ratio of word probabilities predicted from brain for cat and eye

cat	eye
top 10 words in brain distribution (in article): species animal male breed female bird human hunt cat live	top 10 words in brain distribution (in article): water light human form time type produce term surface world
top 10 words in brain distribution (not in article): wolf egg elephant drink genus lamp deer ant hybrid bee	top 10 words in brain distribution (not in article): drink lamp wine ice beer state rock people key process
times more probable under cat 30 20 10 6 4 2.5 1.25 1 1.25 2.5 4 6 10 20 30 times more probable under eye (words not in the model)
The cat'" ("Felis catus"), also known as the domestic cat'" or "'house cat'" to distinguish it from other felines and felids, is a small predatory carnivorous species of crepuscular mammal that is valued by humans for its companionship and its ability to hunt vermin, snakes, scorpions, and other unwanted household pests. It has been associated with humans for at least 9,500 years. A skilled predator, the cat is known to hunt over 1,000 species for food. It can be trained to obey simple commands. Individual cats have also been known to learn on their own to manipulate simple mechanisms, such as doorknobs. Cats use a variety of vocalizations and types of body language for communication, including meowing, purring, hissing, growling, squeaking, chirping, clicking, and grunting. Cats may be the most popular pet in the world, with over 600 million in homes all over the world. They are also bred and shown as registered pedigree pets. This hobby is known as the "cat fancy." Until recently the cat was commonly believed to have been domesticated in ancient Egypt, where it was a cult animal. However a 2007 study found that the lines of descent of all house cats probably run through as few as five self-domesticating African Wildcats "(Felis silvestris lybica)" circa 8000 BC, in the Near East. Size. Cats typically weigh between 2.5 and 7 kg (5.5–16 pounds); however, some, such as the Maine Coon, can exceed. Some have been known to reach up to due to overfeeding. Conversely, very small cats (less than) have been reported. The largest cat ever was officially reported to have weighed in at about (46 lb 15.25 oz). The smallest cat ever officially recorded weighed around 3 lbs (1.36 kg). Cats average about in height and in head body length (males being larger than females), with tails averaging in length. Skeleton. Cats have 7 cervical vertebrae like almost all mammals, 13 thoracic vertebrae (humans have 12), 7 lumbar vertebrae (humans have 5), 3 sacral vertebrae like most mammals (humans have 5 because of their bipedal posture), and, except for Manx cats, 22 or 23 caudal vertebrae (humans have 3 to 5, fused into an internal coccyx). The extra lumbar and thoracic vertebrae account for the cat's enhanced spinal mobility and flexibility, compared with humans. The caudal vertebrae form the tail, used by the cat as a counterbalance to the body during quick movements. Cats also have free-floating clavicle bones, which allows them to pass their body through any space into which they can fit their heads. Mouth. Cats have highly specialized teeth for the killing of prey and the tearing of meat. The premolar and first molar together compose the carnassial pair on each side of the mouth, which efficiently functions to shear meat like a pair of scissors. While this is present in canids, it is highly developed in felines. The cat's tongue has sharp spines, or papillae, useful for retaining and ripping flesh from a carcass. These papillae are small backward-facing hooks that contain keratin which also assist in their grooming. As facilitated by their oral structure, cats use a variety of vocalizations and types of body language for communication, including meowing, purring, hissing, growling, squeaking, chirping, clicking, and grunting. Ears. Thirty-two individual muscles in each ear allow for a manner of directional hearing: a cat can move each ear independently of the other. Because of this mobility, a cat can move its body in one direction and point its ears in another direction. Most cats have straight ears pointing upward. Unlike dogs, flap-eared breeds are extremely rare (Scottish Folds are one such exceptional mutation.) When angry or frightened, a cat will lay back its ears, to accompany the growling or hissing sounds it makes. Cats also turn their ears back when they are playing, or to listen to a sound coming from behind them. The angle of cats' ears is an important clue to their mood. Legs. Cats, like dogs, are digitigrades. They walk directly on their toes, with the bones of their feet making up the lower part of the visible leg. Cats are capable of walking very precisely, because like all felines they directly register; that is, they place each hind paw (almost) directly in the print of the corresponding forepaw, minimizing noise and visible tracks. This also provides sure footing for their hind paws when they navigate rough terrain. Claws. Like nearly all members of family Felidae, cats have protractable claws. In their normal, relaxed position the claws are sheathed with the skin and fur around the toe pads. This keeps the claws sharp by preventing wear from contact with the ground and allows the silent stalking of prey. The claws on the forefeet are typically sharper than those on the hind feet. Cats can voluntarily extend their claws on one or more paws. They may extend their claws in hunting or self-defense, climbing, "kneading", or for extra traction