Levetin−McMahon: Plants III. Plants As a Source of 11.

Origins of Agriculture © The McGraw−Hill

and Society, Fifth Edition Food Companies, 2008

C H A PT E R OU T L I N E
Foraging Societies and Their Diets 178
Early Foragers 178
Modern Foragers 179
A CLOSER LOOK 11.1 Forensic
Botany 180
Agriculture: Revolution or Evolution? 181
Early Sites of Agriculture 181
The Near East 181
The Far East 182
The New World 183
Characteristics of Domesticated
Plants 184
Centers of Plant Domestication 184
Chapter Summary 185
Review Questions 186
Further Reading 186

K EY C O N CE P T S
1. Early human societies were based on a
foraging lifestyle in which wild plants
were gathered and animals were hunted.
2. Agriculture evolved independently in
several areas of the world, most likely
as a natural consequence of intensified
foraging.
3. The earliest evidence of agriculture
dates back at least 10,000 years in the
Near East, Far East, and Mesoamerica.
4. Domesticated plants are genetically
different from their wild counterparts,
and many can no longer survive without
human intervention.

C H A P T E R

11
ORIGINS OF
AGRICULTURE
Starch grains, which exhibit a characteristic cross in polarized light, can be used to identify
species and distinguish wild from cultivated forms. Starch grains of arrowroot
(Maranta arundinacea) have been found in Ecuador that are 6000 years old. 177
 Levetin−McMahon: Plants III. Plants As a Source of
and Society, Fifth Edition Food
178 UNIT III Plants as a Source of Food
T
he human species, known as Homo sapiens, has
existed for about 250,000 years. For most of that time,
humans survived as foragers or hunter-gatherers,
gathering wild plants and hunting animals in their natural
environment (fig 11.1). Around 10,000 years ago in many
areas of the world, there was a shift in human endeavor from
foraging to farming. Most authorities agree that agriculture
arose independently in different areas over several thousand
years. Why this shift occurred can only be theorized, but the
development of agriculture formed the basis of advanced
civilization in both the Old and the New Worlds. Over the
centuries, agricultural societies spread into those environ-
ments that could be easily adapted to agriculture, and forag-
ers gradually became restricted to marginal areas. By the late
twentieth century, foraging societies had largely disappeared,
constituting only a tiny percentage of the human population
and limited to a few tropical rain forests, deserts, savannas,
tundras, and boreal forests.
FORAGING SOCIETIES
AND THEIR DIETS
Foraging societies are by no means all alike in the types of
food they eat. Some groups, such as the Arctic Inuit, subsist
almost entirely on meat; at the other extreme, the Hadza of
(a)
Figure 11.1 Native North Americans of the Pacific Northwest foraging in the early twentieth century. (a) A Hupa man hunts salmon
with a spear. (b) A Pomo woman uses a beater to gather seeds into a basket.
11. Origins of Agriculture © The McGraw−Hill
Companies, 2008
Tanzania are largely vegetarian, rarely hunting for meat.
Other groups such as the !Kung (the exclamation point is
pronounced as a click) of southern Africa, however, have had
a more varied diet, largely plant based but supplemented by
eggs, insects, fish, small animals, and meat from the hunt.
Many hunter-gatherers have utilized plants by gathering
seeds, flowers, roots, and tubers and have a fairly thorough
knowledge of the botany in the area. From experience, they
know which plants are edible, which are poisonous, which
have medicinal properties, which are sources of dyes, which
could be used for weaving or building materials, and even
which have psychoactive properties. By looking for certain
visible clues such as flowering on so-called calendar plants,
foragers know if tubers are ready to be dug up or if turtles are
laying their eggs. They have developed remarkable methods
to prepare edible foods, even from plants with toxins, such as
cassava, which contains poisonous hydrocyanic acid.
Early Foragers
Archeological investigations have supplied knowledge about
the diet of early humans from many sources, and radiocar-
bon dating of artifacts can provide an estimated time frame.
Fossilized remains of both plants and animals have been
found in early settlements (fig. 11.2). Plants in the diet have
been identified from charred seeds and preserved fruits or
(b)
 Levetin−McMahon: Plants III. Plants As a Source of 11. Origins of Agriculture
and Society, Fifth Edition Food
(a) (b)
Figure 11.2 (a) A clay impression fossil of wheat as compared
with (b) a present-day species of wild wheat.
other plant parts while bones, teeth, feathers, scales, fur, and
shells indicate the animals in the diet. Microscopic remains
include plant fibers, plant crystals, and pollen, with crystals
and pollen especially useful in identification. Coprolites,
fossilized fecal materials, provide direct evidence of the diet
because some plant materials, especially seeds and pollen,
can pass through the digestive tract largely intact. Often
middens, or dump sites, from a human encampment provide
a concentrated source of plant and animal remains. (See A
Closer Look 11.1—Forensic Botany.)
Grinding stones, sickles, and digging implements pro-
vide information on the food in the diet but do not indicate
if the plants harvested were wild or domesticated. Tools of
the hunt also provide some information on the size of the
animals hunted and the method of the kill. Depictions in
cave paintings (of animals as well as hunting and gathering
activities), pottery fragments, and clay figurines are other
windows to the past.
In some excavations at Wadi Kubbaniya in the Nile Valley
of Upper Egypt, archeologists have dated plant remains of a
hunter-gatherer settlement from 17,000 to 18,000 years ago.
The charred remains of fruits, seeds, and tubers from 25 plant
species have been found; interestingly, contemporary forag-
ing societies have utilized these same plants or closely related
species. The most abundant plant remains found were the
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CHAPTER 11 Origins of Agriculture 179
Figure 11.3 A rock painting of the !Kung San of the Kalahari
Desert of southern Africa depicts hunters and prey.
tubers of wild nut grass, a type of sedge. Analysis of modern
samples of these tubers indicates that they are high in carbo-
hydrates and fiber but low in protein; however, the protein is
of good quality because it is high in lysine, one of the essen-
tial amino acids. Mature nut grass tubers also contain toxins,
but these could be easily removed by various methods still in
use today. It is believed that this tuber served as one of the
dietary staples, along with acacia seeds, cattail rhizomes, and
palm fruits. Evidence from this and other Paleolithic (Old
Stone Age or preagricultural societies) sites indicates that
early foraging groups had a remarkably varied plant diet.
Modern Foragers
Much of the knowledge about the foraging way of life has
been documented by studying modern foraging societies such
as the !Kung San of the Kalahari Desert of southern Africa.
These people live in the tropical savannas that border the
Kalahari in what is now southeastern Angola, northeastern
Namibia, and northwestern Botswana. The !Kung have for-
aged in this area for at least 10,000 years, continuing the
hunter-gatherer way of life until recent times (fig. 11.3).
Extensive studies of the !Kung during the 1960s revealed
that they utilized over 100 species of plants and 50 animal
species; two-thirds of their diet was plant based. The plants
 Levetin−McMahon: Plants III. Plants As a Source of 11. Origins of Agriculture © The McGraw−Hill
and Society, Fifth Edition Food Companies, 2008

