ratio of word probabilities predicted from brain for dog and eye

dog	eye
top 10 words in brain distribution (in article): animal horse species wear breed cat wolf human hunt male	top 10 words in brain distribution (in article): water form light animal time type surface cause produce region
top 10 words in brain distribution (not in article): lion elephant century cell woman deer cattle forest hybrid saddle	top 10 words in brain distribution (not in article): ice rock drink river flow sea lamp occur soil ocean
times more probable under dog 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 dog'" ("Canis lupus familiaris",) is a domesticated subspecies of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The domestic dog has been one of the most widely kept working and companion animals in human history. The domestication of the gray wolf took place in a handful of events roughly 15,000 years ago in central Asia. The dog quickly became ubiquitous across culture in all parts of the world, and was extremely valuable to early human settlements. For instance, it is believed that the successful emigration across the Bering Strait might not have been possible without sled dogs. As a result of the domestication process, the dog developed a sophisticated intelligence that includes unparalleled social cognition and a simple theory of mind that is important to their interaction with humans. These social skills have helped the dog to perform in myriad roles, such as hunting, herding, protection, and, more recently, assisting handicapped individuals. Currently, there are estimated to be 400 million dogs in the world. Over the 15,000 year span that the dog had been domesticated, it diverged into only a handful of landraces, groups of similar animals whose morphology and behavior have been shaped by environmental factors and functional roles. Humans did not take an active, intentional role in this process until the last few hundred years. As the modern understanding of genetics developed, humans began to intentionally breed dogs for a wide range of specific traits. Through this process, the dog has developed into hundreds of varied breeds, and shows more behavioral and morphological variation than any other land mammal. For example, height measured to the withers ranges from a few inches in the Chihuahua to a few feet in the Irish Wolfhound; color varies from white through grays (usually called "blue'") to black, and browns from light (tan) to dark ("red" or "chocolate") in a wide variation of patterns; coats can be short or long, coarse-haired to wool-like, straight, curly, or smooth. It is common for most breeds to shed this coat, but non-shedding breeds are also popular. Etymology and related terminology. "Dog" is the common use term that refers to members of the subspecies "Canis lupus familiaris". The term is sometimes used to refer to a wider range of species: it can be used to refer to any mammal belonging to the family Canidae, which includes wolves, foxes, jackals, and coyotes; it can be used to refer to the subfamily of Caninae, or the genus Canis, also often called the "true dogs". Some members of the family have "dog" in their common names, such as the raccoon dog and the African wild dog. A few animals have "dog" in their common names but are not canids, such as the prairie dog and the dog fish. The English word "dog" can be traced back to the Old English "docga", a "powerful breed of canine". The term may derive from Proto-Germanic "*dukkōn", represented in Old English "finger-docce" ("finger-muscle"). Due to the linguistically archaic structure of the word, the term "dog" may ultimately derive from the earliest layer of Proto-Indo-European vocabulary, reflecting the role of the dog as the earliest domesticated animal. The English word "hound", which refers to a specific breed group in English, means "dog" in general in other Germanic languages; it is cognate to German "hund", Dutch "hond", common Scandinavian "hund", and Icelandic "hundur". "Hound" itself is derived from the Proto-Indo-European "*kwon-", which is also the direct root of the Greek κυων (kuōn) and the indirect root of the Latin "canis" through the variant form "*kani-". In breeding circles, a male canine is referred to as a "dog", while a female is called a "bitch". A group of offspring is a "litter". The father of a litter is called the "sire", and the mother is called the "dam". Offspring are generally called "pups" or "puppies" until they are about a year old. The process of birth is "whelping". Taxonomy and evolution. The domestic dog was originally classified as "Canis familiaris" and "Canis familiarus domesticus" by Linnaeus in 1758, and is currently classified as "Canis lupus familiaris", a subspecies of the gray wolf "Canis lupus", by the Smithsonian Institution and the American Society of Mammalogists. Overwhelming evidence from behavior, vocalizations, morphology, and molecular biology led to the contemporary scientific understanding that a single species, the gray wolf, is the common ancestor for all breeds of domestic dogs, however the timeframe and mechanisms by which dogs diverged are controversial. The current consensus among biologists and archaeologists is that no one can be sure when dogs were domesticated. There is conclusive evidence that dogs genetically diverged from their wolf ancestors at least 15,000 years ago but most believe domestication to have occurred much earlier. The evidence