ratio of word probabilities predicted from brain for horse and eye

horse	eye
top 10 words in brain distribution (in article): species animal horse wear female male time breed food human	top 10 words in brain distribution (in article): water light form time type animal surface produce cause region
top 10 words in brain distribution (not in article): bird egg fish cat ant kill bee build woman nest	top 10 words in brain distribution (not in article): ice drink lamp rock wine river key flow beer state
times more probable under horse 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 horse'" ("Equus ferus caballus") is a hoofed (ungulate) mammal, a subspecies of one of seven extant species of the family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today. Humans began to domesticate horses around 4000 BC, and their domestication is believed to have been widespread by 3000 BC; by 2000 BC the use of domesticated horses had spread throughout the Eurasian continent. Although most horses today are domesticated, there are still endangered populations of the Przewalski's Horse, the only remaining true wild horse, as well as more common feral horses which live in the wild but are descended from domesticated ancestors. There is an extensive, specialized vocabulary used to describe equine-related concepts, covering everything from anatomy to life stages, size, colors, markings, breeds, locomotion, and behavior. Horses are anatomically designed to use speed to escape predators, and have a well-developed sense of balance and a strong fight-or-flight instinct. Related to this need to flee from predators in the wild is an unusual trait: horses are able to sleep both standing up and lying down. Female horses, called mares, carry their young for approximately 11 months, and a young horse, called a foal, can stand and run shortly following birth. Most domesticated horses begin training under saddle or in harness between the ages of two and four. They reach full adult development by age five, and have an average lifespan of between 25 and 30 years. Horse breeds are loosely divided into three categories based on general temperament: spirited "hot bloods" with speed and endurance; "cold bloods," such as draft horses and some ponies, suitable for slow, heavy work; and "warmbloods," developed from crosses between hot bloods and cold bloods, often focusing on creating breeds for specific riding purposes, particularly in Europe. There are over 300 breeds of horses in the world today, developed for many different uses. Horses and humans interact in many ways, not only in a wide variety of sport competitions and non-competitive recreational pursuits, but also in working activities including police work, agriculture, entertainment, assisted learning and therapy. Horses were historically used in warfare. A wide variety of riding and driving techniques have been developed, using many different styles of equipment and methods of control. Many products are derived from horses, including meat, milk, hide, hair, bone, and pharmaceuticals extracted from the urine of pregnant mares. Humans provide domesticated horses with food, water and shelter, as well as attention from specialists such as veterinarians and farriers. Biology. Horse anatomy is described by a large number of specific terms, as illustrated by the chart to the right. Specific terms also describe various ages, colors and breeds. Age. Depending on breed, management and environment, the domestic horse today has a life expectancy of 25 to 30 years. It is uncommon, but a few animals live into their 40s and, occasionally, beyond. The oldest verifiable record was "Old Billy," a 19th-century horse that lived to the age of 62. In modern times, Sugar Puff, who had been listed in the Guinness Book of World Records as the world's oldest living pony, died in 2007, aged 56. Regardless of a horse's actual birth date, for most competition purposes an animal is considered a year older on January 1 of each year in the northern hemisphere and August 1 in the southern hemisphere. The exception is in endurance riding, where the minimum age to compete is based on the animal's calendar age. A very rough estimate of a horse's age can be made from looking at its teeth. The following terminology is used to describe horses of various ages: In horse racing, the definitions of colt, filly, mare, and stallion may differ from those given above. In the UK, Thoroughbred horse racing defines a colt as a male less than five years old, and a filly as a female less than five years old. In the USA, both Thoroughbred racing and harness racing defines colts and fillies as four years old and younger. Size. The English-speaking world measures the height of horses in hands, abbreviated "h" or "hh," for "hands high," measured at the highest point of an animal's withers, where the neck meets the back, chosen as a stable point of the anatomy, unlike the head or neck, which move up and down; one hand is. Intermediate heights are defined by hands and inches, rounding to the lower measurement in hands, followed by a decimal point and the number of additional inches between 1 and 3. Thus a horse described as "15.2 h," is 15 hands, 2 inches in height. The size of horses varies by breed, but can also be influenced by nutrition. The general rule for cutoff in height between what is considered a horse and a pony at maturity is 14.2 hands. An animal 14.2 