NATIONAL

CORN
HANDBOOK

THE CORN CROP NCH-10

Origin, Adaptation, and Types of Corn

W. L. Brown, Pioneer Hi-Bred International, Inc., IA; M. S. Zuber, University of MIssouri;
L. L. Darrah, USDA-ARS, University of MIssouri; and D. V. Glover, Purdue University

Reviewers
R. G. Creech, Mississippi State University A. A. Fleming, University of Georgia
K. F. Schertz, USDA-ARS, Texas A&M University A. F. Troyer, DeKalb-Pfizer Genetics, IL

ORIGIN OF CORN attention were given to determining corn’s origin and

Corn (Zea mays L.) is the only important cereal more to understanding the remarkable variability found
indigenous to the Western Hemisphere. Apparently within the species.
originating in Mexico, it spread northward to Canada
and southward to Argentina. While the possibility of Variability and Races
secondary centers of origin in South America cannot Regardless of origin, corn has proven to be one of
be completely ruled out, the oldest (7000 years) the most adaptable and variable members of the grass
archaeological corn was found in Mexico’s Valley of family. Its evolution, a large part of which apparently
Tehuacan. occurred under domestication, has resulted in biotypes
The earliest “corn” of which there is record is with adaptation ranging from the tropics to the north
unmistakably corn. The female inflorescence of this temperate zone, from sea level to 12,000 feet altitude
5000 B.C. corn had reached a degree of specialization and growing periods (planting to maturity) extending
that precluded the possibility of natural seed dissemi- from 6 weeks to 13 months.
nation. Thus, the oldest corn of record was dependent Almost 300 races of corn have been described
upon man for its survival. from Mexico, Central and South America, and the
Numerous theories of origin have been offered Caribbean. Although many appear synonymous, at
over the years, only two of which receive serious con- least 150 distinct entities have been collected in these
sideration today. One is that teosinte (Zea mexicana) areas. It was from certain of these races that most of
is the wild progenitor of corn; the other is that a wild the corns of North America were ultimately derived.
pod corn, now extinct, was the ancestor of domesti-
cated corn. While perhaps more students of corn seem Spread from Center of Origin
to accept the first theory, others are equally convinced Following discovery, corn moved quickly to Europe,
of the second. Africa and Asia. From Spain, it spread northward to
Aside from its possible role in the origin of corn, the short-growing-season areas of France, Germany,
teosinte has had major impact on its evolution. In Mexico Austria and eastern Europe, where selection for early
particularly, introgression between corn and teosinte maturity has produced some of the earliest commer-
has likely occurred for centuries and continues to this cial varieties of corn now available. In Italy and Spain,
day. The effects are apparent in the morphology and early counterparts of many South American races are
cytology of both species. There is also reason to be- evident even today.
lieve that genes for resistance to certain viruses have Although introduced into Africa soon after discovery,
reached corn through its introgression with teosinte. much of the corn now found in that continent is derived
The origin of corn may never be known with cer- from later introductions from the southern U.S., Mexico
tainty. One reason is that the hypotheses purporting to and parts of eastern South America. Most of southern
explain origin cannot be tested experimentally. There- Africa’s corn traces back to varieties grown in the
fore, science would perhaps be better served if less southern U.S. in colonial and post-colonial times. Corn

