Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (
North American
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Ca ...
and
Australian English), is a
cereal grain
A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
first domesticated by
indigenous peoples
Indigenous peoples are culturally distinct ethnic groups whose members are directly descended from the earliest known inhabitants of a particular geographic region and, to some extent, maintain the language and culture of those original people ...
in southern Mexico about 10,000 years ago.
The leafy stalk of the plant produces
pollen
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...
s (or "tassels") and separate
ovuliferous inflorescences called
ears that when fertilized yield
kernels or seeds, which are
fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
s. The term ''maize'' is preferred in formal, scientific, and international usage as a
common name
In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contra ...
because it refers specifically to this one grain, unlike ''corn'', which has a complex variety of meanings that vary by context and geographic region.
Maize has become a
staple food
A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard Diet (nutrition), diet for a given person or group of people, supplying a large fraction of ...
in many parts of the world, with the
total production of maize surpassing that of
wheat
Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeolog ...
or
rice
Rice is the seed of the grass species '' Oryza sativa'' (Asian rice) or less commonly '' Oryza glaberrima'' (African rice). The name wild rice is usually used for species of the genera '' Zizania'' and ''Porteresia'', both wild and domestica ...
. In addition to being consumed directly by humans (often in the form of
masa
''Masa'' (or ''masa de maíz'') (; ) is a maize dough that comes from ground nixtamalized corn. It is used for making corn tortillas, ''gorditas'', ''tamales'', '' pupusas'', and many other Latin American dishes. It is dried and powdered into a ...
), maize is also used for
corn ethanol
Corn ethanol is ethanol produced from corn biomass and is the main source of ethanol fuel in the United States, mandated to be blended with gasoline in the Renewable Fuel Standard. Corn ethanol is produced by ethanol fermentation and distillat ...
,
animal feed
Animal feed is food given to domestic animals, especially livestock, in the course of animal husbandry. There are two basic types: fodder and forage. Used alone, the word ''feed'' more often refers to fodder. Animal feed is an important input to ...
and other
maize products, such as
corn starch
Corn starch, maize starch, or cornflour (British English) is the starch derived from corn (maize) grain. The starch is obtained from the endosperm of the kernel. Corn starch is a common food ingredient, often used to thicken sauces or soup ...
and
corn syrup
Corn syrup is a food syrup which is made from the starch of corn (called maize in many countries) and contains varying amounts of sugars: glucose, maltose and higher oligosaccharides, depending on the grade. Corn syrup is used in foods to soften ...
. The six major types of maize are
dent corn
Dent corn, also known as grain corn, is a type of field corn with a high soft starch content. It received its name because of the small indentation, or "dent", at the crown of each kernel on a ripe ear of corn. Reid's Yellow Dent is a variety d ...
,
flint corn
Flint corn (''Zea mays'' var. ''indurata''; also known as Indian corn or sometimes calico corn) is a variant of maize, the same species as common corn. Because each kernel has a hard outer layer to protect the soft endosperm, it is likened to bei ...
,
pod corn
Pod corn or wild maize is a variety of maize. It is not a wild ancestor of maize but rather a mutant that forms leaves around each kernel.
Pod corn (''tunicata'' Sturt) is not grown commercially, but it is preserved in some localities.
Pod corn ...
,
popcorn
Popcorn (also called popped corn, popcorns or pop-corn) is a variety of corn kernel which expands and puffs up when heated; the same names also refer to the foodstuff produced by the expansion.
A popcorn kernel's strong hull contains the se ...
,
flour corn
Flour corn (''Zea mays'' var. ''amylacea'') is a variety of corn with a soft starchy endosperm and a thin pericarp. It is primarily used to make corn flour.
This type, frequently found in Aztec and Inca graves, is widely grown in the drier par ...
, and
sweet corn
Sweet corn (''Zea mays'' convar. ''saccharata'' var. ''rugosa''), also called sugar corn and pole corn, is a variety of maize grown for human consumption with a high sugar content. Sweet corn is the result of a naturally occurring recessive m ...
. Sugar-rich varieties called sweet corn are usually grown for human consumption as kernels, while
field corn Field corn, also known as cow corn, is a North American term for maize (''Zea mays'') grown for livestock fodder (silage), ethanol, cereal, and processed food products. The principal field corn varieties are dent corn, flint corn, flour corn (als ...
varieties are used for animal feed, various corn-based human food uses (including grinding into
cornmeal
Cornmeal is a meal (coarse flour) or a cell membrane ground from dried corn. It is a common staple food, and is ground to coarse, medium, and fine consistencies, but not as fine as wheat flour can be.Herbst, Sharon, ''Food Lover's Companion'', ...
or
masa
''Masa'' (or ''masa de maíz'') (; ) is a maize dough that comes from ground nixtamalized corn. It is used for making corn tortillas, ''gorditas'', ''tamales'', '' pupusas'', and many other Latin American dishes. It is dried and powdered into a ...
, pressing into
corn oil
Corn oil (North American) or maize oil (British) is oil extracted from the germ of corn (maize). Its main use is in cooking, where its high smoke point makes refined corn oil a valuable frying oil. It is also a key ingredient in some margarin ...
, fermentation and distillation into alcoholic beverages like
bourbon whiskey
Bourbon () is a type of barrel-aged American whiskey made primarily from corn. The name derives from the French Bourbon dynasty, although the precise source of inspiration is uncertain; contenders include Bourbon County in Kentucky and Bourbon ...
), and as feedstocks for the chemical industry. Maize is also used in making
ethanol
Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a h ...
and other
biofuels.
Maize is widely cultivated throughout the world, and a greater weight of maize is produced each year than any other grain.
[ In 2021, total world production was 1.2 billion ]tonne
The tonne ( or ; symbol: t) is a unit of mass equal to 1000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton ( United State ...
s. Maize is the most widely grown grain crop
A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydropo ...
throughout the Americas, with 384 million metric tons
The tonne ( or ; symbol: t) is a unit of mass equal to 1000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton (United States c ...
grown in the United States alone in 2021. Genetically modified maize
Genetically modified maize (corn) is a genetically modified crop. Specific maize strains have been genetically engineered to express agriculturally-desirable traits, including resistance to pests and to herbicides. Maize strains with both tra ...
made up 85% of the maize planted in the United States in 2009.[Genetically modified plants: Global Cultivation Area Maize]
GMO Compass, March 29, 2010, retrieved August 10, 2010 Subsidies in the United States help to account for its high level of cultivation of maize and its position as the largest producer in the world.