on soft surfaces (bedspreads, thick rugs, etc.). It is also possible to make a cooperative cat extend its claws by carefully pressing both the top and bottom of the paw. The curved claws may become entangled in carpet or thick fabric, which may cause injury if the cat is unable to free itself. Most cats have five claws on their front paws, and four or five on their rear paws. Because of an ancient mutation, however, domestic and feral cats are prone to polydactylyism, (particularly in the east coast of Canada and north east coast of the United States) and may have six or seven toes. The fifth front claw (the dewclaw) is proximal to the other claws. More proximally, there is a protrusion which appears to be a sixth "finger". This special feature of the front paws, on the inside of the wrists, is the carpal pad, also found on the paws of big cats and dogs. It has no function in normal walking, but is thought to be an anti-skidding device used while jumping. Skin. Cats possess rather loose skin; this allows them to turn and confront a predator or another cat in a fight, even when it has a grip on them. This is also an advantage for veterinary purposes, as it simplifies injections. In fact, the lives of cats with kidney failure can sometimes be extended for years by the regular injection of large volumes of fluid subcutaneously, which serves as an alternative to dialysis. The particularly loose skin at the back of the neck is known as the "scruff", and is the area by which a mother cat grips her kittens to carry them. As a result, cats tend to become quiet and passive when gripped there. This behavior also extends into adulthood, when a male will grab the female by the scruff to immobilize her while he mounts, and to prevent her from running away as the mating process takes place. This technique can be useful when attempting to treat or move an uncooperative cat. However, since an adult cat is heavier than a kitten, a pet cat should never be carried by the scruff, but should instead have its weight supported at the rump and hind legs, and at the chest and front paws. Often (much like a small child) a cat will lie with its head and front paws over a person's shoulder, and its back legs and rump supported under the person's arm. Senses. Cat senses are attuned for hunting. Cats have highly advanced hearing, eyesight, taste, and touch receptors, making the cat extremely sensitive among mammals. Cats' night vision is superior to humans although their vision in daylight is inferior. Cat eyes have a tapetum lucidum and cat eyes that are blue typically lack melanin and hence can show the red-eye effect (see odd-eyed cat). Humans and cats have a similar range of hearing on the low end of the scale, but cats can hear much higher-pitched sounds, up to 64 kHz, which is 1.6 octaves above the range of a human, and even one octave above the range of a dog. A domestic cat's sense of smell is about fourteen times as strong as a human's. Due to a mutation in an early cat ancestor, one of two genes necessary to taste sweetness may have been lost by the cat family. To aid with navigation and sensation, cats have dozens of movable vibrissae (whiskers) over their body, especially their face. Metabolism. Cats conserve energy by sleeping more than most animals, especially as they grow older. The daily duration of sleep varies, usually 12–16 hours, with 13–14 being the average. Some cats can sleep as much as 20 hours in a 24-hour period. The term "cat nap" refers to the cat's ability to fall asleep (lightly) for a brief period and has entered the English lexicon – someone who nods off for a few minutes is said to be "taking a cat nap". Due to their crepuscular nature, cats are often known to enter a period of increased activity and playfulness during the evening and early morning, dubbed the "evening crazies", "night crazies", "elevenses", or "mad half-hour" by some. The temperament of a cat can vary depending on the breed and socialization. Cats with oriental body types tend to be thinner and more active, while cats that have a cobby body type tend to be heavier and less active. The normal body temperature of a cat is between 38 and 39 °C (101 and 102.2 °F). A cat is considered febrile (hyperthermic) if it has a temperature of 39.5 °C (103 °F) or greater, or hypothermic if less than 37.5 °C (100 °F). For comparison, humans have a normal temperature of approximately 36.8 °C (98.6 °F). A domestic cat's normal heart rate ranges from 140 to 220 beats per minute, and is largely dependent on how excited the cat is. For a cat at rest, the average heart rate usually is between 150 and 180 bpm, about twice that of a human (average 80 bpm). Genetics. A 2007 study published in the journal "Science" asserts that all house cats are descended from a group of self-domesticating desert wildcats "Felis silvestris lybica" circa 10,000 years ago, in the Near East. The domesticated cat and its closest wild ancestor are both diploid organisms that possess 38 chromosomes, in which over 200 heritable genetic defects have been identified, many homologous to human inborn errors. Specific metabolic defects have been identified underlying many of these feline diseases. There are several genes responsible for the hair color identified. The combination of them gives different phenotypes. Features like hair length, lack of tail, or presence of a very short tail (bobtail cat) are also determined by single alleles and modified by polygenes. The Cat Genome Project, sponsored by the Laboratory of Genomic Diversity at the