A CLOSER LOOK 11.1

Forensic Botany

Archeologists have made extensive use of plant remains in

reconstructing the lifestyles of ancient foraging and early
agricultural peoples. The recovering and processing of plant
remains begin the painstaking task of botanical detection.
Not all parts of a plant are equally well preserved: seeds,
wood, pollen, phytoliths, starch grains, and fibers are among
the most informative recoverable remains. The lignified walls
of wood and fibers are more resistant to decay than purely
herbaceous tissue because lignin can be degraded only by a
few types of fungi. Likewise, the lignin, in the sclerified cells
of the seed coat, or testa, makes seeds among the most
common dietary material recovered from archeological
sites. Buried pollen grains are virtually indestructible, owing
to the chemical properties of the exine (outer pollen wall).
Phytoliths (literally “plant stones”) are crystals formed and (a)
found in many plants (box fig. 11.1). They often are created
Figure 11.1a Calcium oxalate crystal in the vacuole of a leaf
within epidermal cells or between plant cells. The crystals
cell, ⫻270.
may be composed of calcium oxalate or, rarely, calcium
carbonate. Another common component of phytoliths is
opaline silica (silicon dioxide). After a plant decays, the phy-
toliths are released into the soil and can remain intact for
thousands of years. Phytoliths are so distinctive in shape and
size that they can be used to identify the presence of a par-
ticular plant family, genus, or even species. In fact, phytoliths
can be used to separate wild species from domesticated
ones and have proven especially useful in dating the origin
of agriculture in many sites.
These same principles of plant detection are applicable
to forensic science, in which the identification of plants can
be used as incriminating evidence. The first criminal case
that used botanical information was the famous 1935 trial of
Bruno Hauptmann, who was accused and convicted of kid-
napping and murdering the young son of Charles and Anne
Morrow Lindbergh. The botanical evidence centered on a
homemade wooden ladder used during the kidnapping and (b)
left at the scene of the crime. After extensive investigation,
plant anatomist Arthur Koehler showed that parts of the Figure 11.1b Needle-like crystals are recognizable in
pineapple pulp.
ladder were made from wooden planks from Hauptmann’s
attic floor. situation, proper medical treatment depends on accurate
In forensics, even herbaceous plant parts can be useful identification of the plant or mushroom. Trained botanists
for identification. In one case, botanical evidence disputed and mycologists (scientists who study fungi) are routinely
the testimony of an accused rapist. Fragments of tree leaves called to hospital emergency rooms for this purpose.
and bark in his pants cuff indicated that the accused had Botanical analyses of stomach contents have also played
climbed a tree to get into a window of the victim’s home, roles in other types of cases: a hunting guide killed a grizzly
rather than being admitted through the front door as he bear he claimed was eating his supply of alfalfa hay. Botanical
claimed. In cases of suspected plant poisonings, identifica- evidence showed no alfalfa in the bear’s stomach, only native
tion can be made from leaves or fruits of intact plants or vegetation. The guide was fined and imprisoned for killing a
even from analysis of stomach contents. In this type of threatened species.