History
Pre-Columbian development
Maize is a cultigen
A cultigen () or cultivated plant is a plant that has been deliberately altered or selected by humans; it is the result of artificial selection. These plants, for the most part, have commercial value in horticulture, agriculture or forestry. B ...
; human intervention is required for it to propagate. Whether or not the kernels fall off the cob on their own is a key piece of evidence used in archaeology to distinguish domesticated maize from its naturally-propagating teosinte
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild sp ...
ancestor. Genetic evidence can also be used to determine when various lineages split.
Most historians believe maize was domesticated in the Tehuacán Valley "By faith and hope"
,
, image_map =
, mapsize = 300 px
, map_caption = Location of Tehuacán within the state of Puebla.
, image_map1 = Puebla en México.svg
, mapsize1 = 300 px
, ma ...
of Mexico. Recent research in the early 21st century has modified this view somewhat; scholars now indicate the adjacent Balsas River
The Balsas River (Spanish Río Balsas, also locally known as the Mezcala River, or Atoyac River) is a major river of south-central Mexico.
The basin flows through the states of Guerrero, México, Morelos, and Puebla. Downstream of Ciudad Al ...
Valley of south-central Mexico as the center of domestication.
An 2002 study by Matsuoka ''et al.''. has demonstrated that, rather than the multiple independent domestications model, all maize arose from a single domestication in southern Mexico about 9,000 years ago. The study also demonstrated that the oldest surviving maize types are those of the Mexican highlands. Later, maize spread from this region over the Americas along two major paths. This is consistent with a model based on the archaeological record suggesting that maize diversified in the highlands of Mexico before spreading to the lowlands.
Archaeologist Dolores Piperno has said:
Since then, even earlier dates have been published.
According to a genetic study by the Brazilian Agricultural Research Corporation
The Brazilian Agricultural Research Corporation (Embrapa - pt, Empresa Brasileira de Pesquisa Agropecuária) is a state-owned research corporation affiliated with the Brazilian Ministry of Agriculture. Since its inception on April 26, 1973, it ha ...
(Embrapa), corn cultivation was introduced in South America from Mexico, in two great waves: the first, more than 6000 years ago, spread through the Andes
The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
. Evidence of cultivation in Peru has been found dating to about 6700 years ago. The second wave, about 2000 years ago, through the lowlands of South America.
The earliest maize plants grew only small, corn ears, and only one per plant. In Jackson Spielvogel's view, many centuries of artificial selection (rather than the current view that maize was exploited by interplanting with ''teosinte
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild sp ...
'') by the indigenous people of the Americas resulted in the development of maize plants capable of growing several ears per plant, which were usually several centimetres/inches long each. The Olmec
The Olmecs () were the earliest known major Mesoamerican civilization. Following a progressive development in Soconusco, they occupied the tropical lowlands of the modern-day Mexican states of Veracruz and Tabasco. It has been speculated that ...
and Maya
Maya may refer to:
Civilizations
* Maya peoples, of southern Mexico and northern Central America
** Maya civilization, the historical civilization of the Maya peoples
** Maya language, the languages of the Maya peoples
* Maya (Ethiopia), a popu ...
cultivated maize in numerous varieties throughout Mesoamerica
Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
; they cooked, ground and processed it through nixtamalization
Nixtamalization () is a process for the preparation of corn, or other grain, in which the grain is soaked and cooked in an alkaline solution, usually limewater (but sometimes aqueous alkali metal carbonates), washed, and then hulled. The t ...
. It was believed that beginning about 2500 BC, the crop spread through much of the Americas. Research of the 21st century has established even earlier dates. The region developed a trade network based on surplus and varieties of maize crops.
Mapuche
The Mapuche ( (Mapuche & Spanish: )) are a group of indigenous inhabitants of south-central Chile and southwestern Argentina, including parts of Patagonia. The collective term refers to a wide-ranging ethnicity composed of various groups who s ...
s of south-central Chile cultivated maize along with quinoa
Quinoa (''Chenopodium quinoa''; , from Quechua ' or ') is a flowering plant in the amaranth family. It is a herbaceous annual plant grown as a crop primarily for its edible seeds; the seeds are rich in protein, dietary fiber, B vitamins, a ...
and potatoes
The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae.
Wild potato species can be found from the southern United ...
in pre-Hispanic
In the history of the Americas, the pre-Columbian era spans from the original settlement of North and South America in the Upper Paleolithic period through European colonization, which began with Christopher Columbus's voyage of 1492. Usually, t ...
times; however, potato was the staple food
A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard Diet (nutrition), diet for a given person or group of people, supplying a large fraction of ...
of most Mapuches, "specially in the southern and coastal apucheterritories where maize did not reach maturity". Before the expansion of the Inca Empire
The Inca Empire (also known as the Incan Empire and the Inka Empire), called ''Tawantinsuyu'' by its subjects, ( Quechua for the "Realm of the Four Parts", "four parts together" ) was the largest empire in pre-Columbian America. The adm ...
maize was traded and transported as far south as 40°19' S in Melinquina, Lácar Department
Lácar is a department located in the south of Neuquén Province, Argentina.
Geography
The Department limits with Huiliches Department at North, Collón Cura Department at northeast, Rio Negro Province at southeast, Los Lagos Department a ...
. In that location maize remains were found inside pottery dated to 730 ± 80 BP and 920 ± 60 BP. Probably this maize was brought across the Andes from Chile.[ The presence of maize in ]Guaitecas Archipelago
Guaitecas Archipelago is a sparsely populated archipelago in the Aisén region of Chile. The archipelago is made up of eight main islands and numerous smaller ones. The eight largest islands are from northwest to southeast: Gran Guaiteca, Ascen ...
(43°55' S), the southernmost outpost of pre-Hispanic agriculture, is reported by early Spanish explorers.[ However the Spanish may have misidentified the plant.]
By at least 1000 BCE, the Olmec
The Olmecs () were the earliest known major Mesoamerican civilization. Following a progressive development in Soconusco, they occupied the tropical lowlands of the modern-day Mexican states of Veracruz and Tabasco. It has been speculated that ...
in Mesoamerica
Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
had based their calendar, language, myths and worldview with maize at the center of their symbolism.