U.S. National Cancer Institute Frederick Cancer Research and Development Center in Frederick, Maryland, focuses on the development of the cat as an animal model for human hereditary disease, infectious disease, genome evolution, comparative research initiatives within the family Felidae, and forensic potential. All felines, including the big cats, have a genetic anomaly that may prevent them from tasting sweetness, which is a likely factor for their indifference to or avoidance of fruits, berries, and other sugary foods. Feeding and diet. Cats feed on small prey such as insects, birds, and rodents. Feral cats, or house cats who are free-fed, consume about 8 to 16 small meals in a single day. Despite this, adult cats can adapt to being fed once a day. Cats are classified as obligate carnivores, because their physiology is geared toward efficient processing of meat, and lacks efficient processes for digesting plant matter. The cat cannot produce its own taurine (an essential organic acid), and, as it is contained in flesh, the cat must eat flesh to survive (see Taurine and cats). Similarly as with its teeth, a cat's digestive tract has become specialized over time to suit meat eating, having shortened in length only to those segments of intestine best able to break down proteins and fats from animal flesh. This trait severely limits the cat's ability to properly digest, metabolize, and absorb plant-derived nutrients, as well as certain fatty acids. For example, taurine is scarce in plants but abundant in meats. It is a key amino sulfonic acid for eye health in cats. Taurine deficiency can cause a condition called macular degeneration wherein the cat's retina slowly degenerates, eventually causing irreversible blindness. Despite the cat's meat-oriented physiology, it is still quite common for a cat to supplement its carnivorous diet with small amounts of grass, leaves, shrubs, houseplants, or other plant matter. One theory suggests this behavior helps cats regurgitate if their digestion is upset; another is that it introduces fiber or trace minerals into the diet. In this context, caution is recommended for cat owners because some houseplants are harmful to cats. For example, the leaves of the Easter Lily can cause permanent and life-threatening kidney damage to cats, and Philodendron are also poisonous to cats. The Cat Fanciers' Association has a full list of plants harmful to cats. There are several vegetarian or vegan commercially available cat foods supplemented with chemically synthesized taurine and other added nutrients that attempt to address nutritional shortfalls. Cats can be selective eaters (which may be due in some way to the aforementioned mutation which caused their species to lose sugar-tasting ability). However, cats generally cannot tolerate lack of food for more than 36 hours without risking liver damage. Cats have a fondness for catnip, which is sensed by their olfactory systems. Many enjoy consuming catnip, and most will often roll in it, paw at it, and occasionally chew on it. Cats also can also develop odd eating habits. Some cats like to eat or chew on other things like plastic, paper, string, wool, or even coal. This condition is called pica and can threaten the cat's health depending on the amount and toxicity of the non-food items eaten. The condition's name comes from the Latin word for magpie, a bird which is reputed to eat almost anything. Toxic sensitivity. The liver of a cat is less effective at detoxification than those of other animals, including humans and dogs; therefore exposure to many common substances considered safe for households may be dangerous to them. In general, the cat's environment should be examined for the presence of such toxins and the problem corrected or alleviated as much as possible; in addition, where sudden or prolonged serious illness without obvious cause is observed, the possibility of toxicity must be considered, and the veterinarian informed of any such substances to which the cat may have had access. For instance, the common painkiller paracetamol or acetaminophen, sold under brand names such as Tylenol and Panadol, is extremely toxic to cats; because they naturally lack enzymes needed to digest it, even minute portions of doses safe for humans can be fatal and any suspected ingestion warrants immediate veterinary attention. Even aspirin, which is sometimes used to treat arthritis in cats, is much more toxic to them than to humans and must be administered cautiously. Similarly, application of minoxidil (Rogaine) to the skin of cats, either accidental or by well-meaning owners attempting to counter loss of fur, has sometimes proved fatal. In addition to such obvious dangers as insecticides and weed killers, other common household substances that should be used with caution in areas where cats may be exposed to them include mothballs and other naphthalene products, as well as phenol based products often used for cleaning and disinfecting near cats' feeding areas or litter boxes, such as Pine-Sol, Dettol (Lysol), hexachlorophene, "etc." which, although they are widely used without problem, have been sometimes seen to be fatal. Ethylene glycol, often used as an automotive antifreeze, is particularly appealing to cats, and as little as a teaspoonful can be fatal. Essential oils are toxic to cats and there have been reported cases of serious illnesses caused by tea tree oil, and tea tree oil-based flea treatments and shampoos. Many human foods are somewhat toxic to cats; theobromine in chocolate can cause theobromine