Cooperative Extension Service

Iowa State University
Ames, Iowa 50011
May 1985
of tropical middle (lowland) Africa is similar to the low- and moisture levels, (3) humidity of the atmosphere
land and tropical corns of Central and South America. immediately surrounding the plant, (4) wind move-
The most widely used and productive corns of ment, (5) day length, (6) light intensity, (7) air pollu-
Asia are derived from Caribbean-type flints introduced tion, (8) soil type, (9) soil fertility, (10) number of days
in relatively recent times. However, older and quite exceeding 50°F, (11) competition from other plants
distinct types of corn can also be found, for example, including its intended neighbors as well as weeds, and
among the hill people of Mindanao in southern Philip- (12) the disease-insect complex.
pines. Included are some small-eared, early-maturing Many of these factors interact in a complex man-
flints or pops that either have the capacity to grow ner to produce stress on the plant. The plant’s reac-
and mature quickly before being devastated by downy tion to stress is under genetic control, and differences
mildew or carry some genetic resistance to downy among hybrids exist. Corn breeders are continuously
mildew. developing and testing new genetic combinations in
In the eastern Himalayas (Sikkim and Bhutan), a differing environments to find types that give the best
distinctive type of popcorn is found whose Western agronomic performance over a range of geographical
Hemisphere progenitors seem to have disappeared. locations and years.
When and how the ancestor of this corn reached Asia Corn is grown from sea-level to altitudes of more
is not known. However, since it is not present on the than 12,000 feet and from the equator to about 50°
Indian subcontinent to the south, it may have reached north and south latitudes. Compared to environmen-
the Himalayas by way of China and Tibet. In any tal conditions of the U.S. Corn Belt, many producing
event, a search for similar corns in south China and areas would be considered very much substandard. In
Tibet would seem justified. harsh-environment areas, the varieties grown would
be considered adapted because they responded the
Corn of the U.S. Corn Belt best under the unfavorable growing conditions. The
Whereas most of the modern races of corn are original open-pollinated varieties have become adapt-
derived from prototypes developed by early native ed through selection over time by both man and the
agriculturists of Mexico, Central and South America, environment.
one outstanding exception is solely the product of
post-colonial North America—the yellow dent corns Adaptation Within the Corn Belt
that dominate the U.S. Corn Belt, Canada and much of World production of corn in the early 1980s ap-
Europe today. The origin and evolution of this remark- proached 450 million metric tons annually, with the
able race of corn have been clearly documented and U.S. contributing over 48 percent of the total. Of U.S.
confirmed. annual production, the 13 Corn Belt states account for
In the early 1800s, two predominant races of corn about 82 percent.
of North America’s eastern seaboard—the late-matur- No area of the world equals the Corn Belt for high
ing Virginia Gourdseed and the early-maturing North- yields. This is due to a combination of ideal soils and
eastern Flints—were first crossed, and the superiority climate, advanced farmer know-how, and the suc-
of the hybrid recognized and described. The cross was cess of corn breeders in developing hybrids with high
repeated many times during the western migration of genetic potential.
settlers; and out of these mixtures eventually emerged Climate. Corn is considered a warm-weather
the Corn Belt dents, the most productive race of corn crop. In the Corn Belt, average summer temperatures
found anywhere in the world. range from 70° to 80°F daytime and exceed 58°F at
It was the highly selected varieties of Corn Belt night. The average frost-free growing period is over
dents that formed the basis of hybrid corn and the 140 days. Greatest yields are obtained where 30 or
source of the first inbred lines used to produce hybrids. more inches of rain occur during the growing season.
Germplasm from some of these varieties (Reid, Lan- In areas where rainfall is less than 20 inches, yields
caster, Krug, etc.) still figure prominently in the ances- are much reduced unless irrigation is used.
try of hybrids used in the Corn Belt even to this day. Rainfall distribution greatly influences maximum
yields, especially for the 3-week period centered
ADAPTATION OF CORN around tasseling. In the southern part of the Corn Belt,
high-temperature stress and rainfall deficiencies often
Definition and Adaptation Worldwide occur in late July and August. Therefore, farmers of
Adaptation in corn means good performance with this region try to avoid having corn tassel during this
respect to yield and other agronomic characteristics dry, hot period by planting earlier and using earlier-
in a given environment. The environment includes flowering hybrids.
all conditions to which the plant is subjected during Maturity. Maturity of corn hybrids is a genetic
the growing season (from pre-seedling emergence to characteristic and is generally defined as the period
harvest maturity). from germination to when the kernel ceases to increase
The major environmental factors are: (1) daily max- in weight. In the northern Corn Belt, early hybrids often
imum and minimum temperatures, (2) soil atmosphere reach physiological maturity in less than 100 days;
whereas in the extreme south, 150 or more days may