180
 Levetin−McMahon: Plants III. Plants As a Source of 11. Origins of Agriculture
and Society, Fifth Edition Food
included a mixture of fruits, nuts, berries, melons, roots, and
greenery; one particular nut, the mongongo nut, was a very
important high-protein component of their diet. The !Kung
diet was very nutritious; they consumed an average of 2,355
kilocalories per person per day, with 96 grams of protein
and adequate vitamins and minerals. This diet more than
meets the nutritional requirements established for people of
the !Kung stature and physical activity. As in most foraging
societies, the division of labor was along gender lines, with
women doing most of the gathering and men the hunting.
Surprisingly, the amount of time spent on foraging activities
averaged only about 2.5 days per week, which left plenty of
time for leisure and socializing.
Concept Quiz
Humans survived as foragers of wild plants and animals for
hundreds of thousands of years. Agriculture arose relatively
recently, roughly 10,000 years ago, in the course of human
history.
Explain the success of foraging as a survival strategy for early
humans.
AGRICULTURE: REVOLUTION
OR EVOLUTION?
Archeological evidence indicates that about 10,000 years ago
human cultures began the practice of agriculture in several
areas of the world. Over the next few thousand years in the
Near East, the Far East, and Mesoamerica, agriculture flour-
ished. The question that has puzzled archeologists and other
scientists is, “Why, after thousands of years of foraging, did
hunter-gatherers switch to agriculture?”
Many theories have been proposed to answer this ques-
tion. Some state that agriculture was the discovery of a bril-
liant sage who, with a flash of insight, realized that if you sow
seeds, the crop will grow. There were many variations of this
theme. Some held that the brilliant sage realized that plants
growing at middens, or dump sites, were growing from dis-
carded seeds. Others held that the sage observed seeds buried
with the dead (as food for the afterlife) gave rise to plants at
grave sites. These theories viewed agriculture as a revolution;
in fact, the term Agricultural Revolution was used to describe
the transition from foraging to agriculture. It was suggested
that this revolution spread quickly because agriculture was
thought to be an improvement over the hunter-gatherer life-
style in that a dependable food source could be easily grown
rather than collected from the wild.
Beginning in 1960, archeologists questioned that view,
suggesting instead that the origin of agriculture was not
a revolution but the result of a gradual cultural evolution.
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CHAPTER 11 Origins of Agriculture 181
They reasoned that hunter-gatherers knew the wild plants,
knew how they grew, and would incorporate farming along
with foraging as part of an overall food-collection strategy
when necessary. For example, certain aboriginal groups in
coastal Peru abandoned their farming practices whenever fish
became plentiful.
Many archeologists believe that there was a transitional
stage between simple foraging, in which small nomadic
bands followed the wild plants and animals, and agricultural
societies with their sedentary lifestyle. During this transi-
tional stage, foraging groups formed settlements but sent out
members to hunt and gather. This more complex strategy
resulted in changes in the social organization of the groups
and permitted populations to increase. This transitional stage
lasted for several thousand years in some locations until
resource stress or environmental change led to the switch to
agriculture. Archeologists believe that in the Near East, for
example, the climatic dry period around 11,000 years ago
brought about a change in the distribution of cereal grains
(wheat and barley). Applying their botanical knowledge,
these foragers gradually changed from collecting these wild
cereals to cultivating them.
EARLY SITES OF AGRICULTURE
Archeological excavations have documented many sites of
early agriculture in both the Old and New Worlds (fig. 11.4).
The evidence indicates that the earliest sites were in the Far
East, dating back approximately 11,500 years.
The Near East
Some of the oldest sites of agriculture are in southwestern
Asia, in the foothills around the area known as the Fertile
Crescent, which today includes parts of Iran, Iraq, Turkey,
Syria, Lebanon, and Israel (fig. 11.4). From sites such as
Jarmo, Jericho, Ali Kosh, Cayonu, and others, remains of
both plants and animals date back 9,000 to 14,000 years.
Early plant domesticates include einkorn wheat, emmer
wheat, barley, pea, lentil, and vetch while dogs, goats, and
sheep were among the domesticated animals. Research sug-
gests that the animals were domesticated several thousand
years before the plants. Evidence indicates that barley may
have been the first crop domesticated in the Near East,
approximately 10,000 years ago.
One of the sites that had been particularly well studied
