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top 10 words in brain distribution (in article): animal species breed male hunt human female size population bear |
top 10 words in brain distribution (in article): design wood type plant size key form fruit common allow |
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top 10 words in brain distribution (not in article): cat wolf dog wild lion iron blade elephant steel tiger |
top 10 words in brain distribution (not in article): material tree build lock frame paint machine switch wall card |
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times more probable under ant 30 20 10 6 4 2.5 1.25 1 1.25 2.5 4 6 10 20 30 times more probable under corn (words not in the model) | |
| A phylogeny of the extant ant subfamilies. "'Ants'" are social insects of the family "'Formicidae'", and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,000 species are classified with upper estimates of about 14,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist. Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because ants appear to operate as a unified entity, collectively working together to support the colony. Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15–25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study. Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have establish themselves in new areas where they may be accidentally introduced. Taxonomy and evolution. The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago. In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant ("Sphecomyrma freyi") that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. "Sphecomyrma" was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface. During the Cretaceous period, only a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20–40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Termites, though sometimes called "white ants", are not ants and belong to the order Isoptera. The termites are actually more closely related to cockroaches and mantids. The fact that ants and termites are both eusocial came about by Convergent evolution. Velvet ants look like large ants, but are wingless female wasps. Etymology. The word "ant" is derived from "ante" of Middle English which is derived from "æmette" and "emmett" of Old English and is related to the Old High German "āmeiza" from which comes "Ameise", the German word for ant. The family name "Formicidae" is derived from the Latin "formīca" ("ant") from which derived Portuguese "formiga", Spanish "hormiga", Romanian "furnică", French "fourmi", etc. Distribution and diversity. Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15–20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates. Ants range in size from. Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies. Morphology. Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarisation. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae. All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species. The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests. Polymorphism. In the colonies of a few ant species, there are physical castes—workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution. Some other species show continuous variation in the size of workers. The smallest and largest workers in "Pheidologeton diversus" show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called "repletes". This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in "Acromyrmex" sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of "Solenopsis invicta" can decide whether the colony will have single or multiple queens. The Australian jack jumper ant ("Myrmecia pilosula"), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects. Development and reproduction. The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes. Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. During the short breeding period, the reproductives, excluding the colony queen, are carried outside where other colonies of similar species are doing the same. Then, all the winged breeding ants take flight. Mating occurs in flight and the males die shortly afterwards. Females of some species mate with multiple males. Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, a process akin to swarming in honeybees. Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size. Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity. Communication. Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path. Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group ("e.g.", foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species. Defence===. Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants ("Paraponera"), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, "Solenopsis" spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people. Trap-jaw ants of the genus "Odontomachus" are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of "Odontomachus bauri" recorded peak speeds of between 126 and 230 h (78 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: "Anochetus", "Orectognathus", and "Strumigenys", plus some members of the Dacetini tribe, which are viewed as examples of convergent evolution. In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or "necrophory", the disposal of dead nest-mates. Oleic acid has been identified as the compound released by dead ants that triggers undertaking behaviour in "Atta mexicana". Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of "Cataulacus muticus", an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Learning. Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of "Temnothorax albipennis" leads a naive nest-mate to newly discovered food by the excruciatingly slow process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close. Controlled experiments with colonies of "Cerapachys biroi" suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care. Nest construction. Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; "Temnothorax albipennis" will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats. The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Weaver ant ("Oecophylla" spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of "Polyrhachis". Food cultivation. Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants ("Atta" and "Acromyrmex") feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called "gongylidia". Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi. Navigation. Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants ("Cataglyphis fortis") use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant ("Cataglyphis bicolor") navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. Locomotion. Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant ("Harpegnathos saltator") is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including "Cephalotes atratus"; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling. Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. "Polyrhachis sokolova", a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests. Cooperation and competition. Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey. Some species (such as "Tetramorium caespitum") attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked. Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like "Strumigenys xenos" are entirely parasitic and do not have workers, but instead rely on the food gathered by their "Strumigenys perplexa" hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in "Wasmannia auropunctata", where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father. Relationships with other organisms. The spider "Myrmarachne plataleoides" (here a female) mimics weaver ants to avoid predators. Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines. Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids | Maize'" ("Zea mays" L. ssp. "mays"), known as corn'" in some countries, is a cereal grain domesticated in Mesoamerica and subsequently spread throughout the American continents. After European contact with the Americas in the late 15th and early 16th century, maize spread to the rest of the world. Maize is the most widely grown crop in the Americas (332 million tonnes annually in the United States alone). Hybrid maize, due to its high grain yield as a result of heterosis ("hybrid vigor"), is preferred by farmers over conventional varieties. While some maize varieties grow up to 7 metres (23 ft) tall, most commercially grown maize has been bred for a standardized height of 2.5 metres (8 ft). Sweet corn is usually shorter than field-corn varieties. Naming conventions. The term "maize" derives from the Spanish form ("maíz") of the indigenous Taino term for the plant, and was the form most commonly heard in the United Kingdom. In the United States, Canada (maïs in French speaking Canadian regions) and Australia, the usual term is "corn", which originally referred to any grain, but which now refers exclusively to maize, having been shortened from the form "Indian corn" (which currently, at least in the U.S. & Canada, is often used to refer specifically to multi-colored "field corn" cultivars). Physiology. Maize stems superficially resemble bamboo canes and the internodes can reach 20–30 centimetres (8–12 in). Maize has a very distinct growth form; the lower leaves being like broad flags, 50–100 centimetres long and 5–10 centimetres wide (2–4 ft by 2–4 in); the stems are erect, conventionally 2–3 metres (7–10 ft) in height, with many nodes, casting off flag-leaves at every node. Under these leaves and close to the stem grow the ears. They grow about 3 milimetres a day. The ears are female inflorescences, tightly covered over by several layers of leaves, and so closed-in by them to the stem that they do not show themselves easily until the emergence of the pale yellow silks from the leaf whorl at the end of the ear. The silks are elongated stigmas that look like tufts of hair, at first green, and later red or yellow. Plantings for silage are even denser, and achieve an even lower percentage of ears and more plant matter. Certain varieties of maize have been bred to produce many additional developed ears, and these are the source of the "baby corn" that is used as a vegetable in Asian cuisine. Maize is a facultative long-night plant and flowers in a certain number of growing degree days >50 °F (10 °C) in the environment to which it is adapted. The magnitude of the influence that long nights have on the number of days that must pass before maize flowers is genetically prescribed and regulated by the phytochrome system. Photoperiodicity can be eccentric in tropical cultivars, while the long days characteristic of higher latitudes allow the plants to grow so tall that they do not have enough time to produce seed before being killed by frost. These attributes, however, may prove useful in using tropical maize for biofuels. The apex of the stem ends in the tassel, an inflorescence of male flowers. Each silk may become pollinated to produce one kernel of corn. Young ears can be consumed raw, with the cob and silk, but as the plant matures (usually during the summer months) the cob becomes tougher and the silk dries to inedibility. By the end of the growing season, the kernels dry out and become difficult to chew without cooking them tender first in boiling water. Modern farming techniques in developed countries usually rely on dense planting, which produces on average only about 0.9 ears per stalk because it stresses the plants. The kernel of corn has a pericarp of the fruit fused with the seed coat, typical of the grasses. It is close to a multiple fruit in structure, except