2
be needed. These day periods are measures of rela- Dent Corn
tive maturity. The U.S. Corn Belt dents originated from the
Hybrids have traditionally been classified into hybridization of the Southern Dent or late-flowering
15 maturity groups ranging from Agricultural Experiment maize race called Gourdseed, and the early-flowering
Station (AES) 100 to AES 1500 (earliest to latest). Northern Flints. Dent corn is characterized by the
However, maturity classification can be made more presence of corneous, horny endosperm at the sides
precise by determining the total heat units required and back of the kernels, while the central core is a
from emergence to physiological maturity. A heat unit soft, floury endosperm extending to the crown of the
measure commonly used accumulates the daily ex- endosperm where, upon drying, it collapses to produce
cess of average temperature over 50°F, where: a distinct indentation. Degree of denting varies with
average temperature = (maximum + minimum) ÷2. the genetic background. Nearly all varieties grown in
the U.S. are yellow, with only a few white endosperm
Early hybrids grown in the northern Corn Belt clas- types grown.
sified as 100-day maturity may require 90 or less days Dent corn is used primarily as animal food, but
to reach maturity when grown further south where heat also serves as a raw material for industry and as a sta-
unit accumulation is more rapid. ple food. Upwards of 93 percent of dent corn produced
Adapted hybrid development. Hybrids have (including the corn equivalent of by-product feeds from
been developed that are adapted from Nebraska to corn processing) is used as animal feeds. However, it
Ohio. However, the environmental conditions differ is still an important human food and industrial material,
widely from west to east especially rainfall patterns, entering into many specialized products via the dry- or
daily minimum and maximum temperatures, and the wet-milling industry in the U.S.
disease complex. For example, the hotter, drier condi- Yellow dent corn sells at market price as it enters
tions of the western Corn Belt are less favorable for the normal feed grain or milling channels. However,
leaf diseases caused by Helminthosporium spp. and white dent often receives a premium price in the dry-
anthracnose, but more conducive to viral infection, milling industry, where it is utilized for certain human
bacterial wilt and smut. For this reason, development food products because of its whiter starch.
of hybrids adapted from west to east has been rela-
tively difficult and requires large-scale testing over a Flint Corn
number of locations and years. The flint corns mostly have a thick, hard, vitreous
Number of frost-free days decreases from the south- (glassy) or corneous endosperm layer surrounding a
ern to northern Corn Belt; however, hours of daylight on small, soft granular center. The relative amounts of
June 22nd are much longer in the north than in the south. soft and corneous starch, however, vary in different
Sunlight intensity is greater in the western and northern varieties. Generally, the kernels are smooth and
Corn Belt because these sections have less cloudy weather. rounded, and the ears long and slender with a com-
The result of intensive corn breeding efforts over paratively small number of rows or kernels. In temper-
the past four decades has been better adaptation to the ate zones, flint corn often matures earlier, germinates
many environments under which corn is grown. better, has more spring vigor, more tillers and fewer
prop roots than dent strains.
TYPES OF CORN Very little flint corn is produced and utilized in the
Corn variation may be artificially defined accord- U.S. today, although it was undoubtedly grown ex-
ing to kernel type as follows: dent, flint, flour, sweet, tensively up through colonial times. Generally, yields
pop and pod corn. Except for pod corn, these divisions are lower than our Corn Belt dents, in part because of
are based on the quality, quantity and pattern of en- relatively little breeding work done. Flints are more ex-
dosperm composition in the kernel and are not indica- tensively grown in Argentina and other areas of South
tive of natural relationships. America, Latin America and southern Europe where
Endosperm composition may be changed by a they are used for feed and food.
single gene difference, as in the case of floury (fl)
versus flint (FI), sugary (su) versus starchy (Su), waxy Flour Corn
(wx) versus non-waxy (Wx), and other single reces- This is one of the. oldest types of corn, tracing
sive gene modifiers that have been used in breeding back to the ancient Aztecs and Incas. American
special-purpose types of corn. The quantity or volume Indians ground the soft kernels for flour. Floury maize
of endosperm conditioning the size of the kernel (e.g., types have soft starch throughout, with practically no
the difference between dent and flint corns or flint corn hard, vitreous endosperm and thus are opaque in ker-
and popcorn) is polygenic and, in the latter example, is nel phenotype. Kernels tend to shrink uniformly upon
of some taxonomic significance. drying, so usually have little or no denting. When dry,
The pod corn trait is monogenic and more of an or- they are easy to grind, but may mold on the mature
namental type. The major gene involved (Tu) produces ear in wet areas.
long glumes enclosing each kernel individually, such
as occurs in many other grasses.