was Jarmo in northeastern Iraq, in the foothills of the Zagros
Mountains. This area was inhabited approximately 9,000
years ago; it was a permanent farming village with about 24
mud-walled houses and a population of about 150 people.
The charred grains of wheat and barley found there were a
domesticated type already changed from the wild type; for
this genetic change to have taken place, initial cultivation
must have predated the age of the remains. In addition to the
domesticated plants and animals, the inhabitants of Jarmo
 Levetin−McMahon: Plants III. Plants As a Source of
and Society, Fifth Edition Food
182 UNIT III Plants as a Source of Food
Eastern North
America
Mesoamerica
South American
Highlands
Figure 11.4 Evidence of the beginnings of agriculture have been found in eastern North America, the Tehuacan and Oaxaca valleys in
Mexico (Mesoamerica), the South American highlands, the Fertile Crescent of the Near East, the Yangtze and Yellow River valleys of China
(Far East), and the New Guinea Highlands.
also continued foraging, as evidenced by the bones of wild
animals, snail shells, acorns, and pistachio nuts. Artifacts
uncovered at the site include flint sickles and grinding stones
for harvesting and milling cereal grains as well as clay figu-
rines, woven baskets, and rugs.
The domestication of cats apparently took place earlier
than previously thought. Most authorities placed the domesti-
cation of cats in Egypt around 4,000 years ago. A cat skeleton
discovered on the island of Cyprus indicates that people there
domesticated the cat around 9,500 years ago. The 8-month-
old cat (Felix silvestris) was buried in a small grave next to
a large grave containing a human skeleton. The joint burial
suggests a special association between human and cat. Cats
were most likely valued in agricultural societies to protect
grain stores from mice.
Some researchers question several of the archaeological
findings at some ancient sites of early agriculture. Nicole
Miller, an expert in archaeobotany at the University of
Pennsylvania Museum of Archeology and Anthropology,
studies sites in the Near East for evidence of plants used
by people during ancient times. She documented a 5,000-
year-old site in southwestern Iran that showed an increas-
ing presence of charred wild seeds. In previous analyses,
the presence of wild seeds at the sites of ancient cook fires
was interpreted as spilled food and hence indicated that the
plants were consumed by ancient people. Miller has another
view. She explains that as the surrounding forests were
cleared for fuel and pastureland, the inhabitants turned to
the dung of animals as cooking fuel, and thus, the presence
of charred wild seeds reveals more about the human impact
on the environment than their diet. Similarly, she disagreed
with the conclusion that the diversity of wild seeds pointed
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Companies, 2008
Fertile
Cresent Yangtze and Yellow
River Basins
New Guinea
Highlands
to a varied diet for the inhabitants of an 11,000-year-old
site in Syria. She deduced that the cooking fuel used was
the dung of gazelles, and the wild seeds found were a food
source for an animal that was plentiful at this location in
the past.
The Far East
Excavations of dozens of sites in Asia indicate that agricul-
ture arose at several locations in the Far East. These sites
include the Yellow River and Yangtze River valleys in China
(fig. 11.4). Among the earliest plants domesticated in the Far
East were rice, foxtail millet, broomcorn millet, rape, and
hemp, with evidence of domesticated cattle, pigs, dogs, and
poultry as well.
Current archeological studies in China indicate that
rice cultivation began approximately 11,500 years ago
along the middle reaches of the Yangtze River. From there,
agriculture spread both upstream and downstream. Among
the artifacts examined were samples of rice grains, husks,
plant remains, and impressions of rice grains in pottery.
If these dates are confirmed, this would mean that the
domestication of rice predates the domestication of crops
in the Near East by about 1,500 years. In settlements to the
north, along the Yellow River, foxtail millet was domesti-
cated approximately 8,000 years ago and became a dietary
staple; it remained the dominant crop in North China until
the historical period. Broomcorn millet was also cultivated
but not as extensively. Tilling tools, harvesting tools, and
grain-processing tools made of stone, bone, shell, or wood
have also been recovered from these sites along with bones
of domesticated animals.
 Levetin−McMahon: Plants III. Plants As a Source of 11. Origins of Agriculture
and Society, Fifth Edition Food
Newly discovered evidence strongly indicates that
agriculture may have begun independently in the Papua
New Guinea Highlands nearly 7,000 years ago. The Kuk
Swamp site in the Waghi Valley of central New Guinea
shows several indications of early agriculture. Analysis of
soil sediments for pollen and phytolith microfossils (see A