cited for an earlier divergence comes from archaeological findings and mitochondrial DNA studies, both of which are inconclusive. The archaeological evidence demonstrates that the domestication of dogs occurred prior to 15,000 years ago. Some genetic evidence indicates that the domestication of dogs from their wolf ancestors began in the late Upper Paleolithic close to the Pleistocene Holocene boundary, between 17,000 and 14,000 years ago. The earliest dog fossils, two large skulls from Russia and a mandible from Germany, date from roughly 14,000 years ago. Their likely ancestor is the large Eurasian wolf ("Canis lupus lupus"). Remains of smaller dogs from Natufian cave deposits in the Middle East have been dated to around 10,000 to 12,000 years ago. There is a great deal of archealogical evidence for dogs throughout Europe and Asia around this period and through the next two thousand years (roughly 8,000 to 10,000 years ago), with fossils uncovered in Germany, the French Alps, and Iraq, and cave paintings in Turkey. DNA studies have provided a wider range of possible divergence dates, from 15,000 to 40,000 years ago, to as much as 100,000 to 140,000 years ago. This evidence depends on a number of assumptions that others claim are violated. Genetic studies are based in comparisons of genetic diversity between species, and depend on a calibration date, such as the wolf-coyote divergence date, which is estimated to be roughly 1 million years ago. If this divergence date is closer to 750,000 or 2 million years ago, then genetic analyses would be interpreted very differently. Furthermore, it is believed that the genetic diversity of wolves has been in decline for the last 200 years, and that the genetic diversity of dogs has been reduced by selective breeding, which could bias DNA analyses to support an earlier divergence. The genetic evidence for the domestication event occurring in East Asia is also subject to violations of assumptions. These conclusions are based on the location of maximal genetic divergence, assumes that hybridization does not occur, and that breeds remain geographically localized. Although these assumptions hold for many species, there is good reason to believe that they do not hold for canines. Genetic analyses indicate all dogs are likely descended from a handful of domestication events with a small number of founding females, although there is evidence that domesticated dogs interbred with local populations of wild wolves on several occasions. Data suggests that dogs first diverged from wolves in East Asia, and that these domesticated dogs then quickly migrated throughout the world, reaching the North American continent around 8000 B.C. The oldest groups of dogs, which show the greatest genetic variability and are the most similar to their wolf ancestors, are primarily Asian and African breeds, including the Basenji, Saluki, Afghan Hound, Tibetan Terrier, Lhasa Apso, Chow Chow, Pekingese, Shar-Pei, Shi Tzu, Akita, Shiba Inu, Alaskan Malamute, Siberian Husky, and Samoyed. Some breeds that were thought to be very old, such as the Pharaoh Hound, Ibizan Hound, and Norwegian Elkhound, are now known to have been recreated more recently. There is a great deal of controversy surrounding the evolutionary framework for the domestication of dogs. At least three early species of the "Homo" genus began spreading out of Africa roughly 400,000 years ago, and thus lived for a considerable period in contact with canine species. Despite this, there is no evidence of any adaptation of these canine species to the presence of the close relatives of modern man. If dogs were domesticated, as believed, roughly 15,000 years ago, the event (or events) would have coincided with a large expansion in human territory and the development of agriculture. This has led some biologists to suggest that one of the forces that led to the domestication of dogs was a shift in human lifestyle in the form of established human settlements. Permanent settlements would have coincided with a greater amount of disposable food and would have created a barrier between wild and anthropogenic canine populations. Biology. Domestic dogs have been selectively bred for millennia for various behaviors, sensory capabilities, and physical attributes. Modern dog breeds show more variation in size, appearance, and behavior than any other domestic animal. Nevertheless, their morphology is based on that of their wild ancestors, gray wolves. Dogs are predators and scavengers, and like many other predatory mammals, the dog has powerful muscles, fused wrist bones, a cardiovascular system that supports both sprinting and endurance, and teeth for catching and tearing. Dogs are highly variable in height and weight. The smallest known dog was a Yorkshire Terrier, who stood only 6.3 cm (2.5 in) at the shoulder, 9.5 cm (3.75 in) in length along the head-and-body, and weighed only 113 grams (4 ounces). The largest known dog was an English Mastiff which weighed 155.6 kg (343 lbs) and was 250 cm (8.2 feet) from the snout to the tail. The tallest dog is a Great Dane that stands 106.7 cm (42.2 in) at the shoulder. Sight. The dog's visual system is engineered to serve the purposes of a hunter. While a dog's visual acuity is poor (that of a poodle's has been estimated to translate to a Snellen rating of 20 75), their visual discrimination for moving objects is very high; dogs have been shown to be able to discriminate between