h or over is usually considered to be a horse and one less than 14.2 h a pony. However, there are exceptions to the general rule. Some breeds which typically produce individuals both under and over 14.2 h are considered horses regardless of their height. Conversely, some pony breeds may have features in common with horses, and individual animals may occasionally mature at over 14.2 h, but are still considered to be ponies. The distinction between a horse and pony is not simply a difference in height, but takes account of other aspects of "phenotype" or appearance, such as conformation and temperament. Ponies often exhibit thicker manes, tails and overall coat. They also have proportionally shorter legs, wider barrels, heavier bone, shorter and thicker necks, and short heads with broad foreheads. They often have calmer temperaments than horses and also a high level of equine intelligence that may or may not be used to cooperate with human handlers. In fact, small size, by itself, is sometimes not a factor at all. While the Shetland pony stands on average 10 hands high, the Falabella and other miniature horses, which can be no taller than, the size of a medium-sized dog, are classified by their respective registries as very small horses rather than as ponies. Light riding horses such as Arabians, Morgans, or Quarter Horses usually range in height from 14 to 16 hands and can weigh from. Larger riding horses such as Thoroughbreds, American Saddlebreds or Warmbloods usually start at about 15.2 hands and often are as tall as 17 hands, weighing from. Heavy or draft horses such as the Clydesdale, Belgian, Percheron, and Shire are usually at least 16 to 18 hands high and can weigh from about. The largest horse in recorded history was probably a Shire horse named Sampson, who lived during the late 1800s. He stood 21.2½ hands high, and his peak weight was estimated at. The current record holder for the world's smallest horse is Thumbelina, a fully mature miniature horse affected by dwarfism. She is tall and weighs. Colors and markings. Horses exhibit a diverse array of coat colors and distinctive markings, described with a specialized vocabulary. Often, a horse is classified first by its coat color, before breed or sex. Flashy or unusual colors are sometimes very popular, as are horses with particularly attractive markings. Horses of the same color may be distinguished from one another by their markings. The genetics that create many horse coat colors have been identified, although research continues on specific genes and mutations that result in specific color traits. Essentially, all horse colors begin with a genetic base of "red" (chestnut) or "black," with the addition of alleles for spotting, graying, suppression or dilution of color, or other effects acting upon the base colors to create the dozens of possible coat colors found in horses. Horses which are light in color are often misnamed as being "white" horses. A horse that looks pure white is, in most cases, actually a middle-aged or older gray. Grays have black skin underneath their white hair coat (with the exception of small amounts of pink skin under white markings). The only horses properly called white are those with pink skin under a white hair coat, a fairly rare occurrence. There are no truly albino horses, with pink skin and red eyes, as albinism is a lethal condition in horses. Reproduction and development. Pregnancy lasts for approximately 335–340 days and usually results in one foal. Twins are very rare. Colts are carried on average about 4 days longer than fillies. Horses are a precocial species, and foals are capable of standing and running within a short time following birth. Horses, particularly colts, may sometimes be physically capable of reproduction at about 18 months. In practice, individuals are rarely allowed to breed before the age of three, especially females. Horses four years old are considered mature, although the skeleton normally continues to develop until the age of six; the precise time of completion of development also depends on the horse's size, breed, gender, and the quality of care provided by its owner. Also, if the horse is larger, its bones are larger; therefore, not only do the bones take longer to actually form bone tissue, but the epiphyseal plates are also larger and take longer to convert from cartilage to bone. These plates convert after the other parts of the bones, but are crucial to development. Depending on maturity, breed, and the tasks expected, young horses are usually put under saddle and trained to be ridden between the ages of two and four. Although Thoroughbred race horses are put on the track at as young as two years old in some countries, horses specifically bred for sports such as dressage are generally not entered into top-level competition until they are a minimum of four years old, because their bones and muscles are not solidly developed, nor is their advanced training complete. For endurance riding competition, horses are not deemed mature enough to compete until they are a full 60 calendar months (5 years) old. Skeletal system. Horses have a skeleton that averages 205 bones. A significant difference between the horse skeleton, compared to that of a human, is the lack of a collarbone—the horse's front limb system is attached to the spinal column by a powerful set of muscles, tendons and ligaments that attach the shoulder