Closer Look 11.1— Forensic Botany) indicates that the area
was subjected to accelerated forest clearing. Deforestation
led to the establishment of a grass-sedge swampland.
Archeological remains of stakeholes and postholes were
also found and indicate planting, digging, and support-
ing posts, all practices clearly associated with cultivation.
Vestiges of soil mounding and deep channels have been
interpreted as practices to improve drainage in the grassy
wetland. Evidence of taro and banana, historically two of the
most important tropical food crops, have also been found at
Kuk Swamp. The percentage of banana phytoliths is higher
than would be expected in the area and probably indicates
their cultivation. The researchers conclude that this species
of banana, previously thought to have been domesticated
in Southeast Asia, may have instead been domesticated
first in New Guinea and later dispersed to Southeast Asia.
Microfossils of taro, including starch grains and needle-
like calcium oxalate crystals (See A Closer Look 11.1—
Forensic Botany), were recovered from stone tools. Finding
a lowland crop like taro in the highlands confirms that this
crop was deliberately introduced into this site. All evidence
points to New Guinea as one of the primary centers of agri-
culture.
The New World
In contrast to the Old World, the Neolithic (New Stone Age,
or agricultural society) cultures of the New World had domes-
ticated an impressive array of plants but comparatively few
animals. Among the earliest crops domesticated in the New
World were squash, corn, chili peppers, amaranth, avocado,
gourds, beans, and both white and sweet potatoes, with only
dogs, turkeys, llamas, alpacas, guinea pigs, and Muscovy
ducks as domesticated animals. Most of the initial archeologi-
cal evidence for early agriculture in the New World has been
obtained from the highlands of Mexico and Peru.
Probably the most thoroughly documented site is a
group of caves in the Tehuacan Valley of central Mexico.
Research in this area was initiated to obtain information
on the ancestry of corn. Working with ancient corncobs,
archeologists determined that corn had been domesticated
in this region by 5,500 years ago (fig. 11.5). Originally it
was thought that these corncobs were much older (7,000
years old), but newer dating techniques advanced the time
frame. The Tehuacan Valley is one of the few sites where the
transition from foraging to farming can be thoroughly docu-
mented. For thousands of years, people inhabited the caves
seasonally as they foraged for plants and hunted animals. At
first, there was a shift from hunting to a more intensified for-
aging of plants, along with the domestication of squash and
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CHAPTER 11 Origins of Agriculture 183
Figure 11.5 Preserved corncobs from the Tehuacan Valley,
Mexico. The oldest cob, on the left, is approximately 1 inch long.
avocado. Later, the list of domesticated plants expanded to
include corn, bottle gourd, two species of squash, amaranth,
three species of bean, and chili peppers; however, people
still relied on foraging for the majority of their diet. Over
the next few thousand years, agriculture became even more
important as additional plants (tomato, peanut, guava), the
dog, and later the turkey were domesticated. Artifacts such
as stone tools, textiles, and pottery were also found. From
this evidence, archeologists have been able to reconstruct
a picture of the lifestyles of the inhabitants of these caves
over a 12,000-year period. At yet another cave in southern
Mexico, Guilá Naquitz in Oaxaca, seeds and the fruit rind of
a domesticated squash have been dated between 10,000 to
8,000 years ago. This date of domestication is much earlier
than what has been obtained for maize, beans, or any other
New World domesticate. These findings indicate that farm-
ing appeared in the Americas at more or less the same time
that it did in the Fertile Crescent and Asia.
Dolores Piperno of the Smithsonian Tropical Research
Institute has been a leading investigator into the origins of
agriculture in lowland tropical soils of Central and South
America. Using phytolith data (See A Closer Look 11.1—
Forensic Botany), she has discovered that local squash species
were independently domesticated in coastal Ecuador between
9,000 and 10,000 years ago, perhaps slightly predating squash
domestication in the upland regions of Mexico.
Recently, evidence from microfossils has revealed that
chili peppers were domesticated more than 6,000 years ago in
South America. Starch grains of domesticated chili peppers
differ in appearance from those of wild species. The starch
grains of domesticated chili peppers have been preserved on
grinding stones, ceramic shards of cooking vessels, and in
soil sediments from archeological sites in Central America,
South America, and the Bahamas, with the oldest sites in
Ecuador.
 Levetin−McMahon: Plants III. Plants As a Source of 11. Origins of Agriculture © The McGraw−Hill
and Society, Fifth Edition Food Companies, 2008