that the individual fruits (the kernels) never fuse into a single mass. The grains are about the size of peas, and adhere in regular rows round a white pithy substance, which forms the ear. An ear contains from 200 to 400 kernels, and is from 10–25 centimetres (4–10 inches) in length. They are of various colors: blackish, bluish-gray, red, white and yellow. When ground into flour, maize yields more flour, with much less bran, than wheat does. However, it lacks the protein gluten of wheat and, therefore, makes baked goods with poor rising capability and coherence. A genetic variation that accumulates more sugar and less starch in the ear is consumed as a vegetable and is called sweet corn. Immature maize shoots accumulate a powerful antibiotic substance, DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one). DIMBOA is a member of a group of hydroxamic acids (also known as benzoxazinoids) that serve as a natural defense against a wide range of pests including insects, pathogenic fungi and bacteria. DIMBOA is also found in related grasses, particularly wheat. A maize mutant (bx) lacking DIMBOA is highly susceptible to be attacked by aphids and fungi. DIMBOA is also responsible for the relative resistance of immature maize to the European corn borer (family Crambidae). As maize matures, DIMBOA levels and resistance to the corn borer decline. Due to its shallow roots of only one to two inches deep, maize is susceptible to droughts, intolerant of nutrient-deficient soils, and prone to be uprooted by severe winds. Allergy. Maize contains lipid transfer protein, an undigestable protein which survives cooking. This protein has been linked to a rare and understudied allergy to maize in humans. The allergic reaction can cause skin rash, swelling or itching of mucus membranes, diarrhoea, vomiting, asthma and, in severe cases, anaphylactic shock. It has been noted that those with corn allergy almost always have peach allergy as well. It is unclear how common this allergy is in the general populace. Genetics. Many forms of maize are used for food, sometimes classified as various subspecies: This system has been replaced (though not entirely displaced) over the last 60 years by multi-variable classifications based on ever more data. Agronomic data were supplemented by botanical traits for a robust initial classification, then genetic, cytological, protein and DNA evidence was added. Now the categories are forms (little used), races, racial complexes, and recently branches. Maize has 10 chromosomes (n=10). The combined length of the chromosomes is 1500 cM. Some of the maize chromosomes have what are known as "chromosomal knobs": highly repetitive heterochromatic domains that stain darkly. Individual knobs are polymorphic among strains of both maize and teosinte. Barbara McClintock used these knob markers to prove her transposon theory of "jumping genes", for which she won the 1983 Nobel Prize in Physiology or Medicine. Maize is still an important model organism for genetics and developmental biology today. There is a stock center of maize mutants, "The Maize Genetics Cooperation Stock Center", funded by the USDA Agricultural Research Service and located in the Department of Crop Sciences at the University of Illinois at Urbana-Champaign. The total collection has nearly 80,000 samples. The bulk of the collection consists of several hundred named genes, plus additional gene combinations and other heritable variants. There are about 1000 chromosomal aberrations (e.g., translocations and inversions) and stocks with abnormal chromosome numbers (e.g., tetraploids). Genetic data describing the maize mutant stocks as well as myriad other data about maize genetics can be accessed at MaizeGDB, the Maize Genetics and Genomics Database. In 2005, the U.S. National Science Foundation (NSF), Department of Agriculture (USDA) and the Department of Energy (DOE) formed a consortium to sequence the maize genome. The resulting DNA sequence data will be deposited immediately into GenBank, a public repository for genome-sequence data. Sequencing the corn genome has been considered difficult because of its large size and complex genetic arrangements. The genome has 50,000–60,000 genes scattered among the 2.5 billion bases—molecules that form DNA—that make up its 10 chromosomes. (By comparison, the human genome contains about 2.9 billion bases and 26,000 genes.) On February 26, 2008, researchers announced that they had sequenced the entire genome of maize. Origin. There are several theories about the specific origin of maize in Mesoamerica: The first model was proposed by Nobel Prize winner George Beadle in 1939. Though it has experimental support, it has not explained a number of problems, among them: The domestication of maize is of particular interest to researchers — archaeologists, geneticists, ethnobotanists, geographers, etc. The process is thought by some to have started 7,500 to 12,000 years ago (corrected for solar variations). Recent genetic evidence suggests that maize domestication occurred 9,000 years ago in central Mexico, perhaps in the highlands between Oaxaca and Jalisco. The crop wild relative teosinte most similar to modern maize grows in the area of the Balsas River. Archaeological remains of early maize ears, found at Guila Naquitz Cave in the Oaxaca Valley, date back roughly 6,250 years (corrected; 3450 BC, uncorrected); the oldest ears from caves near Tehuacan, Puebla, date ca. 2750 BC. Little change occurred in ear form until ca. 1100 BC when great changes appeared in ears from Mexican caves: maize diversity rapidly increased and archaeological teosinte was first deposited. Perhaps as early as 1500 BC, maize began to spread widely and rapidly. As it was introduced to new cultures, new uses were developed and new varieties selected to better serve