humans (i.e., identifying their owner) from distances up to a mile. As crepuscular hunters, dogs often rely on their vision in low light situations: they have very large pupils, a high density of rods in the fovea, an increased flicker rate, and a tapetum lucidum. The tapetum is a reflective surface behind the retina that reflects light back to give the photoreceptors a second chance to catch the photons. Like most mammals, dogs are dichromats and have color vision equivalent to red-green color blindness in humans. The eyes of different breeds of dogs have different shapes, dimensions, and retina configurations. Many long-nosed breeds have a "visual streak" — a wide foveal region that runs across the width of the retina and gives them a very wide field of excellent vision. Some long-muzzled breeds, particularly the sighthounds, have a field of vision up to 270° (compared to 180° for humans). Short-nosed breeds, on the other hand, have an "area centralis": a central patch with up to three times the density of nerve endings as the visual streak, giving them detailed sight much more like a human's. Some broad-headed breeds with short noses have a field of vision similar to that of humans. Most breeds have good vision, but some show a genetic predisposition for myopia — such as Rottweilers, where one out of every two has been found to be myopic. Hearing. The frequency range of dog hearing is approximately 40 Hz to 60,000 Hz, which means that dogs can detect sounds outside both ends of the human auditory spectrum. Additionally, dogs have ear mobility which allows them to rapidly pinpoint the exact location of a sound. Eighteen or more muscles can tilt, rotate, raise, or lower a dog's ear. A dog can identify a sound's location much faster than a human can, as well as hear sounds at four times the distance. Smell. While the human brain is dominated by a large visual cortex, the dog brain is largely dominated by an olfactory cortex. The olfactory bulb in dogs is roughly forty times bigger than the olfactory bulb in humans, relative to total brain size, with 125 to 220 million smell-sensitive receptors. The bloodhound exceeds this standard with nearly 300 million receptors. Dogs can discriminate odors at concentrations nearly 100 million times lower than humans can. Coat. The coats of domestic dogs are either "double", made up of a coarse topcoat and a soft undercoat, like a wolf, or "single", with the topcoat only. Dogs with double coats tend to originate in colder climates. Domestic dogs often display the remnants of countershading, a common natural camouflage pattern. The general theory of countershading is that an animal that is lit from above will appear lighter on its upper half and darker on its lower half, where it will usually be in its own shade. This is a pattern that predators can learn to watch for. A countershaded animal will have dark coloring on its upper surfaces and light coloring below, which reduces its general visibility. Thus many breeds will have an occasional "blaze", stripe, or "star" of white fur on their chest or underside. Tail. There are many different shapes for dog tails: straight, straight up, sickle, curled, or cork-screw. In some breeds, the tail is traditionally docked to avoid injuries (especially for hunting dogs). In some breeds, puppies can be born with a short tail or no tail at all. This occurs more frequently in those breeds that are frequently docked and thus have no breed standard regarding the tail. Types and breeds. While all dogs are genetically very similar, natural selection and selective breeding have reinforced certain characteristics in certain populations of dogs, giving rise to dog types and dog breeds. Dog types are broad categories based on function, genetics, or characteristics. Dog breeds are groups of animals that possess a set of inherited characteristics that distinguishes them from other animals within the same species. Modern dog breeds are non-scientific classifications of dogs kept by modern kennel clubs. Purebred dogs of one breed are genetically distinguishable from purebred dogs of other breeds, but the means by which kennel clubs classify dogs is unsystematic. Systematic analyses of the dog genome has revealed only four major types of dogs that can be said to be statistically distinct. These include the "old world dogs" (e.g., Malamute and Shar-Pei), "Mastiff"-type (e.g., Labrador Retriever), "herding"-type (e.g., St. Bernard), and "all others" (also called "modern"- or "hunting"-type). Health. Dogs are susceptible to various diseases, ailments, and poisons, some of which can affect humans. To defend against many common diseases, dogs are often vaccinated. Some breeds of dogs are prone to certain genetic ailments such as elbow or hip dysplasia, blindness, deafness, pulmonic stenosis, cleft palate, and trick knees. Two serious medical conditions particularly affecting dogs are pyometra, affecting unspayed females of all types and ages, and bloat, which affects the larger breeds or deep-chested dogs. Both of these are acute conditions, and can kill rapidly. Dogs are also susceptible to parasites such as fleas, ticks, and mites, as well as hookworms, tapeworms, roundworms, and heartworms. Dogs are also vulnerable to some of the same health conditions as humans, including diabetes, dental and heart disease, epilepsy, cancer, hypothyroidism, and arthritis. Mortality. The typical lifespan of dogs varies widely among