184 UNIT III Plants as a Source of Food

Much attention has been focused on the early site of

agriculture in the eastern forests of North America. On the
basis of the latest evidence, the eastern half of what is now
the United States and Canada was another New World center
of plant domestication, developing independently at least four
domesticated seed crops: sunflower, marsh elder, goosefoot,
and wild gourd. These farming societies flourished for some
2,000 years before the arrival of maize and other domesti-
cated crops from the Southwest in A.D. 1–200 (fig. 11.4).

CHARACTERISTICS OF
DOMESTICATED PLANTS
Plants that have been domesticated are genetically distinct
from their wild progenitors. Through the process of natural
selection (see Chapter 8), wild plants have evolved mecha-
nisms that ensure their survival in the environment, but once
a plant has been domesticated, traits are artificially selected
to suit human needs and do not necessarily have a survival
value. In fact, some of these traits might be detrimental to sur-
vival in the natural environment. For example, modern corn,
with its ensheathing husks, cannot disperse its seeds; also, Figure 11.6 Two-rowed wild barley (left) is contrasted with
domesticated wheat and other cereals have fruiting heads that six-rowed domesticated barley (right).
are nonshattering, a trait that limits seed dissemination.
Most wild grasses have shattering fruiting heads, which
will break apart at a slight touch or breeze and scatter their
seeds over a wide area. A recessive gene is responsible for a Concept Quiz
tough spike with a nonshattering head. It would be natural Domesticated plants are genetically different from their wild
for early foragers to gather those seeds attached to the tougher relatives.
spikes. When agriculture began, the seeds most easily gath-
ered would be planted and so pass on the nonshattering trait. The fruits of wild cherry trees are sour tasting, yet the
In 2006, researchers pinpointed the exact gene mutation in domesticated varieties derived from wild populations produce
domesticated rice which codes for the nonshattering fruiting deliciously sweet cherries. Explain the difference between wild
head. By comparing the DNA sequences of domesticated and domesticated cherries by explaining the difference between
rice to its wild relatives with the shattering trait, researchers artificial and natural selection.
found a single gene on chromosome 4 with one difference in
the nucleotide bases. The wild rice species had a G (guanine)
which had been replaced by a T (thymine) in domesticated
rice. After the researchers genetically engineered domesticated
rice and restored the G, the shattering trait was also restored. CENTERS OF PLANT
Likewise, early foragers would select for larger seeds,
fruits, or tubers, and over time the domesticated varieties DOMESTICATION
would become larger than their wild counterparts. For exam- Within each area of the world where agriculture evolved, the
ple, wild barley has two rows of grains while the domesti- native peoples developed indigenous crops for a staple food
cated varieties have six rows (fig. 11.6). In fact, archeological supply. Crops that were particularly suitable for agriculture
evidence of six-row barley is indicative of agriculture at that slowly spread to surrounding regions as people traded with
excavation. Loss of seed dormancy is another general char- others or migrated to new areas, bringing their crops with
acteristic of domesticated plants. Seeds of most wild plants them. This diffusion led to the emergence of principle crops
are dormant. For example, wild seeds formed at the end of associated with major centers of the world. In the Near East,
autumn typically do not germinate immediately. Instead ger- wheat and barley were the dietary staples; in the Far East,
mination is delayed via chemical means (hormonal control) rice; in Africa, it was sorghum and millet; in Mesoamerica,
or mechanical means (a tough seed coat) until the next spring corn; and in South America, it was the potato and other root
when the environmental conditions are favorable for seedling crops. As civilization continued to develop, trading and