poisoning, for instance, although few cats will eat chocolate. Toxicity in cats ingesting relatively large amounts of onions or garlic has also been reported. Even such seemingly safe items as cat food packaged in pull tab tin cans have been statistically linked to hyperthyroidism; although the connection is far from proven, suspicion has fallen on the use of bisphenol A-based plastics, another phenol based product as discussed above, to seal such cans. Many houseplants are at least somewhat toxic to many species, cats included and the consumption of such plants by cats is to be avoided. Sociability. For cats, life in close proximity with humans (and other animals kept by humans) amounts to a "symbiotic social adaptation" which has developed over thousands of years. It has been suggested that, ethologically, the human keeper of a cat functions as a sort of surrogate for the cat's mother, and that adult domestic cats live their lives in a kind of extended kittenhood, a form of behavioral neoteny. Cats may express affection towards their human companions, especially if they imprint on them at a very young age and are treated with consistent affection. Regardless of the average sociability of any given cat or of cats in general, there are still any number of cats who meet or exceed the negative feline stereotype insofar as being poorly socialized. Older cats have also been reported to sometimes develop aggressiveness towards kittens, which may include biting and scratching; this type of behavior is known as Feline Asocial Aggression. Cohabitation. One may see natural house cat behavior by observing feral domestic cats, which are social enough to form colonies. Each cat in a colony holds a distinct territory, with sexually active males having the largest territories, and neutered cats having the smallest. Between these territories are neutral areas where cats watch and greet one another without territorial conflicts. Outside these neutral areas, territory holders usually aggressively chase away stranger cats, at first by staring, hissing, and growling, and if that does not work, by short but noisy and violent attacks. Despite cohabitation in colonies, cats do not have a social survival strategy, or a pack mentality. This mainly means that an individual cat takes care of all basic needs on its own (e.g., finding food, and defending itself), and thus cats are always lone hunters; they do not hunt in groups as dogs or lions do. Cats frequently tonguebathe themselves (see hygiene). The chemistry of their saliva, expended during their frequent grooming, appears to be a natural deodorant. Thus, a cat's cleanliness would aid in decreasing the chance a prey animal could notice the cat's presence. By contrast, dog odor is an advantage in hunting, for a dog is a pack hunter; part of the pack stations itself upwind, and its odor drives prey towards the rest of the pack stationed downwind. This requires a cooperative effort, which in turn requires communication skills. No such communication skills are required of a lone hunter. Fighting. When engaged in feline-to-feline combat for self-defense, territory, reproduction, or dominance, fighting cats make themselves appear more impressive and threatening by raising their fur and arching their backs, thus increasing their apparent size. Cats also behave this way while playing. Attacks usually comprise powerful slaps to the face and body with the forepaws as well as bites, but serious damage is rare; usually the loser runs away with little more than a few scratches to the face, and perhaps the ears. Cats will also throw themselves to the ground in a defensive posture to rake with their powerful hind legs. Normally, serious negative effects will be limited to possible infections of the scratches and bites, though these have been known to sometimes kill cats if untreated. In addition, such fighting is believed to be the primary route of transmission of feline immunodeficiency virus (FIV). Sexually active males will usually be in many fights during their lives, and often have decidedly battered faces with obvious scars and cuts to the ears and nose. Not only males will fight; females will also fight over territory or to defend their kittens. Play. Domestic cats, especially young kittens, are known for their love of play. This behavior mimics hunting and is important in helping kittens learn to stalk, capture, and kill prey. Many cats cannot resist a dangling piece of string, or a piece of rope drawn randomly and enticingly across the floor. This well known love of string is often depicted in cartoons and photographs, which show kittens or cats playing with balls of yarn. It is probably related to hunting instincts, including the common practice of kittens hunting their mother's and each other's tails. If string is ingested, however, it can become caught in the cat’s stomach or intestines, causing illness, or in extreme cases, death. Due to possible complications caused by ingesting a string, string play is sometimes replaced with a laser pointer's dot, which some cats will chase. While caution is called for, there are no documented cases of feline eye damage from a laser pointer, and the combination of precision needed and low energy involved make it a remote risk. A common compromise is to use the laser pointer to draw the cat to a prepositioned toy so the cat gets a reward at the end of the chase. A regular flashlight with a well-focused light spot has been commonly used in such play for decades, preceding the availability of consumer