Columbian exchange
After the arrival of Europeans in 1492, Spanish settlers consumed maize, and explorers and traders carried it back to Europe and introduced it to other countries. Spanish settlers much preferred wheat
Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeolog ...
bread to maize, cassava
''Manihot esculenta'', commonly called cassava (), manioc, or yuca (among numerous regional names), is a woody shrub of the spurge family, Euphorbiaceae, native to South America. Although a perennial plant, cassava is extensively cultivated a ...
, or potatoes
The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae.
Wild potato species can be found from the southern United ...
. Maize flour could not be substituted for wheat for communion bread, since in Christian
Christians () are people who follow or adhere to Christianity, a monotheistic Abrahamic religion based on the life and teachings of Jesus Christ. The words '' Christ'' and ''Christian'' derive from the Koine Greek title ''Christós'' (Χρ ...
belief only wheat could undergo transubstantiation
Transubstantiation (Latin: ''transubstantiatio''; Greek: μετουσίωσις '' metousiosis'') is, according to the teaching of the Catholic Church, "the change of the whole substance of bread into the substance of the Body of Christ and of ...
and be transformed into the body of Christ. Some Spaniards worried that by eating indigenous foods, which they did not consider nutritious, they would weaken and risk turning into Indians. "In the view of Europeans, it was the food they ate, even more than the environment in which they lived, that gave Amerindians and Spaniards both their distinctive physical characteristics and their characteristic personalities." Despite these worries, Spaniards did consume maize. Archeological evidence
Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landsca ...
from Florida sites indicate they cultivated it as well.[Earle, ''The Body of the Conquistador'', p. 144.]
Maize spread to the rest of the world because of its ability to grow in diverse climates. It was cultivated in Spain just a few decades after Columbus's voyages and then spread to Italy, West Africa and elsewhere.
Widespread cultivation most likely began in southern Spain in 1525, after which it quickly spread to the rest of the Spanish Empire
The Spanish Empire ( es, link=no, Imperio español), also known as the Hispanic Monarchy ( es, link=no, Monarquía Hispánica) or the Catholic Monarchy ( es, link=no, Monarquía Católica) was a colonial empire governed by Spain and its prede ...
including its territories in Italy (and, from there, to other Italian states). Maize had many advantages over wheat and barley; it yielded two and a half times the food energy
Food energy is chemical energy that animals (including humans) derive from their food to sustain their metabolism, including their muscular activity.
Most animals derive most of their energy from aerobic respiration, namely combining the carbohy ...
per unit cultivated area, could be harvested in successive years from the same plot of land, and grew in wildly varying altitudes and climates, from relatively dry regions with only of annual rainfall to damp regions with over . By the 17th century it was a common peasant food in Southwestern Europe, including Portugal, Spain, southern France, and Italy. By the 18th century, it was the chief food of the southern French and Italian peasantry, especially in the form of polenta
Polenta (, ) is a dish of boiled cornmeal that was historically made from other grains. The dish comes from Italy. It may be served as a hot porridge, or it may be allowed to cool and solidify into a loaf that can be baked, fried, or grilled. ...
in Italy.
Names
The word ''maize'' derives from the Spanish form of the indigenous Taíno
The Taíno were a historic indigenous people of the Caribbean whose culture has been continued today by Taíno descendant communities and Taíno revivalist communities. At the time of European contact in the late 15th century, they were the pri ...
word for the plant, ''mahiz''. Using the maize common name, Linnaeus included it as the species epithet in ''Zea mays''. It is known by other names including "corn" in some English speaking countries.
''Maize'' is preferred in formal, scientific, and international usage as a common name
In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contra ...
because it refers specifically to this one grain, unlike ''corn'', which has a complex variety of meanings that vary by context and geographic region. International groups such as the Centre for Agriculture and Bioscience International
CABI (legally CAB International, formerly Commonwealth Agricultural Bureaux) is a nonprofit intergovernmental development and information organisation focusing primarily on agricultural and environmental issues in the developing world, and the ...
also consider maize the preferred common name. According to Ohio State University, the US and a handful of other English-speaking countries primarily use corn, but the rest of the world calls this maize or maíz. The word maize is considered interchangeable with corn in the West; during early British and American trade, all grains were considered corn. Maize retained the name corn in the West as the primary grain in these trade relationships.
The word "corn" outside the US, Canada, Australia, and New Zealand is synonymous with grain
A grain is a small, hard, dry fruit ( caryopsis) – with or without an attached hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and legu ...
referring to any cereal
A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more foo ...
crop with its meaning understood to vary geographically to refer to the local staple
Staple may refer to:
*Staple food, a foodstuff that forms the basic constituent of a diet
*Staple (fastener), a small formed metal fastener
**Surgical staple
Arts, entertainment, and media
* Staple (band), a Christian post-hardcore band
** ''Stap ...
. In the United States, Canada, Australia, and New Zealand, ''corn'' primarily means maize; this usage started as a shortening of "Indian corn". "Indian corn" primarily means maize (the staple grain of indigenous Americans), but can refer more specifically to multicolored "flint corn
Flint corn (''Zea mays'' var. ''indurata''; also known as Indian corn or sometimes calico corn) is a variant of maize, the same species as common corn. Because each kernel has a hard outer layer to protect the soft endosperm, it is likened to bei ...
" used for decoration. Other common names include barajovar, makka, silk maize, and zea.
In a 1999 journal article, Betty Fussell describing calling maize as corn was "to plunge into tragi-farcial mistranslations of language and history." Similar to the British, the Spanish referred to maize as panizo, a generic term for cereal grains, as did Italians with the term polenta. The British later referred to maize as Turkey wheat, Turkey corn, or Indian corn with Fusell commenting that "they meant not a place but a condition, a savage rather than a civilized grain", especially with Turkish people
The Turkish people, or simply the Turks ( tr, Türkler), are the world's largest Turkic ethnic group; they speak various dialects of the Turkish language and form a majority in Turkey and Northern Cyprus. In addition, centuries-old ethnic Tu ...
later naming it kukuruz, or barbaric.
The word ''maize'' is used by agricultural bodies and research institutes such as the UN's FAO
The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
, the International Maize and Wheat Improvement Center
The International Maize and Wheat Improvement Center (known - even in English - by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de Maíz y Trigo'') is a non-profit research-for-development organization that develops im ...
based out of Mexico, and the Indian Institute of Maize Research. National agricultural and industry associations often include the word ''maize'' in their name such as the Maize Association of Australia, and the National Maize Association of Nigeria.
In Southern Africa, maize is commonly called ''mielie'' (Afrikaans
Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans g ...