breeds, but for most the median longevity, the age at which half the dogs in a population have died and half are still alive, ranges from 10 to 13 years. Individual dogs may live well beyond the median of their breed. The breed with the shortest lifespan (among breeds for which there is a questionnaire survey with a reasonable sample size) is the Dogue de Bordeaux, with a median longevity of about 5.2 years, but several breeds, including Miniature Bull Terriers, Bulldogs, Nova Scotia Duck-Tolling Retrievers, Bloodhounds, Irish Wolfhounds, Greater Swiss Mountain Dogs, Great Danes, and Mastiffs, are nearly as short-lived, with median longevities of 6 to 7 years. The longest-lived breeds, including Toy Poodles, Border Terriers, Miniature Dachshunds, Miniature Poodles, and Tibetan Spaniels, have median longevities of 14 to 15 years. The median longevity of mixed breed dogs, taken as an average of all sizes, is one or more years longer than that of purebred dogs when all breeds are averaged. The dog widely reported to be the longest-lived is "Bluey," who died in 1939 and was claimed to be 29.5 years old at the time of his death; however, the Bluey record is anecdotal and unverified. The longest verified records are of dogs living for 24 years. Predation. Although wild dogs, like wolves, are apex predators, they can be killed in territory disputes with wild animals. Furthermore, in areas where both dogs and other large predators live, dogs can be a major food source for big cats or canines. Reports from Croatia indicate that dogs are killed more frequently than sheep. Wolves in Russia apparently limit feral dog populations. In Wisconsin, more compensation has been paid for dog losses than livestock. Some wolf pairs have been reported to prey on dogs by having one wolf lure the dog out into heavy brush where the second animal waits in ambush. In some instances, wolves have displayed an uncharacteristic fearlessness of humans and buildings when attacking dogs, to the extent that they have to be beaten off or killed. Coyotes and big cats have also been known to attack dogs. Leopards in particular are known to have a predilection for dogs, and have been recorded to kill and consume them regardless of the dog's size or ferocity. Tigers in Manchuria, Indochina, Indonesia, and Malaysia, are reputed to kill dogs with the same vigor as leopards. Striped Hyenas are major predators of village dogs in Turkmenistan, India, and the Caucasus. Diet. Despite its descent from wolves, the domestic dog is an omnivore, though it is classified in the order Carnivora. Unlike an obligate carnivore, such as a member of the cat family with its shorter small intestine, a dog is neither dependent on meat-specific protein nor a very high level of protein in order to fulfill its basic dietary requirements. Dogs are able to healthily digest a variety of foods, including vegetables and grains, and can consume a large proportion of these in their diet. In the wild, canines often eat available plants and fruits. Reproduction. In domestic dogs, sexual maturity begins to happen around age six to twelve months for both males and females, although this can be delayed until up to two years old for some large breeds. This is the time at which female dogs will have their first estrous cycle. They will experience subsequent estrous cycles biannually, during which the body prepares for pregnancy. At the peak of the cycle, females will come into estrus, being mentally and physically receptive to copulation. Because the ova survive and are capable of being fertilized for a week after ovulation, it is possible for a female to mate with more than one male. Adolescence for most domestic dogs is around 12 to 15 months, beyond which they are for the most part more adult than puppy. Domestication has selectively bred for higher libido and earlier and more frequent breeding cycles in dogs than in their wild ancestors, and dogs remain reproductively active until old age. Dogs bear their litters roughly 56 to 72 days after fertilization, with an average of 63 days, although the length of gestation can vary. An average litter consists of about six puppies, though this number may vary widely based on the breed of dog. Toy dogs generally produce from one to four puppies in each litter, while much larger breeds may average as many as twelve. Some dog breeds have acquired traits through selective breeding that interfere with reproduction. Male French Bulldogs, for instance, are incapable of mounting the female. For many dogs of this breed, the female must be artificially inseminated in order to reproduce. Spaying and neutering. Neutering refers to the sterilization of animals, usually by removal of the male's testicles or the female's ovaries and uterus, in order to eliminate the ability to procreate and reduce sex drive. Because of the overpopulation of dogs in some countries, animal control agencies, such as the American Society for the Prevention of Cruelty to Animals (ASPCA), advise that dogs not intended for further breeding should be neutered, so that they do not have undesired puppies that may have to be destroyed later. According to the Humane Society of the United States, 3–4 million dogs and cats are put down each year in the United States and many more are confined to cages in shelters because there are many more animals than there are homes. Spaying or castrating dogs helps keep overpopulation down. Local humane societies, SPCAs, and