REVIEW ARTICLES

Origin of agriculture and domestication ofplants and animals linked to early Holocene
climate amelioration
Anil K. Gupta
Department of Geology and Geophysics, Indian Institute of Technology, Kharagpur 721 302, India

Domestication of plants and animals was necessary
for the evolution of agriculture, spatial expansion andpopulation increase of humans during the Holocene,
which facilitated the evolution of technically innovative
societies. The agricultural practices enabled people toestablish permanent settlements and expand urbanbased societies. Domestication of plants and animals
transformed the profession of the early humans fromhunting and gathering to selective hunting, herding
and settled agriculture. The earliest archaeologicalevidences, found throughout the tropical and subtropical areas of southwestern and southern Asia, northernand central Africa and Central America, suggest rapid
and large-scale domestication of plants and animals
ca. 10,000–7000 cal years BP. This interval corresponds
to an intense humid phase and equable climates, as observed in numerous paleo records across the regions. I
suggest that domestication of plants and animals andsubsequent beginning of agriculture were linked to climate amelioration in the early Holocene.
CLIMATE has played a critical role in the evolution of
fauna and flora. Proxy data suggest occurrence of widespread repeated, abrupt climate changes throughout thegeological record1. Persistent abrupt changes in climate mayalter ecological landscape, leading to faunal adaptation.
Human adaptation and migration in response to severe climate changes are known from the paleo records2. Recent
archaeological findings from Yana River, Siberia, indicatethat humans adapted to the harsh, frigid climate of the
Arctic during the late Pleistocene about 27,000 year ago3.
Likewise, they adapted to arid conditions in the Thar4 andSahara Deserts5 in the late Holocene. Although some human societies have adapted to rapid climate changes in
the past, there are also examples of civilizational collapse
under the persistent influence of climate change. The collapse of the Akkadian, Classic Maya, Mochica and Tiwanaku
was related to persistent multi-century shifts in climate6,7.
The rise and fall of human civilizations in South Asia havealso been climatically mediated8. The effect of persistent
abrupt climate changes has been equally drastic on thee-mail: anilg@gg.iitkgp.ernet.in
54

flora, that brought shifts from one type to another type ofvegetation in the past geological records9.
The anthropogenic pressures in response to climatechange have brought significant changes in fauna andflora. The most significant examples of human impact on
the evolution of ecological niches come from domestication of animals and plants. Domestication refers to theprocess of reciprocation, by which animal and plant species come to depend on humans for survival, while providing humans with numerous benefits in turn. The processof domestication has been markedly important for spatial
expansion and population increase of humans during the
Holocene. Domesticated plants and animals are of prime
importance for agriculture. Without agriculture, the complex, technically innovative societies and large humanpopulations that exist today could not have evolved. Agriculture allowed people to become sedentary (living for a
prolonged period in one place), establish permanent villages and towns and develop classified societies that included specialized and dedicated segments such as farmers,
artisans, soldiers, religious leaders, teachers and governors.
Agriculture is thus defined as the cultivation of domesticated plants and animals for use by the human societies,
as many domesticated plants and animals would not survive without human intervention10.
Early hominids were hunters and gatherers who reliedon naturally occurring vegetation, fruits, nuts, carrion andgame for subsistence. Hunters and gatherers do not establish permanent settlements such as villages. They movetheir camps in response to changes in the season and climate. Subsistence by hunting and gathering is still practised by ancient tribes throughout the world. As the climate
ameliorated, there was an increase in vegetation and peoplechanged their food habits and lifestyle. Dogs were thefirst to have been domesticated by early humans (Table
1)10, which might have been used for hunting of wild animals. Goats and sheep were next to have been domesticated in southern and southwestern Asia during the early
Holocene8,10,11. The remains of sheep and goats found at
hominid sites that are older than about 10,000 cal years show
no evidence of domestication10, suggesting that hunterswere killing the whole population of goats and sheep at thattime. However, there was a shift towards greater abundance of skeletons of younger goat and sheep at hominid
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sites after about 10,000 years ago10. This suggests that
immediately after 10,000 cal years, people were keeping
herds of these animals and by 9000 cal years, sheep andgoat were being kept and raised by people at locations
away from the natural realms of wild goats and sheep10.
Thus, it is believed that human interaction with domesticated herding animals transformed them from hunting
to selective hunting and finally to herding. Over the past
10,000 years, many animals have been domesticated indifferent parts of the world, as known through reasonablearchaeological evidences on the early period of domestication (Table 1)12. For example, pigs and cattle were domesticated in southwestern Asia about 9000–7000 cal years
ago. The horse and bactrian camel were domesticated incentral Asia between 5000 and 3500 cal years ago, whereas
water buffalo was domesticated in India and southeastern
Asia ~ 4500 cal years ago12. In areas such as central Eurasia, parts of the Near East, northern Eurasia and parts ofNorth Africa, human cultures developed, which were organized around herding domesticated animals such as sheep,
horses, cattle, camels and reindeer. These people supplemented the meat and milk obtained from their herds withhunting and gathering. The human herders migrated andmoved along with their herds during seasonal changes ingrazing areas – the so-called pastoral nomadicism10.
The domestication of plants and animals is believed tohave begun in the tropical and subtropical areas in the early
half of the Holocene (Figure 1). Tropics provide enoughmoisture for the growth of vegetation that provides foodto both humans and animals. Some of the earliest evidences for the domestication of plants come from southwestern
Asia, dating back to the same period as the domesticationof goats and sheep. A variety of food plants, including
wheat, rye, barley, lentils, chickpeas and peas were all
domesticated in the Near East and South Asia about 11,000
to 9000 cal years ago10,11. In southwestern Asia, the archaeological sites with evidence of early plant domestication