) or ''mealie'' (English), words possibly derived from the Portuguese word for maize, ''milho'', but more probably from Dutch ''meel'' or English ''meal'', meaning the edible part of a grain or pulse
In medicine, a pulse represents the tactile arterial palpation of the cardiac cycle (heartbeat) by trained fingertips. The pulse may be palpated in any place that allows an artery to be compressed near the surface of the body, such as at the ...
.
Structure and physiology
The maize plant is often in height, though some natural strains can grow , and the tallest recorded plant reached almost . The stem is commonly composed of 20 internodes of length. The leaves arise from the nodes, alternately on opposite sides on the stalk, and have entire margins.
The apex of the stem ends in the tassel, an inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...
of male flowers. When the tassel is mature and conditions are suitably warm and dry, anthers on the tassel dehisce and release pollen. Maize pollen is anemophilous
Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous pla ...
(dispersed by wind), and because of its large settling velocity, most pollen falls within a few meters of the tassel.
Ears develop above a few of the leaves in the midsection of the plant, between the stem and leaf sheath, elongating by around per day, to a length of with being the maximum alleged in the subspecies. They are female inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...
s, tightly enveloped by several layers of ear leaves commonly called husks.
Elongated stigmas, called silks, emerge from the whorl of husk leaves at the end of the ear. They are often pale yellow and in length, like tufts of hair in appearance. At the end of each is a carpel, which may develop into a "kernel" if fertilized by a pollen grain. The pericarp
Fruit anatomy is the plant anatomy of the internal structure of fruit. Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits. Ag ...
of the fruit is fused with the seed coat referred to as "caryopsis
In botany, a caryopsis (plural caryopses) is a type of simple fruit—one that is monocarpellate (formed from a single carpel) and indehiscent (not opening at maturity) and resembles an achene, except that in a caryopsis the pericarp is f ...
", typical of the grasses
Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
, and the entire kernel is often referred to as the "seed
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
". The cob 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 pea
The pea is most commonly the small spherical seed or the seed-pod of the flowering plant species ''Pisum sativum''. Each pod contains several peas, which can be green or yellow. Botanically, pea pods are fruit, since they contain seeds and d ...
s, and adhere in regular rows around a white, pithy substance, which forms the cob. The maximum size of kernels is reputedly . An ear commonly holds 600 kernels. They are of various colors: blackish, bluish-gray, purple
Purple is any of a variety of colors with hue between red and blue. In the RGB color model used in computer and television screens, purples are produced by mixing red and blue light. In the RYB color model historically used by painters, ...
, green, red, white and yellow. When ground into flour
Flour is a powder made by grinding raw grains, roots, beans, nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredient of bread, which is a staple food for many c ...
, maize yields more flour with much less bran
Bran, also known as miller's bran, is the hard outer layers of cereal grain. It consists of the combined aleurone and pericarp. Corn (maize) bran also includes the pedicel (tip cap). Along with germ, it is an integral part of whole grains ...
than wheat does. It lacks the protein gluten
Gluten is a structural protein naturally found in certain cereal grains. Although "gluten" often only refers to wheat proteins, in medical literature it refers to the combination of prolamin and glutelin proteins naturally occurring in all gra ...
of wheat and, therefore, makes baked goods with poor rising capability. A genetic variant Genetic variant may refer to:
* Single-nucleotide polymorphism (SNP), in a case it is a common genetic variant
* Mutation, in a case where it is a rare genetic variant
* Copy-number variation
* Variant (biology)
See also
* Genetic variation (dis ...
that accumulates more sugar and less starch
Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
in the ear is consumed as a vegetable and is called sweet corn
Sweet corn (''Zea mays'' convar. ''saccharata'' var. ''rugosa''), also called sugar corn and pole corn, is a variety of maize grown for human consumption with a high sugar content. Sweet corn is the result of a naturally occurring recessive m ...
. 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
A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant growth. Wh ...
, the kernels dry out and become difficult to chew without cooking.
File:Cornsilk 7091.jpg, Female inflorescence, with young silk
Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the ...
File:Corn blooming.jpg, Mature silk
File:GreenCorn.JPG, Stalks, ears and silk
File:Männliche Blüte einer Maispflanze 2009-08-19.JPG, Male flowers
File:ZeaMays.jpg, Full-grown maize plants
File:Klip kukuruza uzgojen u Međimurju (Croatia).JPG, Mature maize ear on a stalk
Planting density affects multiple aspects of maize. Modern farming techniques in developed countries
A developed country (or industrialized country, high-income country, more economically developed country (MEDC), advanced country) is a sovereign state that has a high quality of life, developed economy and advanced technological infrastr ...
usually rely on dense planting, which produces one ear per stalk. Stands of silage
Silage () is a type of fodder made from green foliage crops which have been preserved by fermentation to the point of acidification. It can be fed to cattle, sheep and other such ruminants ( cud-chewing animals). The fermentation and storage ...
maize are yet denser, and achieve a lower percentage of ears and more plant matter.
Maize is a facultative short-day plant and flowers in a certain number of growing degree day
Growing degree days (GDD), also called growing degree units (GDUs), are a heuristic tool in phenology. GDD are a measure of heat accumulation used by horticulturists, gardeners, and farmers to predict plant and animal development rates such as the ...
s > 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 flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanism ...
s is genetically prescribed and regulated by the phytochrome
Phytochromes are a class of photoreceptor in plants, bacteria and fungi used to detect light. They are sensitive to light in the red and far-red region of the visible spectrum and can be classed as either Type I, which are activated by far-re ...
system. Photoperiodicity can be eccentric in tropical cultivar
A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
s such that 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.
Immature maize shoots accumulate a powerful antibiotic substance, 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA
DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) is a naturally occurring hydroxamic acid, a benzoxazinoid. DIMBOA is a powerful antibiotic present in maize, wheat, rye, and related grasses,
DIMBOA was first identified in maize in 1962 a ...
). DIMBOA is a member of a group of hydroxamic acid
A hydroxamic acid is a class of organic compounds bearing the functional group RC(O)N(OH)R', with R and R' as organic residues and CO as a carbonyl group. They are amides (RC(O)NHR') wherein the NH center has an OH substitution. They are often us ...
s (also known as benzoxazinoids) that serve as a natural defense against a wide range of pests, including insects, pathogenic
In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ger ...
fungi and bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
. DIMBOA is also found in related grasses, particularly wheat. A maize mutant (bx) lacking DIMBOA is highly susceptible to attack by aphid
Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A ...
s and fungi
A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
. DIMBOA is also responsible for the relative resistance of immature maize to the European corn borer
The European corn borer (''Ostrinia nubilalis''), also known as the European corn worm or European high-flyer, is a moth of the family Crambidae which includes other grass moths. It is a pest of grain, particularly maize (''Zea mays''). The ins ...