laser pointers. Cats will also engage in play fighting, with each other and with human partners. Humans "wrestling" with a supine cat, however, should be wary: if the cat is overstimulated or startled it may decide that the play has turned serious and cease to pull its punches; this can lead to serious scratches and occasionally even bites. Hunting. Much like their big cat relatives, domestic and feral cats are very effective predators. Domestic felines ambush or pounce upon and immobilize vertebrate prey using tactics similar to those of leopards and tigers. Having overpowered such prey, a cat delivers a lethal neck bite with its long canine teeth that either severs the prey's spinal cord with irreversible paralysis to prey, causes fatal bleeding by puncturing the carotid artery or the jugular vein, or asphyxiates the prey by crushing its trachea. One poorly understood element of cat hunting behavior is the presentation of prey to human owners. Ethologist Paul Leyhausen proposed that cats adopt humans into their social group, and share excess kill with others in the group according to the local pecking order, in which humans are placed at or near the top. However, anthropologist and animal scientist Desmond Morris in his 1986 book "Catwatching" suggests that when cats bring home mice or birds they have caught, they are teaching their human to hunt, or helping their human as if feeding (his words) "an elderly, inept kitten". Another possibility is that presenting the kill might be a relic of a kitten's behavior of demonstrating for its mother's approval that it has developed the necessary skill for hunting. Indoor cats will often retain their hunting instinct and deliver small household items to their owners, such as watches, pens, pencils, and other objects they can carry in their mouths. Reproduction. Cats are seasonally polyestrous, which means they may have many periods of heat over the course of a year. A heat period lasts about 4 to 7 days if the female is bred; if she is not, the heat period lasts longer. Multiple males will be attracted to a female in heat. The males will fight over her, and the victor wins the right to mate. At first, the female will reject the male, but eventually the female will allow the male to mate. The female will give a loud yowl as the male pulls out of her. After mating, the female will give herself a thorough wash. If a male attempts to breed with her at this point, the female will attack him. Once the female is done grooming, the cycle will repeat. The male cat's penis has spines which point backwards. Upon withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation. Because this does not always occur, females are rarely impregnated by the first male with which they mate. Furthermore, cats are superfecund; that is, a female may mate with more than one male when she is in heat, meaning different kittens in a litter may have different fathers. The gestation period for cats is approximately 63–65 days. The size of a litter averages three to five kittens, with the first litter usually smaller than subsequent litters. Kittens are weaned at between six and seven weeks, and cats normally reach sexual maturity at 5–10 months (females) and to 5–7 months (males). Cats are ready to go to new homes at about 12 weeks old (the recommended minimum age by Fédération Internationale Féline), or when they are ready to leave their mother. Cats can be surgically sterilized (spayed or castrated) as early as 6–8 weeks to limit unwanted reproduction. This surgery also prevents undesirable sex-related behavior, such as territory marking (spraying urine) in males and yowling (calling) in females. If a cat is neutered after such behavior has been learned, however, then the behavior may persist. Hygiene. Cats are known for their fastidious cleanliness. They groom themselves by licking their fur, employing their hooked papillae and saliva. As mentioned, their saliva is a powerful cleaning agent and deodorant. Many cats also enjoy grooming humans or other cats. Sometimes the act of grooming another cat is initiated as an assertion of superior position in the pecking order of a group (dominance grooming). Some cats occasionally regurgitate hairballs of fur that have collected in their stomachs as a result of their grooming. Longhaired cats are more prone to this than shorthaired cats. Hairballs can be	Eyes'" are organs that detect light, and send signals along the optic nerve to the visual and other areas of the brain. Complex optical systems with resolving power have come in ten fundamentally different forms, and 96% of animal species possess a complex optical system. Image-resolving eyes are present in cnidaria, mollusks, chordates, annelids and arthropods. The simplest "eyes", in even unicellular organisms, do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms. From more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment. Overview. Complex eyes can distinguish shapes and colors. The visual fields of many organisms, especially predators, involve large areas of binocular vision to improve depth perception; in other organisms, eyes are located so as to maximise the field of view, such as in rabbits and horses. The first proto-eyes evolved among animals 540 million years ago, about the time of the so-called Cambrian explosion. 