growth and survival. Plant breeders continue to select for migration expanded the range of crops far from their origin,
desired traits today, using traditional as well as more sophis- and today many crops are even more successful outside their
ticated genetic manipulations (see Chapter 15). native range. Potatoes, which became associated with the
 Levetin−McMahon: Plants III. Plants As a Source of
and Society, Fifth Edition Food
Strawberry
Sunflower
Sugarbeet
Corn
Cotton
Tomato
Dry bean
Tobacco
Dry bean
Potato
Peanut
Strawberry
Figure 11.7 Centers of origin as first defined by N. I Vavilov are indicated by the shaded areas. More recent work has shown that
some crops were domesticated outside these areas or were domesticated independently in several regions.
Irish, are actually native to the highlands of Peru; coffee,
actually native to the mountains of Ethiopia, is most fre-
quently linked to Colombia and Brazil; and the tomato, so
essential to Italian cuisine, was first domesticated in the New
World. The agricultural harvest of the United States would be
meager if limited to commercial crops of native origin such as
blueberries, cranberries, sunflower, pecan, and maple syrup.
Pinpointing the exact origin of important crops has
intrigued scientists for many years. The name most frequently
associated with this endeavor is Nikolai I. Vavilov, a Russian
botanist. Vavilov directed plant-collecting expeditions around
the world and examined thousands of plants, looking for pat-
terns of variation in crop plants and their wild relatives. He
reasoned that areas that had the greatest diversity of a par-
ticular crop would most likely be the center of origin for that
crop. On the basis of his research from 1916 to 1936, Vavilov
proposed eight centers of origin for the major domesticated
plants, six in the Old World and two in the New World.
Examples of crops known to have originated in these centers
are indicated in Figure 11.7. Vavilov’s life ended tragically
in a Soviet gulag. Biology in the Soviet Union under Stalin
was dominated by Trofim Lysenko, who rejected established
genetic theory in favor of his own outdated views. Vavilov’s
adherence to Mendelian genetics clashed with the established
policies of the Soviet state, and he was sentenced to a Soviet
prison camp where he died in 1943, a martyr to the cause of
scientific freedom.
11. Origins of Agriculture © The McGraw−Hill
Companies, 2008
CHAPTER 11 Origins of Agriculture 185
Rye
Almond
Grape Apple
Soybean
Rice
Orange
Alfalfa
Sorghum Cotton
Sugarcane
Barley
Onion
Wheat
Recent work has expanded the number of centers and
questioned Vavilov’s conclusions. Evidence suggests that
although some crops have been domesticated more than once
in different places, others did not originate where Vavilov
indicated, and still others were developed over vast regions.
For example, certain New World crops such as cotton and cas-
sava appear to have been independently domesticated in both
Mesoamerica and South America. This search for the origin of
certain crops is even more important today as plant geneticists
strive to improve the gene pool of domesticated plants by tap-
ping the genetic resources of wild strains (see Chapter 15).
CHAPTER SUMMARY
1. The first human societies were based on a foraging life-
style, gathering wild plants and hunting animals for food.
Archeological investigations have determined that the
diet of Stone Age foragers was varied, especially in the
variety of plants consumed. Studies on the diet and life-
style of extant foragers, especially the !Kung San of the
Kalahari Desert, reinforce the viewpoint that the foraging
lifestyle more than satisfies the nutritional requirements
yet allows time for activities not directed to food gather-
ing and preparation.
2. The archeological record indicates that, at least in some
parts of the world, certain groups began to shift from
the nomadic, foraging lifestyle to the sedentary one of
 Levetin−McMahon: Plants III. Plants As a Source of
and Society, Fifth Edition Food
186 UNIT III Plants as a Source of Food
agriculture. Many theories to explain this shift have been
presented and discarded over the years, but the currently
held view believes that the switch to agriculture was