3
 In the U.S., flour corn has limited production and is readily identifiable by cultivar name alone. At least
restricted to the drier sections. It is grown widely in the 13 endosperm mutants, in combination with sugary,
Andean region of South America. have been studied for improving sweet corn. Except
for sugary, the genes used in breeding act differently
Sweet Corn to produce the taste and texture deemed desirable for
The following genetic model featuring primary iso- sweet corn.
lation groups for naming “vegetable corns” has been Augmented sugary kernel types. In these sweet
suggested by the industry: corns, the sugars are modified (increased) by the action
I. Sugary mutants of other genes, either partially or completely.
A. Standard sugary (su) Major modifier genes of kernel sweetness are
B. Augmented sugary shrunken-2 (sh2) and sugary enhancer (se). In partial
1 . Partial modification modifications, the sugary (su) kernels are modified
a. Heterozygous shrunken -2 (sh2) by the segregation of major modifier genes such that
b. Heterozygous sugary enhancer (se) about 25 percent of the kernels are double-mutant
c. Heterozygous shrunken -2 and sugary endosperm types possessing the enhanced benefits of
enhancer (sh2 and se) the modifier. The addition of the sugary enhancer (se)
2. Complete (100%) modification gene along with one of the major modifier genes (e.g.,
a. Homozygous sugary enhancer (se) sh2) will further modify some of the sugary kernels
II. Shrunken-2 (sh2) to about 44 percent double-mutant endosperm types
III. Brittle (bt) rather than 25 percent.
IV. Brittle -2 (bt2) In complete (100 percent) modification, the sugary
V. Amylose-extender (ae) Dull (du) Waxy (wx) (su) kernels are all modified with the sugary enhancer
VI Dent (vegetable) (se) gene to produce the double combination (su se)
VII. Additional classes as new genes are used for obtaining maximum benefit from the se gene. Other
Isolation will be required between major groups major modifier genes of kernel sweetness are: brittle
identified by a Roman numeral. Isolation is suggested (bt), brittle-2 (bt2), shrunken (sh) and shrunken-4
but not required between subgroups within a major (Sh4). Other genes with minor modifying effects of
group. No isolation is needed for cultivars within the kernel sweetness are: dull (du), floury (fI), floury-2 (fl2)
same classification. opaque (o), opaque-2 (o2), sugary-2 (su2), and waxy
Standard sugary kernel types. Sweet corn, com- (wx). Some are known to be present in sweet corn
monly referred to as the standard sugary (su) corn, backgrounds either in the segregating or homozygous
is thought to have originated from a mutation in the state.
Peruvian race Chullpi. Most certainly it was grown and Other mutants producing sugary kernels include
used by native American Indians in pre-Columbian the single-mutant endosperm genes shrunken-2 (sh2),
times. brittle (bt) and brittle-2 (bt2), and the multiple-mutant
In sweet corn, the sugary gene prevents or retards endosperm genes amylose-extender, dull, waxy (ae du
the normal conversion of sugar into starch during wx).
endosperm development, and the kernel accumulates Precautions with modified endosperm sweet
a water-soluble polysaccharide called “phytoglycogen.” corn to avoid xenia. Isolation of “sweet corn” cultivar
As a result, the dry, sugary kernels are wrinkled and plantings of different genetic types is necessary to
glassy. The higher content of water-soluble polysac- prevent cross-pollination. Xenia is the immediate effect
charide adds a texture quality factor in addition to of foreign pollen on a variety; on sweet corn (su), it will
sweetness. In the U.S., sweet corn is eaten in the produce a starchy kernel. Isolation can be obtained by
immature milk stage and is one of the most popular planting at a different time, planting cultivars of different
vegetables. maturities, planting “upwind” of prevailing wind direc-
Sweet corn in the U.S. is more important economi- tion, or providing barriers and border rows. All of these
cally than its limited commercial production would methods will reduce the isolation distances necessary.
indicate, because it is consumed directly as human On a practical basis, commercial growers should pro-
food (fresh market or canned and frozen products) vide at least 50 feet separation, plant upwind of normal
rather than indirectly as livestock feed. The bulk of field corn, and use four or more border rows.
sweet corn production is confined to the northern tier
of states and to southern Florida as a winter crop. Popcorn
In the broader sense, vegetable corns include all Popcorns are perhaps the most primitive of the
corn harvested and eaten while the kernels are still surviving races of maize. This corn type is characterized
tender and before all of the sugars are converted to by a very hard, corneous endosperm containing only a
starch. This definition includes “roasting ears” of se- small portion of soft starch. Popcorns are essentially
lected field corns. small-kerneled flint types. The kernels may be either
Today, the standard sugary corns are being modified pointed (rice-like) or round (pearl-like). Some of the
with other endosperm genes and gene combinations more recently developed popcorns have thick pericarps
that control sweetness to develop new cultivars. As (seed coats), while some primitive semi-popcorns,
a result, growers must consider genetic type when such as the Argentine popcorns, have thin pericarps.
making selections for planting. The genetic type is not
4