(family Crambidae
The Crambidae are the grass moth family of lepidopterans. They are variable in appearance, the nominal subfamily Crambinae (grass moths) taking up closely folded postures on grass stems where they are inconspicuous, while other subfamilies includ ...
). As maize matures, DIMBOA levels and resistance to the corn borer decline.
Because of its shallow roots, maize is susceptible to droughts, intolerant of nutrient-deficient soils, and prone to be uprooted by severe winds.
File:Ab food 06.jpg, Maize kernels
File:Maize plant diagram.svg, Maize plant diagram
File:Aa maize ear irregular 01.jpg, Ear of maize with irregular rows of kernels
While yellow maizes derive their color from lutein
Lutein (;["Lutein"](_blank)
zeaxanthin
Zeaxanthin is one of the most common carotenoids in nature, and is used in the xanthophyll cycle. Synthesized in plants and some micro-organisms, it is the pigment that gives paprika (made from bell peppers), corn, saffron, goji (wolfberries), ...
, in red-colored maizes, the kernel coloration is due to anthocyanin
Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical co ...
s and phlobaphene
Phlobaphenes (or phlobaphens, CAS No.:71663-19-9) are reddish, alcohol-soluble and water-insoluble phenolic substances. They can be extracted from plants, or be the result from treatment of tannin extracts with mineral acids (tanner's red). The nam ...
s. These latter substances are synthesized in the flavonoids synthetic pathway from polymerization of flavan-4-ol
The flavan-4-ols (3-deoxyflavonoids) are flavone-derived alcohols and a family of flavonoids. Flavan-4-ols are colorless precursor compounds that polymerize to form red phlobaphene pigments. They can be found in the sorghum. Glycosides (abacopterin ...
s by the expression of maize pericarp color1 (p1) gene which encodes an R2R3 myb-like transcriptional activator
A transcriptional activator is a protein (transcription factor) that increases transcription of a gene or set of genes. Activators are considered to have ''positive'' control over gene expression, as they function to promote gene transcription and ...
of the A1 gene encoding for the dihydroflavonol 4-reductase
In enzymology, a dihydrokaempferol 4-reductase () is an enzyme that catalyzes the chemical reaction
:cis-3,4-leucopelargonidin + NADP+ \rightleftharpoons (+)- dihydrokaempferol + NADPH + H+
Thus, the two substrates of this enzyme are cis-3,4- ...
(reducing dihydroflavonol
The flavanonols (with two "o"s a.k.a. 3-hydroxyflavanone or 2,3-dihydroflavonol) are a class of flavonoids that use the 3-hydroxy-2,3-dihydro-2-phenylchromen-4-one (IUPAC name) backbone.
Some examples include:
* Taxifolin (or Dihydroquercetin)
* ...
s into flavan-4-ols) while another gene (Suppressor of Pericarp Pigmentation 1 or SPP1) acts as a suppressor
A silencer, also known as a sound suppressor, suppressor, or sound moderator, is a muzzle device that reduces the acoustic intensity of the muzzle report (sound of a gunshot) and muzzle rise when a gun (firearm or air gun) is discharged, by ...
. The p1 gene encodes an Myb-homologous transcriptional activator of genes required for biosynthesis of red phlobaphene pigments, while the P1-wr allele specifies colorless kernel pericarp and red cobs, and unstable factor for orange1 (Ufo1) modifies P1-wr expression to confer pigmentation in kernel pericarp, as well as vegetative tissues, which normally do not accumulate significant amounts of phlobaphene pigments. The maize P gene encodes a Myb homolog that recognizes the sequence CCT/AACC, in sharp contrast with the C/TAACGG bound by vertebrate Myb proteins.
The ''ear leaf'' is the leaf most closely associated with a particular developing ear. This leaf and above contribute 70% to 75% to 90% of grain fill. Therefore fungicide application is most important in that region in most disease environments.
Abnormal flowers
Maize flowers may sometimes exhibit mutations that lead to the formation of female flowers in the tassel. These mutations, ''ts4'' and ''Ts6'', prohibit the development of the stamen while simultaneously promoting pistil development. This may cause inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed ...
s containing both male and female flowers, or hermaphrodite
In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes.
Many taxonomic groups of animals (mostly invertebrates) do not have ...
flowers.
Genetics
Maize is an annual grass in the family Gramineae
Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns and ...
, which includes such plants as wheat
Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeolog ...
, rye
Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is a member of the wheat tribe ( Triticeae) and is closely related to both wheat (''Triticum'') and barley (genus ''Hordeum''). Rye grain is ...
, barley
Barley (''Hordeum vulgare''), a member of the grass family, is a major cereal grain grown in temperate climates globally. It was one of the first cultivated grains, particularly in Eurasia as early as 10,000 years ago. Globally 70% of barley ...
, rice
Rice is the seed of the grass species '' Oryza sativa'' (Asian rice) or less commonly '' Oryza glaberrima'' (African rice). The name wild rice is usually used for species of the genera '' Zizania'' and ''Porteresia'', both wild and domestica ...
, sorghum
''Sorghum'' () is a genus of about 25 species of flowering plants in the grass family ( Poaceae). Some of these species are grown as cereals for human consumption and some in pastures for animals. One species is grown for grain, while many ot ...
, and sugarcane. There are two major species of the genus '' Zea'' (out of six total): ''Z. mays'' (maize) and '' Z. diploperennis'', which is a perennial
A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
type of teosinte. The annual
Annual may refer to:
* Annual publication, periodical publications appearing regularly once per year
**Yearbook
**Literary annual
* Annual plant
*Annual report
*Annual giving
*Annual, Morocco, a settlement in northeastern Morocco
*Annuals (band), ...
teosinte variety called '' Z. m. mexicana'' is the closest botanical relative to maize. It still grows in the wild as an annual in Mexico and Guatemala.
Many forms of maize are used for food, sometimes classified as various subspecies related to the amount of starch each has:
* Flour corn: ''Z. m.'' var. ''amylacea''
* Popcorn
Popcorn (also called popped corn, popcorns or pop-corn) is a variety of corn kernel which expands and puffs up when heated; the same names also refer to the foodstuff produced by the expansion.