blade to the torso. The horse's legs and hooves are also unique structures. Their leg bones are proportioned differently from those of a human. For example, the body part that is called a horse's "knee" is actually made up of the carpal bones that correspond to the human wrist. Similarly, the hock, contains the bones equivalent to those in the human ankle and heel. The lower leg bones of a horse correspond to the bones of the human hand or foot, and the fetlock (incorrectly called the "ankle") is actually the proximal sesamoid bones between the cannon bones (a single equivalent to the human metacarpal or metatarsal bones) and the proximal phalanges, located where one finds the "knuckles" of a human. A horse also has no muscles in its legs below the knees and hocks, only skin, hair, bone, tendons, ligaments, cartilage, and the assorted specialized tissues that make up the hoof. Hooves. The critical importance of the feet and legs is summed up by the traditional adage, "no foot, no horse". The horse hoof begins with the distal phalanges, the equivalent of the human fingertip or tip of the toe, surrounded by cartilage and other specialized, blood-rich soft tissues such as the laminae. The exterior hoof wall and horn of the sole is made of essentially the same material as a human fingernail. The end result is that a horse, weighing on average, travels on the same bones as a human on tiptoe. For the protection of the hoof under certain conditions, some horses have horseshoes placed on their feet by a professional farrier. The hoof continually grows, and needs to be trimmed (and horseshoes reset, if used) every five to eight weeks. Teeth. Horses are adapted to grazing. In an adult horse, there are 12 incisors, adapted to biting off the grass or other vegetation, at the front of the mouth. There are 24 teeth adapted for chewing, the premolars and molars, at the back of the mouth. Stallions and geldings have four additional teeth just behind the incisors, a type of canine teeth that are called "tushes." Some horses, both male and female, will also develop one to four very small vestigial teeth in front of the molars, known as "wolf" teeth, which are generally removed because they can interfere with the bit. There is an empty interdental space between the incisors and the molars where the bit rests directly on the bars (gums) of the horse's mouth when the horse is bridled. The incisors show a distinct wear and growth pattern as the horse ages, as well as change in the angle at which the chewing surfaces meet. The teeth continue to erupt throughout life as they are worn down by grazing, so a very rough estimate of a horse's age can be made by an examination of its teeth, although diet and veterinary care can affect the rate of tooth wear. Digestion. Horses are herbivores with a digestive system adapted to a forage diet of grasses and other plant material, consumed steadily throughout the day. Therefore, compared to humans, they have a relatively small stomach but very long intestines to facilitate a steady flow of nutrients. A horse will eat of food per day and, under normal use, drink to of water. Horses are not ruminants, so they have only one stomach, like humans, but unlike humans, they can also digest cellulose from grasses due to the presence of a "hind gut" called the cecum, or "water gut," which food goes through before reaching the large intestine. Unlike humans, horses cannot vomit, so digestion problems can quickly cause colic, a leading cause of death. Senses. The horse's senses are generally superior to those of a human. As prey animals, they must be aware of their surroundings at all times. They have the largest eyes of any land mammal, and because their eyes are positioned on the sides of their heads, horses have a range of vision of more than 350°, with approximately 65° of this being binocular (seen with both eyes) and the remaining 285° monocular (seen with only one eye). Horses have excellent day and night vision, but studies indicate that they have two-color, or dichromatic vision; their color vision is somewhat like red-green color blindness in humans. This means that certain colors, especially red and related colors, appear more green. Their hearing is good, and the pinna of each ear can rotate up to 180°, giving the potential for 360° hearing without having to move the head. Their sense of smell, while much better than that of humans, is not their strongest asset; they rely to a greater extent on vision. Horses have a great sense of balance, due partly to their ability to feel their footing and partly to highly developed proprioceptive abilities (the unconscious sense of where the body and limbs are at all times). A horse's sense of touch is well developed. The most sensitive areas are around the eyes, ears and nose. Via touch, horses perceive and respond immediately to changes in their environment, sensing contact as subtle as an insect landing anywhere on the body. Horses have an advanced sense of taste that allows them to sort through grains and grasses to choose what they would most like to eat, and their prehensile lips can easily sort even the smallest grains. Horses generally will not eat poisonous plants. However, there are exceptions and horses will occasionally eat toxic amounts of poisonous plants even when there is adequate healthy food. Movement. All horses move naturally with four basic gaits: the four-beat