in those preparations. Maize was the staple food, or a major staple, of most the pre-Columbian North American, Mesoamerican, South American, and Caribbean cultures. The Mesoamerican civilization was strengthened upon the field crop of maize; through harvesting it, its religious and spiritual importance and how it impacted their diet. Maize formed the Mesoamerican people’s identity. During the 1st millennium AD, maize cultivation spread from Mexico into the U.S. Southwest and a millennium later into U.S. Northeast and southeastern Canada, transforming the landscape as Native Americans cleared large forest and grassland areas for the new crop. It is unknown what precipitated its domestication, because the edible portion of the wild variety is too small and hard to obtain to be eaten directly, as each kernel is enclosed in a very hard bi-valve shell. However, George Beadle demonstrated that the kernels of teosinte are readily "popped" for human consumption, like modern popcorn. Some have argued that it would have taken too many generations of selective breeding in order to produce large compressed ears for efficient cultivation. However, studies of the hybrids readily made by intercrossing teosinte and modern maize suggest that this objection is not well founded. In 2005, research by the USDA Forest Service indicated that the rise in maize cultivation 500 to 1,000 years ago in what is now the southeastern United States contributed to the decline of freshwater mussels, which are very sensitive to environmental changes. Production quantities and methods. Maize is widely cultivated throughout the world, and a greater weight of maize is produced each year than any other grain. While the United States produces almost half of the world's harvest(~42.5%), other top producing countries include China, Brazil, Mexico, Argentina, India and France. Worldwide production was around 800 million tonnes in 2007—just slightly more than rice (~650 million tonnes) or wheat (~600 million tonnes). In 2007, over 150 million hectares of maize were planted worldwide, with a yield of 4970.9 kilogram hectare. Because it is cold-intolerant, in the temperate zones maize must be planted in the spring. Its root system is generally shallow, so the plant is dependent on soil moisture. As a C4 plant (a plant that uses C4 carbon fixation), maize is a considerably more water-efficient crop than C3 plants (plants that use C3 carbon fixation) like the small grains, alfalfa and soybeans. Maize is most sensitive to drought at the time of silk emergence, when the flowers are ready for pollination. In the United States, a good harvest was traditionally predicted if the corn was "knee-high by the Fourth of July," although modern hybrids generally exceed this growth rate. Maize used for silage is harvested while the plant is green and the fruit immature. Sweet corn is harvested in the "milk stage," after pollination but before starch has formed, between late summer and early to mid-autumn. Field corn is left in the field very late in the autumn in order to thoroughly dry the grain, and may, in fact, sometimes not be harvested until winter or even early spring. The importance of sufficient soil moisture is shown in many parts of Africa, where periodic drought regularly causes famine by causing maize crop failure. Maize was planted by the Native Americans in hills, in a complex system known to some as the Three Sisters: beans used the corn plant for support and in turn provided nitrogen from nitrogen-fixing bacteria which live on the roots of beans and other legumes; and squashes provided ground cover to stop weeds and inhibit evaporation by providing shade over the soil. This method was replaced by single species hill planting where each hill 60–120 cm (2–4 ft) apart was planted with 3 or 4 seeds, a method still used by home gardeners. A later technique was "checked corn" where hills were placed 40 inches apart in each direction, allowing cultivators to run through the field in two directions. In more arid lands this was altered and seeds were planted in the bottom of 10–12 cm (4–5 in) deep furrows to collect water. Modern technique plants maize in rows which allows for cultivation while the plant is young, although the hill technique is still used in the cornfields of some Native American reservations. In North America, fields are often planted in a two-crop rotation with a nitrogen-fixing crop, often alfalfa in cooler climates and soybeans in regions with longer summers. Sometimes a third crop, winter wheat, is added to the rotation. Fields are usually ploughed each year, although no-till farming is increasing in use. Many of the maize varieties grown in the United States and Canada are hybrids. Over half of the corn area planted in the United States has been genetically modified using biotechnology to express agronomic traits such as pest resistance or herbicide resistance. Before about World War II, most maize in North America was harvested by hand (as it still is in most of the other countries where it is grown). This often involved large numbers of workers and associated social events. Some one- and two-row mechanical pickers were in use but the corn combine was not adopted until after the War. By hand or mechanical picker, the entire ear is harvested which then requires a separate operation of a corn sheller to remove the kernels from the ear. Whole ears of corn were often stored in "corn cribs" and these whole ears are a sufficient form for some livestock feeding use. Few modern farms store maize in this manner. Most harvest the grain from the field and store it in bins. The combine with a corn head (with points and snap rolls instead of a reel) does not cut the stalk; it simply pulls the stalk down. The stalk continues downward and is crumpled in to a mangled pile on the ground. The