A popcorn kernel's strong hull contains the se ...
: ''Z. m.'' var. ''everta''
* Dent corn
Dent corn, also known as grain corn, is a type of field corn with a high soft starch content. It received its name because of the small indentation, or "dent", at the crown of each kernel on a ripe ear of corn. Reid's Yellow Dent is a variety d ...
: ''Z. m.'' var. ''indentata''
* Flint corn
Flint corn (''Zea mays'' var. ''indurata''; also known as Indian corn or sometimes calico corn) is a variant of maize, the same species as common corn. Because each kernel has a hard outer layer to protect the soft endosperm, it is likened to bei ...
: ''Z. m.'' var. ''indurata''
* Sweet corn
Sweet corn (''Zea mays'' convar. ''saccharata'' var. ''rugosa''), also called sugar corn and pole corn, is a variety of maize grown for human consumption with a high sugar content. Sweet corn is the result of a naturally occurring recessive m ...
: ''Z. m.'' var. ''saccharata'' and ''Z. m.'' var. ''rugosa''
* Waxy corn: ''Z. m.'' var. ''ceratina''
* Amylomaize: ''Z. mays''
* Pod corn
Pod corn or wild maize is a variety of maize. It is not a wild ancestor of maize but rather a mutant that forms leaves around each kernel.
Pod corn (''tunicata'' Sturt) is not grown commercially, but it is preserved in some localities.
Pod corn ...
: ''Z. m.'' var. ''tunicata'' Larrañaga ex A. St. Hil.
* Striped maize: ''Z. m.'' var. ''japonica''
This system has been replaced (though not entirely displaced) over the last 60 years by multivariable classifications based on ever more data. Agronomic data were supplemented by botanical traits for a robust initial classification, then genetic, cytological
Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living a ...
, protein and DNA evidence was added. Now, the categories are forms (little used), races, racial complexes, and recently branches.
Maize is a diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respecti ...
with 20 chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins ar ...
s (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
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild sp ...
.
Barbara McClintock
Barbara McClintock (June 16, 1902 – September 2, 1992) was an American scientist and cytogeneticist who was awarded the 1983 Nobel Prize in Physiology or Medicine. McClintock received her PhD in botany from Cornell University in 1927. There ...
used these knob markers to validate her transposon
A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
theory of "jumping genes", for which she won the 1983 Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine ( sv, Nobelpriset i fysiologi eller medicin) is awarded yearly by the Nobel Assembly at the Karolinska Institute, Nobel Assembly at the Karolinska Institute for outstanding discoveries in physiology or ...
. Maize is still an important model organism for genetics and developmental biology
Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of ste ...
today.
The centromere
The centromere links a pair of sister chromatids together during cell division. This constricted region of chromosome connects the sister chromatids, creating a short arm (p) and a long arm (q) on the chromatids. During mitosis, spindle fibers ...
s have two types of structural components, both of which are found only in the centromeres: Large arrays of CentC, a short satellite DNA Satellite DNA consists of very large arrays of tandemly repeating, non-coding DNA. Satellite DNA is the main component of functional centromeres, and form the main structural constituent of heterochromatin.
The name "satellite DNA" refers to the p ...
; and a few of a family of retrotransposon
Retrotransposons (also called Class I transposable elements or transposons via RNA intermediates) are a type of genetic component that copy and paste themselves into different genomic locations (transposon) by converting RNA back into DNA through ...
s. The B chromosome, unlike the others, contains an additional repeat which extends into neighboring areas of the chromosome. Centromeres can accidentally shrink during division and still function, although it is thought this will fail if it shrinks below a few hundred kilobase. Kinetochore
A kinetochore (, ) is a disc-shaped protein structure associated with duplicated chromatids in eukaryotic cells where the spindle fibers attach during cell division to pull sister chromatids apart. The kinetochore assembles on the centromere ...
s contain RNA originating from centromeres. Centromere regions can become inactive, and can continue in that state if the chromosome still has another active one.
The Maize Genetics Cooperation Stock Center, funded by the USDA Agricultural Research Service
The Agricultural Research Service (ARS) is the principal in-house research agency of the United States Department of Agriculture (USDA). ARS is one of four agencies in USDA's Research, Education and Economics mission area. ARS is charged with ex ...
and located in the Department of Crop Sciences at the University of Illinois at Urbana-Champaign
The University of Illinois Urbana-Champaign (U of I, Illinois, University of Illinois, or UIUC) is a public land-grant research university in Illinois in the twin cities of Champaign and Urbana. It is the flagship institution of the Uni ...
, is a stock center of maize mutants. 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., tetraploid
Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
s). Genetic data describing the maize mutant stocks as well as myriad other data about maize genetics can be accessed a
MaizeGDB
the Maize Genetics and Genomics Database.
In 2005, the US National Science Foundation
The National Science Foundation (NSF) is an independent agency of the United States government that supports fundamental research and education in all the non-medical fields of science and engineering. Its medical counterpart is the National ...
(NSF), Department of Agriculture (USDA
The United States Department of Agriculture (USDA) is the federal executive department responsible for developing and executing federal laws related to farming, forestry, rural economic development, and food. It aims to meet the needs of com ...
) and the Department of Energy A Ministry of Energy or Department of Energy is a government department in some countries that typically oversees the production of fuel and electricity; in the United States, however, it manages nuclear weapons development and conducts energy-relat ...
(DOE) formed a consortium to sequence the B73 maize genome
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
. The resulting DNA sequence data was deposited immediately into GenBank
The GenBank sequence database is an open access, annotated collection of all publicly available nucleotide sequences and their protein translations. It is produced and maintained by the National Center for Biotechnology Information (NCBI; a part ...
, a public repository for genome-sequence data. Sequences and genome annotations have also been made available throughout the project's lifetime at the project's official site.
Primary sequencing of the maize genome was completed in 2008. On November 20, 2009, the consortium published results of its sequencing effort in ''Science''. The genome, 85% of which is composed of transposon
A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
s, was found to contain 32,540 genes (By comparison, the human genome
The human genome is a complete set of nucleic acid sequences for humans, encoded as DNA within the 23 chromosome pairs in cell nuclei and in a small DNA molecule found within individual mitochondria. These are usually treated separately as the ...
contains about 2.9 billion bases and 26,000 genes). Much of the maize genome has been duplicated and reshuffled by helitrons—group of rolling circle
Rolling circle replication (RCR) is a process of unidirectional nucleic acid replication that can rapidly synthesize multiple copies of circular molecules of DNA or RNA, such as plasmids, the genomes of bacteriophages, and the circular RNA genom ...
transposons.