walk, which averages four miles per hour; the two-beat trot or jog, which averages per hour (faster for harness racing horses); and the leaping gaits known as the canter or lope (a three-beat gait that is per hour), and the gallop. The gallop averages per hour. The world record for a horse galloping over a short, sprint distance is per hour. Besides these basic gaits, some horses perform a two-beat pace, instead of the trot. In addition, there are several four-beat "ambling" gaits that are approximately the speed of a trot or pace, though smoother to ride. These include the lateral slow gait, rack, running walk, and tölt as well as the diagonal fox trot. Ambling gaits are often genetic traits in specific breeds, known collectively as gaited horses. In most cases, gaited horses replace the standard trot with one of the ambling gaits. Behavior. Horses are prey animals with a well-developed fight-or-flight instinct. Their first response to threat is to startle and usually flee, although they are known to stand their ground and defend themselves or their offspring in cases where flight is not possible, or when their young are threatened. They also tend to be curious; when startled, they will often hesitate an instant to ascertain the cause of their fright, and may not always flee from something that they perceive as non-threatening. Through selective breeding, some breeds of horses are quite docile, particularly certain large draft horses. Most light horse riding breeds were developed for speed, agility, alertness and endurance; natural qualities that extend from their wild ancestors. Horses are herd animals, with a clear hierarchy of rank, led by a dominant animal (usually a mare). They are also social creatures who are able to form companionship attachments to their own species and to other animals, including humans. They communicate in various ways, including vocalizations such as nickering or whinnying, mutual grooming, and body language. Many horses will become difficult to manage if they are isolated. Through proper training, it is possible to teach any horse to accept a human as a type of companion, and thus be comfortable away from other horses. When confined with insufficient companionship, exercise or stimulation, individuals may develop stable vices, an assortment of bad habits, mostly psychological in origin, that include wood chewing, wall kicking, "weaving" (rocking back and forth) and other problems. Intelligence and learning. In the past, horses were considered unintelligent, with no abstract thinking ability, unable to generalize, and driven primarily by a herd mentality. However, recent studies show that they perform a number of cognitive tasks on a daily basis, and frequently engage in mental challenges that include food procurement and social system identification. They have also been shown to have good spatial discrimination abilities. Studies have assessed equine intelligence in the realms of problem solving, learning speed, and knowledge retention. Results show that horses excel at simple learning, but also are able to solve advanced cognitive challenges that involve categorization and concept learning. They have been shown to learn from habituation, desensitization, Pavlovian conditioning, and operant conditioning. They respond to and learn from both positive and negative reinforcement. Domesticated horses tend to face greater mental challenges than wild horses, due to living in artificial environments that stifle instinctual behavior while learning tasks that are not natural. Horses are creatures of habit that respond and adapt well to regimentation, and respond best when the same routines and techniques are used consistently. Some trainers believe that "intelligent" horses are reflections of intelligent trainers who effectively use response conditioning techniques and positive reinforcement to train in the style that fits best with an individual animal's natural inclinations. Others who handle horses regularly note that personality also may play a role separate from intelligence in determining how a given animal responds to various experiences. Temperament. Thoroughbred race horses are a "hot blooded" breed. Horses are mammals, and as such are "warm-blooded" creatures, as opposed to reptiles, which are cold-blooded. However, these words have developed a separate meaning in the context of equine terminology, used to describe temperament, not body temperature. For example, the "hot-bloods," such as many race horses, exhibit more sensitivity and energy, while the "cold-bloods," such as most draft breeds, are quieter and calmer. "Hot blooded" breeds include "oriental horses" such as the Akhal-Teke, Barb, Arabian horse and now-extinct Turkoman horse, as well as the Thoroughbred, a breed developed in England from the older oriental breeds. Hot bloods tend to be spirited, bold, and learn quickly. They are bred for agility and speed. They tend to be physically refined—thin-skinned, slim, and long-legged. The original oriental breeds were brought to Europe from the Middle East and North Africa when European breeders wished to infuse these traits into racing and light cavalry horses. The "cold blooded" draft breeds are powerful and heavily-muscled Muscular, heavy draft horses are known as "cold bloods," as they are bred not only for strength, but also to have the calm, patient temperament needed to pull a plow or a heavy carriage