ear of corn is too large to pass through a slit in a plate and the snap rolls pull the ear of corn from the stalk so that only the ear and husk enter the machinery. The combine separates out the husk and the cob, keeping only the kernels. Pellagra. When maize was first introduced into other farming systems than those used by traditional native-American peoples, it was generally welcomed with enthusiasm for its productivity. However, a widespread problem of malnutrition soon arose wherever maize was introduced as a staple. This was a mystery since these types of malnutrition were not normally seen among the indigenous Americans, to whom Maize was the principal staple food. It was eventually discovered that the indigenous Americans learned long ago to add alkali—in the form of ashes among North Americans and lime (calcium carbonate) among Mesoamericans—to corn meal, which liberates the B-vitamin niacin, the lack of which was the underlying cause of the condition known as pellagra. This alkali process is known by its Nahuatl (Aztec)-derived name: nixtamalization. Besides the lack of niacin, pellagra was also characterized by protein deficiency, a result of the inherent lack of two key amino acids in pre-modern maize, lysine and tryptophan. Nixtamalisation was also found to increase the lysine and tryptophan content of maize to some extent, but more importantly, the indigenous Americans had learned long ago to balance their consumption of maize with beans and other protein sources such as amaranth and chia, as well as meat and fish, in order to acquire the complete range of amino acids for normal protein synthesis. Since maize had been introduced into the diet of non-indigenous Americans without the necessary cultural knowledge acquired over thousands of years in the Americas, the reliance on maize in other cultures was often tragic. In the late 19th century pellagra reached endemic proportions in parts of the deep southern U.S., as medical researchers debated two theories for its origin: the deficiency theory (eventually shown to be true) posited that pellagra was due to a deficiency of some nutrient, and the germ theory posited that pellagra was caused by a germ transmitted by stable flies. In 1914 the U.S. government officially endorsed the germ theory of pellagra, but rescinded this endorsement several years later as evidence grew against it. By the mid-1920s the deficiency theory of pellagra was becoming scientific consensus, and the theory was proved in 1932 when niacin deficiency was determined to be the cause of the illness. Once alkali processing and dietary variety was understood and applied, pellagra disappeared. The development of high lysine maize and the promotion of a more balanced diet has also contributed to its demise. Insect pests. The susceptibility of maize to the European corn borer, and the resulting large crop losses, led to the development of transgenic expressing the "Bacillus thuringiensis" toxin. "Bt corn" is widely grown in the United States and has been approved for release in Europe. Food. Corn and cornmeal (corn flour) constitutes a staple food in many regions of the world. Corn meal is made into a thick porridge in many cultures: from the polenta of Italy, the angu of Brazil, the mămăligă of Romania, to mush in the U.S. or the food called sadza, nshima, ugali, tuwan-masara and mealie pap in Africa. Corn meal is also used as a replacement for wheat flour, to make cornbread and other baked products. Masa (cornmeal treated with lime water) is the main ingredient for tortillas, atole and many other dishes of Mexican food. Popcorn is kernels of certain varieties that explode when heated, forming fluffy pieces that are eaten as a snack. Chicha and "chicha morada"(purple chicha) are drinks made usually from particular types of maize. The first one is fermented and alcoholic, the second one is a soft drink commonly drunk in Peru Corn flakes are a common breakfast staple in the United States, and are increasingly popular all over the world. Maize can also be prepared as hominy, in which the kernels are soaked with lye; or grits, which are coarsely ground hominy. These are commonly eaten in the Southeastern United States, foods handed down from Native Americans. The Brazilian dessert canjica is made by boiling maize kernels in sweetened milk. Roasted dried corn cobs with semi-hardened kernels, coated with a seasoning mixture of fried chopped spring onions with salt added to the oil, is a popular snack food in Vietnam. Maize can also be harvested and consumed in the unripe state, when the kernels are fully grown but still soft. Unripe corn must usually be cooked to become palatable; this may be done by simply boiling or roasting the whole ears and eating the kernels right off the cob. Such corn on the cob is a common dish in the United States, United Kingdom and some parts of South America, but virtually unheard of in some European countries. The cooked unripe kernels may also be shaved off the cob and served as a vegetable in side dishes, salads, garnishes, etc. Alternatively, the raw unripe kernels may also be grated off the cobs and processed into a variety of cooked dishes, such as corn purée, tamales, pamonhas, curau, cakes, ice creams, etc. Sweetcorn, a genetic variety that is high in sugars and low in starch, is usually consumed in the unripe state. Maize is a major source of starch, a major ingredient in home cooking and in many industrialized food products. It is also a major source of cooking oil (corn oil) and of corn gluten. Maize starch can be hydrolyzed and enzymatically treated to produce syrups, particularly high fructose corn syrup, a sweetener; and also fermented and distilled to produce grain alcohol. Grain alcohol from maize is traditionally the source of bourbon whiskey. Maize is used to make chicha, a