four non-neuronal cone cells.
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.
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.
Drosophila melanogaster is a species of fly in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly", "pomace fly", or "banana fly". In the wild, D. melanogaster are attracted to rotting fruit and fermenting beverages, and are often found in orchards, kitchens and pubs.
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.
Retinitis pigmentosa (RP) is a member of a group of genetic disorders called inherited retinal dystrophy (IRD) that cause loss of vision. Symptoms include trouble seeing at night and decreasing peripheral vision. As peripheral vision worsens, people may experience "tunnel vision". Complete blindness is uncommon. Onset of symptoms is generally gradual and often begins in childhood.
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.
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 into signals that can stimulate biological processes. To be more specific, photoreceptor proteins in the cell absorb photons, triggering a change in the cell's membrane potential.
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 peripheral vision. On average, there are approximately 92 million rod cells in the human retina. Rod cells are more sensitive than cone cells and are almost entirely responsible for night vision. However, rods have little role in color vision, which is the main reason why colors are much less apparent in dim light.
The fovea centralis is a small, central pit composed of closely packed cones in the eye. It is located in the center of the macula lutea of the retina.
A retinal ganglion cell (RGC) is a type of neuron located near the inner surface of the retina of the eye. It receives visual information from photoreceptors via two intermediate neuron types: bipolar cells and retina amacrine cells. Retina amacrine cells, particularly narrow field cells, are important for creating functional subunits within the ganglion cell layer and making it so that ganglion cells can observe a small dot moving a small distance. Retinal ganglion cells collectively transmit image-forming and non-image forming visual information from the retina in the form of action potential to several regions in the thalamus, hypothalamus, and mesencephalon, or midbrain.
Electroretinography measures the electrical responses of various cell types in the retina, including the photoreceptors, inner retinal cells, and the ganglion cells. Electrodes are placed on the surface of the cornea or on the skin beneath the eye to measure retinal responses. Retinal pigment epithelium (RPE) responses are measured with an EOG test with skin-contact electrodes placed near the canthi. During a recording, the patient's eyes are exposed to standardized stimuli and the resulting signal is displayed showing the time course of the signal's amplitude (voltage). Signals are very small, and typically are measured in microvolts or nanovolts. The ERG is composed of electrical potentials contributed by different cell types within the retina, and the stimulus conditions can elicit stronger response from certain components.
A simple eye or ocellus 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 multiple lenses. They are not necessarily simple in the sense of being uncomplicated or basic.
Visual phototransduction is the sensory transduction process of the visual system by which light is detected by photoreceptor cells in the vertebrate retina. A photon is absorbed by a retinal chromophore, which initiates a signal cascade through several intermediate cells, then through the retinal ganglion cells (RGCs) comprising the optic nerve.
In neurobiology, lateral inhibition is the capacity of an excited neuron to reduce the activity of its neighbors. Lateral inhibition disables the spreading of action potentials from excited neurons to neighboring neurons in the lateral direction. This creates a contrast in stimulation that allows increased sensory perception. It is also referred to as lateral antagonism and occurs primarily in visual processes, but also in tactile, auditory, and even olfactory processing. Cells that utilize lateral inhibition appear primarily in the cerebral cortex and thalamus and make up lateral inhibitory networks (LINs). Artificial lateral inhibition has been incorporated into artificial sensory systems, such as vision chips, hearing systems, and optical mice. An often under-appreciated point is that although lateral inhibition is visualised in a spatial sense, it is also thought to exist in what is known as "lateral inhibition across abstract dimensions." This refers to lateral inhibition between neurons that are not adjacent in a spatial sense, but in terms of modality of stimulus. This phenomenon is thought to aid in colour discrimination.
The pigmented layer of retina or retinal pigment epithelium (RPE) is the pigmented cell layer just outside the neurosensory retina that nourishes retinal visual cells, and is firmly attached to the underlying choroid and overlying retinal visual cells.
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.
Vision is the most important sense for birds, since good eyesight is essential for safe flight. Birds have a number of adaptations which give visual acuity superior to that of other vertebrate groups; a pigeon has been described as "two eyes with wings". Birds are theropods, and the avian eye resembles that of other sauropsids, with ciliary muscles that can change the shape of the lens rapidly and to a greater extent than in the mammals. Birds have the largest eyes relative to their size in the animal kingdom, and movement is consequently limited within the eye's bony socket. In addition to the two eyelids usually found in vertebrates, bird's eyes are protected by a third transparent movable membrane. The eye's internal anatomy is similar to that of other vertebrates, but has a structure, the pecten oculi, unique to birds.
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.
Sevenless (sev) is a gene in the fruit fly Drosophila melanogaster that encodes a receptor tyrosine kinase protein essential to the development of the R7 photoreceptor cells in the Drosophila embryonic eye. The Drosophila ommatidium contains 8 distinct retinula or R cells, each of which has a different spectral sensitivity. The R7 photo receptor, located in each of several ommatidia in the fly's compound eye, is used to detect ultraviolet light. The R8 photoreceptor contains an activator of the RTK for on a precursor R7 cell, called the bride of sevenless (BOSS). The binding of BOSS to sevenless stimulates a complex series of reactions involving the RTK (sevenless), MAP kinases, Ras and many more molecules to differentiate that precursor R7 photo receptor to a fully functional R7 photo receptor that can see UV light. Much of this knowledge was gained by examining flies with a mutant sevenless which still produced a fully functional R7 photoreceptor when a dominant Ras was injected into the mutant R7 precursor.
Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions. They have a camera-type eye which consists of an iris, a circular lens, vitreous cavity, pigment cells, and photoreceptor cells that translate light from the light-sensitive retina into nerve signals which travel along the optic nerve to the brain. For the past 140 years, the camera-type cephalopod eye has been compared with the vertebrate eye as an example of convergent evolution, where both types of organisms have independently evolved the camera-eye trait and both share similar functionality. Contention exists on whether this is truly convergent evolution or parallel evolution. Unlike the vertebrate camera eye, the cephalopods' form as invaginations of the body surface, and consequently the cornea lies over the top of the eye as opposed to being a structural part of the eye. Unlike the vertebrate eye, a cephalopod eye is focused through movement, much like the lens of a camera or telescope, rather than changing shape as the lens in the human eye does. The eye is approximately spherical, as is the lens, which is fully internal.