full of people. They are sometimes nicknamed "gentle giants." Well-known draft breeds include the Belgian and the Clydesdale. Some, like the Percheron are lighter and livelier, developed to pull carriages or to plow large fields in drier climates. Others, such as the Shire, are slower and more powerful, bred to plow fields with heavy, clay-based soils. The cold-blooded group also includes some pony breeds. "Warmblood" breeds, such as the Trakehner or Hanoverian, developed when European carriage and war horses were crossed with Arabians or Thoroughbreds, producing a riding horse with more refinement than a draft horse, but greater size and more phlegmatic temperament than a lighter breed. Certain pony breeds with warmblood characteristics have been developed for smaller riders. A modern "Warmblood" horse is large, but agile and athletic. Today, the term "Warmblood" refers to a specific subset of sport horse breeds that have dominated the Olympic Games and international FEI competition in dressage and show jumping since the 1970s. Prior to that time, the term "warm blood" often referred to any cross between cold-blooded and hot-blooded breeds. Examples included breeds such as the Irish Draught or the Cleveland Bay. Less often, the term was even used to refer to breeds of light riding horse other than Thoroughbreds or Arabians, such as the Morgan horse. Sleep patterns. Horses are able to sleep both standing up and lying down. In an adaptation from life in the wild, horses are able to enter light sleep by using a "stay apparatus" in their legs, allowing them to doze without collapsing. Horses sleep better when in groups because some animals will sleep while others stand guard to watch for predators. A horse kept alone will not sleep well because its instincts are to keep a constant eye out for danger. Unlike humans, horses do not sleep in a solid, unbroken period of time, but take many short periods of rest. Horses may spend anywhere from four to fifteen hours a day in standing rest, and from a few minutes to several hours lying down. Total sleep time in a day may range from several minutes to a couple of hours, mostly in short intervals of about 15 minutes each. Horses must lie down to reach REM sleep. They only have to lie down for an hour or two every few days to meet their minimum REM sleep requirements. However, if a horse is never allowed to lie down, after several days it will become sleep-deprived, and in rare cases may suddenly collapse as it involuntarily slips into REM sleep while still standing. This condition differs from narcolepsy, although horses may also suffer from that disorder. Taxonomy and evolution. The horse as it is known today adapted by evolution to survive in areas of wide-open terrain with sparse vegetation, surviving in an ecosystem where other large grazing animals, especially ruminants, could not. Horses and other equids are odd-toed ungulates of the order Perissodactyla, a group of mammals that was dominant during the Tertiary period. In the past, this order contained 14 families and many species, but only three families—Equidae (the horse and related species), the tapir and the rhinoceros—containing 18 known species have survived to the present day. The earliest known member of the Equidae family was the "Hyracotherium", which lived between 45 and 55 million years ago, during the Eocene period and had 4 toes on each front foot, and 3 toes on each back foot. The extra toe on the front feet soon disappeared with the "Mesohippus", which lived 32 to 37 million years ago, and by about 5 million years ago, the modern Equus had developed. The extra side toes shrank in size until they have vanished in modern horses. All that remains is a set of small vestigial bones on the leg above the hoof, known informally as ergots, chestnuts, or splint bones. Their legs also lengthened as their toes disappeared and until they were a hoofed animal capable of running at great speed. Over millions of years, equid teeth also evolved from browsing on soft, tropical plants to adapt to browsing of drier plant material, and grazing of tougher plains grasses. Thus the proto-horses changed from leaf-eating forest-dwellers to grass-eating inhabitants of semi-arid regions worldwide, including the steppes of Eurasia and the Great Plains of North America. For reasons not fully understood, "Equus caballus" disappeared from North America around 10,000 years ago, at the end of the last ice age. The "Four Foundations" theory. Modern DNA evidence suggests that domesticated horses evolved from multiple wild populations. Specifically, the "Four Foundations" theory suggests that the modern horse evolved from multiple ancient wild prototypes, each adapted to a given habitat. However, an older theory holds that there was only one type of wild horse, the Tarpan subtype, and all other types diverged in form after domestication to meet human needs. Under the four foundations theory, all types and breeds of horses are thought to have developed from the following base prototypes: Domestication and surviving wild species. Competing theories exist as to the time and place of initial domestication. The earliest evidence for the domestication of the horse comes from Ukraine and dates to approximately 4,000 BC. It is thought that the horse was completely domesticated by 3000 BC, and by 2000 BC there was	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.