other animal protection organizations urge people to neuter their pets and to adopt animals from shelters instead of purchasing them. Several notable public figures have spoken out against animal overpopulation, including Bob Barker. On his game show, "The Price Is Right", Barker stressed the problem at the end of every episode, saying: "Help control the pet population. Have your pets spayed or neutered." The current host, Drew Carey, makes a similar plea at the conclusion of each episode. Neutering reduces problems caused by hypersexuality, especially in male dogs. Spayed female dogs are less likely to develop some forms of cancer, affecting mammary glands, ovaries, and other reproductive organs. However, neutering increases the risk of urinary incontinence in female dogs, and prostate cancer in males, as well as osteosarcoma, hemangiosarcoma, cruciate ligament rupture, obesity, and diabetes mellitus in either gender. The hormonal changes involved with sterilization can change the animal's personality and metabolism. Recent studies proved that spayed and neutered dogs in general are more aggressive towards people and other dogs, and more fearful and sensitive to touch than dogs than had not been sterilized. Spaying or neutering very young animals ("early-age spay"), can result in increased health concerns later on in life for both sexes. Incontinence in female dogs is made worse by spaying too early. In both males and females, alteration causes changes in hormones during development. This inhibits the natural signals needed for proper body development, leading to larger animals with greater risk for hip dysplasia, osteoporosis, and other joint disorders. Other studies have shown, however, that early-age neutering of male dogs is associated with no major risks compared to neutering at the more traditional age of six months. Behavior and intelligence. Although dogs have been the subject of a great deal of Behaviorist psychology (e.g.,Pavlov's Dog), they do not enter the world with a psychological "blank slate". Rather, dog behavior is affected by genetic factors as well as environmental factors. Domestic dogs exhibit a number of behaviors and predispositions that were inherited from wolves. The grey wolf is a social animal that has evolved a sophisticated means of communication and social structure. The domestic dog has inherited some of these predispositions, but many of the salient characteristics in dog behavior have been largely shaped by selective breeding by humans. Thus, some of these characteristics, such as the dog's highly developed social cognition, are found only in primitive forms in grey wolves. Intelligence. "Intelligence" is an umbrella term that encompasses the faculties involved in a wide range of mental tasks, such as learning, problem-solving, and communication. The domestic dog has a predisposition to exhibit a social intelligence that is uncommon in the animal world. Dogs are capable of learning in a number of ways, such as through simple reinforcement learning (e.g.,classical or operant conditioning) and by observation. Dogs go through a series of stages of cognitive development. They are not born with an understanding of object permanence; the understanding that objects which are not being actively perceived still remain in existence. The understanding of object permanence occurs during as the infant is learning the coordination of secondary circular reactions, as described by Jean Piaget. That is, as the infants learn to interact intentionally with objects around it. For dogs, this occurs at roughly 8 weeks of age. Handlers of working dogs such as herding dogs and sled dogs know that pups learn behaviors quickly by following examples set by experienced dogs. Many of these behaviors are allelomimetic in that they depend on a genetic predisposition to learn and imitate behaviors of other dogs. This form of intelligence is not peculiar to those tasks dogs have been bred to perform, but can be generalized to myriad abstract problems. Adler & Alder demonstrated this by giving Dachshund puppies the task of learning to pull a cart by tugging on an attached piece of ribbon in order to get a reward in the cart. Puppies that watched an experienced dog successfully retrieve the reward in this way learned the task fifteen times faster than those who were left to solve the problem on their own. In addition to learning by example from other dogs, dogs have also been shown to learn by mimicking human behaviors. In another study, puppies were presented with a box, and shown that when a handler pressed a lever, a ball would roll out of the box. The handler then allowed the puppy to play with the ball, making it an intrinsic reward. The pups were then allowed to interact with the box. Roughly three-quarters of the puppies subsequently touched the lever, and over half successfully released the ball, compared to only 6 percent in a control group that did not watch the human manipulate the lever. Furthermore, the ability for dogs to learn by example has been shown to be as effective as operant conditioning. McKinley and Young have demonstrated that dogs show equivalent accuracy and learning times when taught to identify an object by operant conditioning as they do when taught by human example. Their study found that handing an object between experimenters who then use its name in a sentence successfully taught an	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.