 Popcorn is a relatively minor crop compared to pie fillings, sauces, gravies, retorted foods, salad
dent corn. It is used primarily for human consumption dressings, etc. Other waxy products are used as re-
as freshly popped corn or as the basis of popcorn con- moistening adhesives in the manufacture of gummed
fections. Isolated planting is not necessary, since there tape, in adhesives and in the paper industry. Waxy
are no major xenia effects on popping expansion and grain is also grown as a feed for dairy cattle and live-
many popcorns are cross-sterile with field corn. stock.
Most popcorn acreage is grown under contract. Waxy corn is usually grown under contract for the
Although conditions for growing popcorn are the major wet millers and exporters. Premiums are paid
same as for dent corn, special harvesting, drying and to the growers of waxy corn for wet milling because
storage practices are necessary to maintain popping it must be isolated during production, harvesting,
quality. transporting and storing. Since waxy is a recessive
characteristic, isolation from dent corn is necessary to
Pod Corn prevent loss of its peculiar starch properties.
Pod corn (tunicate maize) is more of an ornamen- High-amylose corn. Amylomaize is the generic
tal type. The major gene involved (Tu) produces long name for corn that has an amylose content higher than
glumes enclosing each kernel individually, such as 50 percent. The endosperm mutant amylose extender
occurs in many other grasses. The ear is also enclosed (ae) found by R. P. Bear in 1950, increases the amylose
in husks, as with other types of corn. content of the endosperm to about 60 percent in many
Homozygous pod corn usually is highly self-sterile, dent backgrounds. Modifying factors alter the amylose
and the ordinary type of pod corn is heterozygous. Pod contents as well as desirable agronomic characteristics
corn may be dent, sweet, waxy, pop, flint or floury in of the grain. The amylose-extender gene expression is
endosperm characteristics. It is merely a curiosity and characterized by a tarnished, translucent, sometimes
is not grown commercially. semi -full kernel appearance.
High-amylose grain is grown exclusively for wet
Special-Purpose Corns milling. The two types produced commercially are
Corn may be altered by genetic means to produce Class V (amylose content, 50-60 percent) and Class
modifications in starch, protein, oil and other prop- VII (amylose content, 70-80 percent). The starch from
erties. As a result of modifications of ordinary dent high-amylose corn is used in the textile industry, in
types, new corn specialties have been created. Among gum candies (where its tendency to form a gel aids
them are waxy-maize, amylomaize, and high-lysine or production), and as an adhesive in the manufacture of
modified-protein corn. corrugated cardboard.
Waxy corn. This special-purpose type was intro- High-amylose corn yields vary depending upon
duced to the U.S. from China in 1908. Although China location, but average only 65-75 percent of that of
was the original source, waxy (wx) mutations have ordinary dents. Present production acreage is limited
since been found in American dent strains. Its name to that grown under contract arrangements for wet
derives from the waxy appearance of the endosperm millers. Premiums are paid to growers because of
exposed in a cleanly cut cross-section. Common corn decreased yields and the necessity to isolate high
starch is approximately 73 percent amylopectin and amylose corn during production, harvesting, transport-
27 percent amylose, whereas waxy starch is com- ing and storing. The premium depends upon class,
posed entirely of amylopectin, which is the branched year and desired acreage.
molecular form. Ordinary cornstarch stains blue with High-lysine corn. This is the generic name for corn
2 percent potassium iodide solution, whereas waxy having an improved amino acid balance, thus a better
cornstarch stains a reddish brown. The waxy gene protein quality for feeding and food use compared to
also expresses itself in the pollen with this staining ordinary dent types. E. T. Mertz in 1964 discovered that
reaction, which is an aid in breeding. the single recessive gene, opaque-2 (o2), reduced zein
Significant advances in yields have been made in the endosperm and increased the percent of lysine to
with the newer waxy hybrids. While the overall aver- improve nutritional quality. Other genes with similar gross
age may run somewhat less than dent corn hybrids, effects on protein quality exist in corn, but attempts to
the newer waxy hybrids are more comparable to the improve corn protein quality have been primarily based
better dents in yields. on use of the opaque-2 gene and modified opaque-2
Waxy corn has carved out a formidable position as germplasm.
the raw material of waxy cornstarch produced by cer- The opaque-2 gene is characterized by a soft,
tain wet-corn millers in the U.S., Canada, Europe, etc., chalky, non-transparent kernel appearance, having
for industry and food uses. Waxy starch and modified practically no hard vitreous or horny endosperm. Unde-
waxy starches are sold extensively worldwide because sirable kernel characteristics (e.g., kernel and ear rots)
of their stability and other properties of their solutions. and insect and rodent damage can be a problem with
Products made from waxy corn are used by the the soft opaque-2 chalky phenotypes. Improvements in
food industry as stabilizers and thickeners for puddings, resistance to ear and kernel rots have been substantial