fermented beverage of Central and South America; and sometimes as the starch source for beer. In the United States and Canada maize is also widely grown to feed for livestock, as forage, silage (made by fermentation of chopped green cornstalks), or grain. Corn meal is also a significant ingredient of some commercial animal food products, such as dog food. Maize is also used as a fish bait, called "dough balls". It is particularly popular in Europe for coarse fishing. Chemicals and medicines. Starch from maize can also be made into plastics, fabrics, adhesives, and many other chemical products. Stigmas from female corn flowers, known popularly as corn silk, are sold as herbal supplements. The corn steep liquor, a plentiful watery byproduct of maize wet milling process, is widely used in the biochemical industry and research as a culture medium to grow many kinds of microorganisms. Biofuel. "Feed corn" is being used increasingly for heating; specialized corn stoves (similar to wood stoves) are available and use either feed corn or wood pellets to generate heat. Corncobs are also used as a biomass fuel source. Maize is relatively cheap and home-heating furnaces have been developed which use maize kernels as a fuel. They feature a large hopper that feeds the uniformly sized corn kernels (or wood pellets or cherry pits) into the fire. Maize is increasingly used as a biomass fuel, such as ethanol, which as researchers search for innovative ways to reduce fuel costs, has unintentionally caused a rapid rise in food costs. This has led to the 2007 harvest being one of the most profitable corn crops in modern history for farmers. Maize is widely used in Germany as a feedstock for biogas plants. Here the maize is harvested, shredded then placed in silage clamps from which it is fed into the biogas plants. A biomass gasification power plant in Strem near Güssing, Burgenland, Austria was begun in 2005. Research is being done to make diesel out of the biogas by the Fischer Tropsch method. Increasingly ethanol is being used at low concentrations (10% or less) as an additive in gasoline (gasohol) for motor fuels to increase the octane rating, lower pollutants, and reduce petroleum use (what is nowadays also known as "biofuels" and has been generating an intense debate regarding the human beings' necessity of new sources of energy, on the one hand, and the need to maintain, in regions such as Latin America, the food habits and culture which has been the essence of civilizations such as the one originated in Mesoamerica; the entry, January 2008, of maize among the commercial agreements of NAFTA has increased this debate, considering the bad labor conditions of workers in the fields, and mainly the fact that NAFTA "opened the doors to the import of corn from the United States, where the farmers who grow it receive multi-million dollar subsidies and other government supports. According to OXFAM UK, after NAFTA went into effect, the price of maize in Mexico fell 70% between 1994 and 2001. The number of farm jobs dropped as well: from 8.1 million in 1993 to 6.8 million in 2002. Many of those who found themselves without work were small-scale maize growers."). However, introduction in the northern latitudes of the U.S. of, and not for human or animal consumption, may potentially alleviate this. As a result of the U.S. federal government announcing its production target of 35 billion gallons of biofuels by 2017, ethanol production will grow to 7 billion gallons by 2010, up from 4.5 billion in 2006, boosting ethanol's share of corn demand in the U.S. from 22.6 percent to 36.1 percent. Ornamental and other uses. Some forms of the plant are occasionally grown for ornamental use in the garden. For this purpose, variegated and colored leaf forms as well as those with colorful ears are used. Additionally, size-superlative varieties, having reached 31 ft (9.4m) tall, or with ears 24 inches (60 cm) long, have been popular for at least a century. Corncobs can be hollowed out and treated to make inexpensive smoking pipes, first manufactured in the United States in 1869. An unusual use for maize is to create a "maize maze" as a tourist attraction. This is a maze cut into a field of maize. The idea of a maize maze was introduced by Adrian Fisher, one of the most prolific designers of modern mazes, with The American Maze Company who created a maze in Pennsylvania in 1993. Traditional mazes are most commonly grown using yew hedges, but these take several years to mature. The rapid growth of a field of maize allows a maze to be laid out using GPS at the start of a growing season and for the maize to grow tall enough to obstruct a visitor's line of sight by the start of the summer. In Canada and the U.S., these are called "corn mazes" and are popular in many farming communities. Corn kernels can be used in place of sand in a sandbox-like enclosure for children's play. Fodder. Maize makes a greater quantity of epigeous mass than other cereal plants, so can be used for fodder. Digestibility and palatability are higher when ensiled and fermented, rather than dried. In art. Maize has been an essential crop in the Andes since the pre-Columbian Era. The Moche culture from Northern Peru made ceramics from earth, water, and fire. This pottery was a sacred substance, formed in significant shapes and used to represent important themes. Maize represented anthropomorphically as well as naturally. In the United States, maize itself is sometimes used for temporary architectural detailing when the intent is to celebrate local agricultural productivity and culture. A well-known example of this use is the Corn Palace in Mitchell, South Dakota, which utilizes cobs of colored maize to implement a design that is recycled annually. External links. 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