In ''Z. mays'' and various other angiosperms the MADS-box
The MADS box is a conserved sequence motif. The genes which contain this motif are called the MADS-box gene family. The MADS box encodes the DNA-binding MADS domain. The MADS domain binds to DNA sequences of high similarity to the motif CC /Tsub>6G ...
motif is involved in floral development. Early study in several angiosperm models including ''Z. mays'' was the beginning of research into the molecular evolution of floral structure in general, as well as their role in nonflowering plants.
Varieties differ in their resistance to insects, including borers. CIMMYT
The International Maize and Wheat Improvement Center (known - even in English - by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de Maíz y Trigo'') is a non-profit research-for-development organization that develops im ...
maintains a large collection of maize/corn accessions tested and cataloged for insect resistance.[
]
Evolution
As with many plants and animals, ''Z. mays'' has a positive correlation between effective population size
The effective population size (''N'e'') is a number that, in some simplified scenarios, corresponds to the number of breeding individuals in the population. More generally, ''N'e'' is the number of individuals that an idealised population w ...
and the magnitude of selection pressure
Any cause that reduces or increases reproductive success in a portion of a population potentially exerts evolutionary pressure, selective pressure or selection pressure, driving natural selection. It is a quantitative description of the amount of ...
. ''Z. m.'' having an EPS of ~650,000, it clusters with others of about the same EPS, and has 79% of its amino acid
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
sites under selection.
Recombination is a significant source of diversity in ''Z. mays''. (Note that this finding supersedes previous studies which showed no such correlation.)
This recombination/diversity effect is seen throughout plants but is also found to not occur – or not as strongly – in regions of high gene density. This is likely the reason that domesticated ''Z. mays'' has not seen as much of an increase in diversity within areas of higher density as in regions of lower density, although there is more evidence in other plants.
Some lines of maize have undergone ancient polyploid
Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contain ...
y events, starting 11m years ago. Over that time ~72% of polyploid duplicated genes have been retained, which is higher than other plants with older polyploidy events. Thus maize ''may'' be due to lose more duplicate genes as time goes along, similar to the course followed by the genomes of other plants. If so - if gene loss has merely not occurred yet - that could explain the lack of observed positive selection
In population genetics, directional selection, is a mode of negative natural selection in which an extreme phenotype is favored over other phenotypes, causing the allele frequency to shift over time in the direction of that phenotype. Under dir ...
and lower negative selection which are observed in otherwise similar plants, i.e. also naturally outcrossing
Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity.
Outcrossing can be a useful ...
and with similar effective population sizes.
Ploidy does not appear to influence EPS or magnitude of selection effect in maize.
Breeding
Maize reproduces sexually each year. This randomly selects half the genes from a given plant to propagate to the next generation, meaning that desirable traits found in the crop (like high yield or good nutrition) can be lost in subsequent generations unless certain techniques are used.
Maize breeding in prehistory resulted in large plants producing large ears. Modern breeding began with individuals who selected highly productive varieties in their fields and then sold seed to other farmers. James L. Reid was one of the earliest and most successful developing Reid's Yellow Dent in the 1860s. These early efforts were based on mass selection. Later breeding efforts included ear to row selection (C. G. Hopkins c. 1896), hybrids made from selected inbred
Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction
Human reproduction is sexual reproduction that result ...
lines (G. H. Shull, 1909), and the highly successful double cross hybrids using four inbred lines (D. F. Jones
Dennis Feltham Jones (15 July 1918 – 1 April 1981) was a British science fiction author who published under the name D.F. Jones. He was a Royal Navy commander during World War II and lived in Cornwall.
His first novel, ''Colossus'' (1966), a ...
c. 1918, 1922). University supported breeding programs were especially important in developing and introducing modern hybrids. By the 1930s, companies such as Pioneer devoted to production of hybrid maize had begun to influence long-term development. Internationally important seed banks such as the International Maize and Wheat Improvement Center
The International Maize and Wheat Improvement Center (known - even in English - by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de Maíz y Trigo'') is a non-profit research-for-development organization that develops im ...
(CIMMYT) and the US bank at the Maize Genetics Cooperation Stock Center University of Illinois at Urbana-Champaign
The University of Illinois Urbana-Champaign (U of I, Illinois, University of Illinois, or UIUC) is a public land-grant research university in Illinois in the twin cities of Champaign and Urbana. It is the flagship institution of the Uni ...
maintain germplasm
Germplasm are living genetic resources such as seeds or tissues that are maintained for the purpose of animal and plant breeding, preservation, and other research uses. These resources may take the form of seed collections stored in seed banks, t ...
important for future crop development.
Since the 1940s the best strains of maize have been first-generation hybrids made from inbred strains that have been optimized for specific traits, such as yield, nutrition, drought, pest and disease tolerance. Both conventional cross-breeding and genetic engineering have succeeded in increasing output and reducing the need for cropland, pesticides, water and fertilizer. There is conflicting evidence to support the hypothesis that maize yield potential has increased over the past few decades. This suggests that changes in yield potential are associated with leaf angle, lodging resistance, tolerance of high plant density, disease/pest tolerance, and other agronomic traits rather than increase of yield potential per individual plant.
Certain varieties of maize have been bred to produce many ears which are the source of the "baby corn
An infant or baby is the very young offspring of human beings. ''Infant'' (from the Latin word ''infans'', meaning 'unable to speak' or 'speechless') is a formal or specialised synonym for the common term ''baby''. The terms may also be used to ...
" used as a vegetable in Asian cuisine
Asian cuisine includes several major regional cuisines: Central Asian, East Asian, North Asian, South Asian, Southeast Asian, and West Asian. A cuisine is a characteristic style of cooking practices and traditions, usually associated wit ...
.
CIMMYT operates a conventional breeding program to provide optimized strains. The program began in the 1980s. Hybrid seeds are distributed in Africa by the Drought Tolerant Maize for Africa project.
Genetic engineering
Genetically engineered
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including t ...
(GE) maize was one of the 26 GE crops grown commercially in 2016. The vast majority of this is Bt maize
Genetically modified maize (corn) is a genetically modified crop. Specific maize strains have been genetically engineered to express agriculturally-desirable traits, including resistance to pests and to herbicides. Maize strains with both trait ...