5
with selection, and a number of good hybrids exist. Current U.S. use of high-lysine corn is restricted
On the average, the opaque-2 hybrids yield about because of (1) yield differentials compared to normal
7-10 percent lower than their normal counterparts. corn and (2) the corn-to-soybean oil/meal price
A promising approach to overcoming some of the relationship. Demand for high-quality protein corn in
deficiencies of the homozygous opaque-2 materials the U.S. is insufficient to command a premium price.
involves the visual selection of specific modifiers of However, high-lysine corn is grown to a limited extent
opaque-2. It is fairly easy to develop modified, vitreous as a feed for poultry, swine, dairy cattle and other live-
opaque-2 materials with good ear rot and grain insect stock production needs. In corn-dependent countries
resistance. Selections must include endosperm chemi- where normal corn is a major staple of the human diet,
cal analyses to maintain high levels of protein quality. or where high-quality protein supplements for animal
Another approach to endosperm textural modifi- feeding are scarce, yield is a secondary consideration.
cation to solve some of the problems associated with Some high-lysine materials are to the point of develop-
opaque-2 corn has been use of the double-mutant ment where it may be cost-efficient to grow quality-
combination, sugary-2 opaque-2 (su2 o2). This modifi- protein corn as a specialty crop.
cation has improved kernel vitreousness, density and Ornamental corn. The so-called ornamental or
resistance to kernel breakage. The improved vitreous “Indian” corns commonly show segregation for alleles of
of su2/o2 is accompanied by protein quality at least several genetic factors that control the production of an-
equal to the unmodified opaque-2 materials; however, thocyanins and related pigments in the aleurone, pericarp
at this point, yields are 80-85 percent of normal dents. and plant tissues of corn. The kernels may be segregat-
High-lysine grain can be an important source of ing for various color expressions; and varigation of color
may even be expressed within a kernel, depending upon
high-quality protein in the diets of nonruminants; and
the genetic factors involved and their interaction during
nutritional studies have confirmed the potential value
development of the kernel.
of high-lysine corn in helping to meet the world’s hu- Ornamental corns may be dent, sweet, pop, flint
man and animal nutritional needs. For the present, or floury endosperm types. Apart from genetic studies,
loss of calories per acre is the trade-off for increased they are a curiosity and are only grown for ornamental
amounts of high-quality protein. and decorative purposes.

A publication of the National Corn Handbook Project

and justice for all

Cooperative Extension Service, Iowa State University of Science and The Iowa Cooperative Extension Service programs and policies are
Technology and the United States Department of Agriculture cooperating. consistent with pertinent federal and state laws and regulations on non-
Robert L. Crom, director, Ames, Iowa. Distributed in furtherance of the Acts of discrimination regarding race, color, national origin, religion, sex, age, and
Congress of May 8 and June 30, 1914. handicap.