Table 1.

occur along an inverted crescent, the Fertile Crescent (Figure 1, Table 1), extending from the mouth of Tigris andEuphrates Rivers, north to eastern Turkey and then south
to the coastal regions of Lebanon and Israel12. One of theearliest evidences of domesticated plant cultivation anduse comes from a site along the Euphrates River in Iraq.
Although numerous important domestic species come fromsouthwestern Asia, people around the world domesticatedwild plants12. For example, corn domestication occurredin Central America and spread to South America. Likewise, domesticated Asian rice is from India and China13
and foxtail millet and cabbage came from China10.

Origin and expansion of agriculture and domestication of animals
Two questions regarding domestication and agriculture
expansion have long hounded archaeologists, paleoecologists and paleoclimatologists. These are: (1) why did hunting and gathering people turn to agriculture and domestication of animals? and (2) what was the spatial pattern ofagricultural origin and expansion? The cause for people
to abandon a hunting and gathering lifestyle and take up
agriculture is not that clear. Experiments suggest that it
was most likely because of the more energy required for
hunting and gathering, than for agricultural practices toobtain the same calories of food energy (e.g. MacDonald10,
p. 357). The external pressure from environmental change
and intrinsic dynamics of human populations, particularlyin the wake of pressure from population growth, are theother factors proposed to explain how and why huntersand gatherers turned to agriculture10. But this hypothesis
is contradictory, because the early Holocene climate conditions were more conducive with enough rain and riverwater available throughout the Asian-African region5,14–16
that might have caused the origination and expansion ofagriculture and not the environmental pressure (Figure 2,

Areas and minimum ages for domestication of selected animals and plants around the world(from Simmons12; Allchin and Allchin11). For location of areas, please see Figure 1.

Areas with minimum ages
Pan Po (3750 BC)
Indian subcontinent (9000 BC)
Central Asia (3000 BC)
Ali Kosh (7500 BC)
Jarmo (6700 BC)
Egypt (3000 BC)
Jericho (7000 BC)
Palegawra cave (12000 BC)
Shanidar (9000 BC)
Cayönü (7000 BC)
Catal Hüyük (6500 BC)
Franchthi (5000 BC)
Argissa (6500 BC)
Ayacucho (6000 BC)
Tehuacan (6000 BC)
Ocampo Cave (7000 BC)
Highland South America (2000 BC)
CURRENT SCIENCE, VOL. 87, NO. 1, 10 JULY 2004