gradual. A prolonged transitional stage ensued in which
groups formed settlements but sent out members to hunt
and gather.
3. The earliest agricultural settlements, approximately
11,500 years old, have been found in the Far East along
the Yellow and Yangtze River valleys. Sites in the Near
East, in an area known as the Fertile Crescent, also docu-
ment early agriculture. The New World dates from the
Tehuacan and Oaxaca valleys of Mexico show that agri-
culture started at approximately the same time.
4. Domesticated plants and animals are genetically different
from their wild relatives because they have been shaped
by artificial selection. Many of the traits, such as nonshat-
tering fruiting heads, do not enhance a plant’s survival
value but have been selected to suit humanity’s needs.
5. Because domesticated plants have been modified greatly
from their wild ancestors after thousands of years of
artificial selection, it has often been difficult to pinpoint
their area of origin. Nikolai I. Vavilov laid the foundation
for detecting the centers of origin of domesticated plants
when he proposed eight centers, six in the Old World and
two in the New.
REVIEW QUESTIONS
1. What has been learned about foraging societies of the past
by studying the !Kung?
2. Describe an early agricultural community in the Near
East.
3. How do archeologists reconstruct diets of prehistoric
peoples?
4. How have the theories about the origin of agriculture
changed in recent decades?
5. What crops were domesticated in the New World?
6. You are an archeologist on a dig and discover in a cave
the preserved seeds and fruits of a type of squash. These
remains are dated to between 8,400 and 10,000 years old.
What characteristics would you look for to determine
whether this squash was a wild type, gathered by forag-
ing, or a domesticated plant, cultivated in the fields?
7. A national seed company offers a reward of $10,000 to
the first gardener who develops a pure white chrysanthe-
mum. How would you go about breeding a chrysanthe-
mum for its color? What is the underlying process and its
mechanism of action?
11. Origins of Agriculture © The McGraw−Hill
Companies, 2008
FURTHER READING
Balter, Michael. 2007. Seeking Agriculture’s Ancient Roots.
Science 316 (5833): 1830–1835.
Braidwood, Robert J. 1960. The Agricultural Revolution.
Scientific American 203(3): 130–148.
Bryant, Vaughn M., and Glenna Williams-Dean. 1975. The
Coprolites of Man. Scientific American 232(1): 100–109.
Diamond, Jared. 1998. Guns, Germs, and Steel: The Fates of
Human Societies. W. W. Norton, New York, NY.
Diamond, Jared and Peter Bellwood. 2003. Farmers and Their
Languages: The First Expansions. Science 300: 597–603.
Fritz, Gayle J. 1995. New Dates and Data on Early Agriculture:
The Legacy of Complex Hunter-Gatherers. Annals of the
Missouri Botanical Garden 82(1): 3–15.
Gilbert, Robert I., and James H. Mielke. 1985. The Analysis
of Prehistoric Diets. Academic Press, Orlando, FL.
Harlan, Jack R. 1976. The Plants and Animals That Nourish
Man. Scientific American 235(3): 88–97.
Harlan, Jack R. 1992. Crops and Man, 2nd Edition. American
Society of Agronomy. Madison, WI
Harris, David R., and Gordan C. Hillman, eds. 1989. Foraging
and Farming: The Evolution of Plant Exploitation.Unwin
Hyman, London.
Heiser, Jr. Charles B. 1979. The Gourd Book. University of
Oklahoma Press, Norman, OK.
Lane, Meredith A., Loran C. Anderson, Theodore M. Barkley,
Jane H. Bock, Ernest M. Gifford, David W. Hall, David
O. Norris, Thomas L. Rost, and William L. Stern. 1990.
Forensic Botany. BioScience 40(1): 34–39.
MacNeish, Richard S. 1964. The Origins of New World
Civilization. Scientific American 211(5): 29–37.
Nash, Steve. 2004. Seeds of Time. Archaeology 57(1): 24–29.
Piperno, Dolores, and Deborah Pearsall. 1998. The Origins of
Agriculture in the Lowland Neotropics.Academic Press,
New York, NY.
Pringle, Heather. 1998. The Slow Birth of Agriculture.
Science 282: 1446–1450.
Smith, Bruce D. 1995. The Emergence of Agriculture. Scientific
American Library. W. H. Freeman and Co., New York, NY.
Yellen, John E. 1990. The Transformation of the Kalahari
!Kung. Scientific American 262(4): 96–105.
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