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 6 of the thirty-something main phyla. In most vertebrates and some mollusks, the eye works by allowing light to enter it and project onto a light-sensitive panel of cells, known as the retina, at the rear of the eye. The cone cells (for color) and the rod cells (for low-light contrasts) in the retina detect and convert light into neural signals for vision. The visual signals are then transmitted to the brain via the optic nerve. Such eyes are typically roughly spherical, filled with a transparent gel-like substance called the vitreous humour, with a focusing lens and often an iris; the relaxing or tightening of the muscles around the iris change the size of the pupil, thereby regulating the amount of light that enters the eye, and reducing aberrations when there is enough light. The eyes of cephalopods, fish, amphibians and snakes usually have fixed lens shapes, and focusing vision is achieved by telescoping the lens — similar to how a camera focuses. Compound eyes are found among the arthropods and are composed of many simple facets which, depending on the details of anatomy, 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, which are arranged hexagonally, and which can give a full 360-degree field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera, have compound eyes of only a few facets, each with a retina capable of creating an image, creating vision. With each eye viewing a different thing, a fused image from all the eyes is produced in the brain, providing very different, high-resolution images. Possessing detailed hyperspectral color vision, the Mantis shrimp has been reported to have the world's most complex color vision system. Trilobites, which are now extinct, had unique compound eyes. They used clear calcite crystals to form the lenses of their eyes. In this, they differ from most other arthropods, which have soft eyes. The number of lenses in such an eye varied, however: some trilobites had only one, and some had thousands of lenses in one eye. In contrast to compound eyes, simple eyes are those that have a single lens. For example, jumping spiders have a large pair of simple eyes with a narrow field of view, supported by an array of other, smaller eyes for peripheral vision. Some insect larvae, like caterpillars, have a different type of simple eye (stemmata) which gives a rough image. Some of the simplest eyes, called ocelli, can be found in animals like some of the snails, which cannot actually "see" in the normal sense. They do have photosensitive cells, but no lens and no other means of projecting an image onto these cells. They can distinguish between light and dark, but no more. This enables snails to keep out of direct sunlight. In organisms dwelling near deep-sea vents, compound eyes have been secondarily simplified and adapted to spot the infra-red light produced by the hot vents in this way the bearers can spot hot springs and avoid being boiled alive. Evolution. Visual pigments appear to have a common ancestor and were probably involved in circadian rhythms or reproductive timing in simple organisms. Complex vision, associated with dedicated visual organs, or eyes, evolved many times in different lineages. Types of eye. Nature has produced ten different eye layouts — indeed every way of capturing an image has evolved at least once in nature, with the exception of zoom and Fresnel lenses. Eye types can be categorized into "simple eyes", with one concave chamber, 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 prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, 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 rhabdomic. These two groups are not monophyletic; the cnidaira also possess cilliated cells, Pit eyes. Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, 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 eye". 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. Pinhole eye. The pinhole eye is an "advanced" form of pit eye incorporating these improvements, most notably a small aperture (which may be adjustable) and deep pit. It is only found in the nautiloids. Without a lens to focus the image, it produces a blurry image, and will blur out a point to the size of the aperture. Consequently, nautiloids can't discriminate between objects with an angular separation of less than 11°. Shrinking the aperture would produce a sharper image, but let in less light. Spherical lensed eye. The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered — hence increasing the resolution obtainable. The most basic form, still 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. Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii. Heterogeneous eyes have evolved at least eight times — four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods. No 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 minimize the effect of eye motion while the animal moves, most such eyes have stabilizing eye muscles. The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. 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. Focussing the image would also cause the sun's image to be focussed 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). Weaknesses. One weakness of this eye construction is that chromatic aberration is still quite high — although for organisms without color vision, this is a very minor concern. A weakness of the vertebrate eye is the blind spot which results from a gap in the retina where the optic nerve exits at the back of the eye; the cephalopod eye has no blind spot as the retina is in the opposite orientation. Multiple lenses. Some marine organisms bear more than one lens; for instance the copeopod "Pontella" has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. "Copilla'"s eyes have two lenses, which move in and out like a telescope. Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): 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 terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air, relieving the lens of the function of reducing the focal length. This has freed it up for fine adjustments of focus, allowing a very high resolution to be obtained. As with spherical lenses, the problem of spherical aberration caused by the lens can be countered either by using an inhomogeneous lens material, or by flattening the lens. Flattening the lens has a disadvantage: the quality of vision is diminished away from the main line of focus, meaning that animals requiring all-round vision are detrimented. Such animals often display an inhomogeneous lens instead. As mentioned above, a refractive cornea is only useful out of water; in water, there is no difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures which have returned to the water — penguins and seals, for example — lose their refractive cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strong 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 rotifers, copeopods and platyhelminths use such organs, but these are too small to produce usable images. Some larger organisms, such as scallops, also use reflector eyes. The scallop "Pecten" has up to 100 millimeter-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses. Compound eyes. A compound eye may consist of thousands of individual photoreception units. 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 polarization of light. Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained. This can only be countered by increasing lens size and number — to see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their head. 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, and are also present in annelids and some bivalved molluscs. Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia. Apposition eyes. 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.) 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. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels. Superposition eyes. The second type is named the superposition eye. The superposition eye is divided into three types; 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 compound eye type, 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 has a transparent gap but uses corner mirrors 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. Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialized zones of ommatidia organized into a fovea area which gives acute vision. In the acute zone the eye are 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. The body of "Ophiocoma wendtii", a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons. Relationship to lifestyle. Eyes are generally adapted to the environment and lifestyle 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 African 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 amphipods 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. 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. Acuity. Visual acuity is often measured in cycles per degree (CPD), which measures an angular 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 gray block, is then the measurement of visual acuity of the eye. For a human eye with excellent acuity, the maximum theoretical resolution would be 50 CPD (1.2 arcminute 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 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, corresponds to 20 20 (normal vision) in humans. Color. All organisms are restricted to a small range of the electromagnetic spectrum; this varies from creature to creature, but is mainly between 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, 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 colors, seeing instead in shades of "grey"; color 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 cells, from which the more sensitive rod cells evolved. Even if organisms are physically capable of discriminating different colors, this does not necessarily mean that they can perceive the different colors; only with behavioral tests can this be deduced. Most organisms with color vision are able to 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 cells used for vision: the rods and the cones. Rods cannot distinguish colors, but are responsible for low-light black-and-white (scotopic) vision; they work well in dim light as they contain a pigment, visual purple, which is sensitive at low light intensity, but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are none at the fovea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina. Cones are responsible for color vision. They require brighter light to function than rods require. 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 colors). The color seen is the combined effect of stimuli 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 this spot, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibers of the optic nerve. When rods and cones are stimulated by light, the nerves send off impulses through these fibers to the brain. Pigment. 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 are the pigments involved in photoreception. Other pigments, such as melanin, 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 cilliary 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 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, 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.