. Grown since 1997 in the United States and Canada, 92% of the US maize crop was genetically modified in 2016 and 33% of the worldwide maize crop was GM in 2016. As of 2011, Herbicide-tolerant maize varieties were grown in Argentina, Australia, Brazil, Canada, China, Colombia, El Salvador, the European Union, Honduras, Japan, Korea, Malaysia, Mexico, New Zealand, Philippines, the Russian Federation, Singapore, South Africa, Taiwan, Thailand, and the United States. Insect-resistant maize was grown in Argentina, Australia, Brazil, Canada, Chile, China, Colombia, Egypt, the European Union, Honduras, Japan, Korea, Malaysia, Mexico, New Zealand, Philippines, South Africa, Switzerland, Taiwan, the United States, and Uruguay.
In September 2000, up to $50 million worth of food products were recalled due to the presence of Starlink
Starlink is a satellite internet constellation operated by SpaceX, providing satellite Internet access coverage to 45 countries. It also aims for global mobile phone service after 2023. SpaceX started launching Starlink satellites in 2019. As ...
genetically modified corn, which had been approved only for animal consumption and had not been approved for human consumption, and was subsequently withdrawn from the market.
For pest and disease resistance
Tropical landrace
A landrace is a domesticated, locally adapted, often traditional variety of a species of animal or plant that has developed over time, through adaptation to its natural and cultural environment of agriculture and pastoralism, and due to isolati ...
s remain an important and underused source of resistance alleles for disease and for herbivores. Notable discoveries of rare alleles for this purpose were made by Dao ''et al.'', 2014 and Sood ''et al.'', 2014. Rashid ''et al.'', 2018 use an association mapping panel from CIMMYT
The International Maize and Wheat Improvement Center (known - even in English - by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de Maíz y Trigo'') is a non-profit research-for-development organization that develops im ...
originally developed for tropical drought tolerance traits to find new genomic
Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
regions providing sorghum downy mildew resistance, and to further characterize SDMR regions already located by others.[ .]
Origin
Maize is the domesticated variant of teosinte
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild sp ...
.[
*] The two plants have dissimilar appearance, maize having a single tall stalk with multiple leaves and teosinte being a short, bushy plant. The difference between the two is largely controlled by differences in just two genes, called grassy tillers-1 (''gt1'', ) and teosinte branched-1 (''tb1'', ).
Several theories had been proposed about the specific origin of maize in Mesoamerica:
# It is a direct domestication
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
of a Mexican
Mexican may refer to:
Mexico and its culture
*Being related to, from, or connected to the country of Mexico, in North America
** People
*** Mexicans, inhabitants of the country Mexico and their descendants
*** Mexica, ancient indigenous people ...
annual teosinte
''Zea'' is a genus of flowering plants in the grass family. The best-known species is ''Z. mays'' (variously called maize, corn, or Indian corn), one of the most important crops for human societies throughout much of the world. The four wild sp ...
, ''Z. m.'' ssp. ''parviglumis'', native to the Balsas River
The Balsas River (Spanish Río Balsas, also locally known as the Mezcala River, or Atoyac River) is a major river of south-central Mexico.
The basin flows through the states of Guerrero, México, Morelos, and Puebla. Downstream of Ciudad Al ...
valley in south-eastern Mexico, with up to 12% of its genetic material
Nucleic acids are biopolymers, macromolecules, essential to all known forms of life. They are composed of nucleotides, which are the monomers made of three components: a 5-carbon sugar, a phosphate group and a nitrogenous base. The two main clas ...
obtained from ''Z. m.'' ssp. ''mexicana'' through introgression
Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intro ...
.
# It has been derived from hybridization between a small domesticated maize (a slightly changed form of a wild maize) and a teosinte of section ''Luxuriantes'', either ''Z. luxurians'' or '' Z. diploperennis''.
# It has undergone two or more domestications either of a wild maize or of a teosinte. (The term "teosinte" describes all species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
and subspecies in the genus ''Zea'', excluding ''Z. m.'' ssp. ''mays''.)
# It has evolved from a hybridization of ''Z. diploperennis'' by ''Tripsacum dactyloides
''Tripsacum dactyloides'', commonly called eastern gamagrass, or Fakahatchee grass, is a warm-season, sod-forming bunch grass. It is widespread in the Western Hemisphere, native from the eastern United States to northern South America. ''.
In the late 1930s, Paul Mangelsdorf suggested that domesticated maize was the result of a hybridization event between an unknown wild maize and a species of ''Tripsacum
''Tripsacum'' is a genus of plants in the grass family native to the Western Hemisphere. Gamagrass is a common name for plants in this genus.
Species
formerly included
see '' Anthephora Apluda Chionachne Coelorachis Elionurus Hackelochloa ...
'', a related genus. This theory about the origin of maize has been refuted by modern genetic testing
Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
, which refutes Mangelsdorf's model and the fourth listed above.
The teosinte origin theory was proposed by the Russian botanist Nikolai Ivanovich Vavilov
Nikolai Ivanovich Vavilov ( rus, Никола́й Ива́нович Вави́лов, p=nʲɪkɐˈlaj ɪˈvanəvʲɪtɕ vɐˈvʲiləf, a=Ru-Nikolay_Ivanovich_Vavilov.ogg; – 26 January 1943) was a Russian and Soviet agronomist, botanist ...
in 1931 and the later American Nobel Prize
The Nobel Prizes ( ; sv, Nobelpriset ; no, Nobelprisen ) are five separate prizes that, according to Alfred Nobel's will of 1895, are awarded to "those who, during the preceding year, have conferred the greatest benefit to humankind." Alfre ...
-winner George Beadle
George Wells Beadle (October 22, 1903 – June 9, 1989) was an American geneticist. In 1958 he shared one-half of the Nobel Prize in Physiology or Medicine with Edward Tatum for their discovery of the role of genes in regulating biochemical e ...
in 1932. It is supported experimentally and by recent studies of the plants' genomes. Teosinte and maize can cross-breed and produce fertile offspring. A number of questions remain concerning the species, among them:
# how the immense diversity of the species of sect. ''Zea'' originated,
# how the tiny archaeological specimens of 3500–2700 BC could have been selected from a teosinte, and
# how domestication could have proceeded without leaving remains of teosinte or maize with teosintoid traits earlier than the earliest known until recently, dating from ca. 1100 BC.
The domestication
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
of maize is of particular interest to researchers— archaeologists, geneticist
A geneticist is a biologist or physician who studies genetics, the science of genes, heredity, and variation of organisms. A geneticist can be employed as a scientis