Domesticated plants, animals
Foxtail millet, cabbage
Wheat, barley, jujube, sheep, goat (water buffalo: 2500 BC)
Horse (bactrian camel: 1500 BC)
Sheep, goat? (eincorn, emmer: 7000 BC)
Einkorn, emmer, barley, lentil, pea
DonkeyEinkorn, emmer, barley, pea, lentil
Dog
Sheep?
Einkorn, pea, lentil, sheep, pig, goat?
CattleSheep, goat, pig, cattle
CattleLima bean, common bean
Maize?, squash, gourds (Maize: 5000 BC)
Squash?, gourd?, scarlet runner bean (Common bean: 4000 BC)
Guinea pig (llama, alpaca: 1500 BC)
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REVIEW ARTICLESTable 1). I thus suggest that it was climate amelioration
during the early Holocene (10,000–7000 cal years BP) andnot the environment pressure that triggered the domestication of animals and plants throughout the tropics andsubtropics. The favourable climate conditions with enoughrains brought significant change in the ecology, leading
to the exuberance of vegetation and diversification of theplant community. This hypothesis thus also opposes V. G.
Childe’s Oasis Theory17 that envisions a humid phaseduring the Last Glacial Maximum and a dry phase in theearly Holocene.
Evidences indicate that floral taxonomic diversity decreased at the peak of the Last Glacial Maximum (LGM),
followed by an increase towards the beginning of the
Holocene and a dramatic change in the assemblage during the Holocene18. As noted earlier, several independent
trajectories of subsistence intensification, often leading toagriculture, began during the Holocene. That agriculture
did not start during the Pleistocene can be explained fromthe fact that last glacial climates were extremely unfavourable to agriculture19, being dry, low in atmosphericCO2, and extremely variable on short time scales. As theclimate ameliorated, domestication followed. Indeed, inthe Holocene, agriculture may have become necessary forthe expansion of humans19.
The increased greenery and diverse vegetation during
the early Holocene enhanced man’s awareness and sensitiveness towards nature. Humans became more curious tolearn about the traits and usefulness of plants and vegetation for their subsistence. The availability of stone andmetal tools helpful in clearing fields and crop harvesting

may have also helped the domestication of wild plants
and beginning of agriculture in the early Holocene. Initially, animals like sheep and goats were domesticated for
milk, meat and wool. Subsequent domestication of larger
animals was probably for their use in agriculture, irrigation and conveyance.
Despite the well-established ability of humans to domesticate plants and animals and to practice agriculture,both archaeological and genetic evidence from crop
plants show that certain areas of the world were centresof domestication for the early development of agriculture(Figure 1, Table 1). Other areas either developed agriculture after contact with people and crops from one of thesecentres or they never practised it at all. Notable centres of
initial plant domestication and agricultural development
include southwestern Asia, a large part of central Africa,
India, eastern Asia and Central and South Americas.
People of northern Europe, North America and Australiapractised agriculture only after it was brought from elsewhere10,12. The origin of agriculture was linked to theavailability of wild plants and animals that were usefulfor domestication. The Fertile Crescent of southwestern
Asia and the Indian subcontinent offered many varieties
of wild plants and animals, which were ideal for domestication (Figure 1).

Evidence from the Indian subcontinent
The Indian subcontinent has been one of the earliest centres for animal and plant domestication, owing to suitableclimate conditions during the early Holocene. The region

Figure 1. Important centres of domestication and agricultural origins (from Simmons12; Table 1). Fertile Crescent(yellow-shaded region) and parts of South and Central Asia appear to be the centres of earliest domestication of plants
and animals that led to the beginning of agricultural practices.
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is characterized by one of the most fascinating climate features of the Earth – the South Asian or Indian Ocean monsoon system that influences the socio-economic life ofpeople of the region15,20. The seasonal changes and decadal to century scale shifts in the monsoon during the Holocene influenced plant population and diversification. For
instance, vegetational changes near Gangotri Himalayas
have been linked to changes in the monsoonal precipitation during the past 2000 years8,21.
The relation between humans and animals in India has
been very friendly and complementary since the beginning of animal domestication, which can be realized from

a

b

c

d

e

Figure 2. Holocene record of climate variability from different regionsof the tropics and subtropics. a, Ti (%) from Cariaco Basin25; b, Terrigenous flux at ODP Site 658, NW Africa5; c, Globigerina bulloides(%) from ODP Site 723, Arabian Sea15; d, Oxygen isotope record fromsouthern Oman16; e, West Pacific26. Shaded area marks the early Holocene humid phase (~ 10,000–7000 cal yrs BP) corresponding to thebeginning of agriculture and domestication of plants and animals in thetropical and subtropical regions.
CURRENT SCIENCE, VOL. 87, NO. 1, 10 JULY 2004

the fact that many Indian sages turned wild animals as
their pets since the Vedic times. Worshipping of different
components of nature (both animals and plants) by Indianssince the Vedic times indicates a deep love and awarenessof Indians towards nature, which they thought important
for their survival. In fact, all ancient civilizations throughout the world were nature worshippers. An excellent example of human–animal relation in India is the domesticationof Asian elephant long ago, whereas the African elephantstill lives a wild life. The relation between plants and people has been equally intimate for people of India. Pipaland barh (banyan) – the two sacred trees to Hindus, foundthroughout the Ganga–Yamuna doab, are some of the oldest wild trees worshipped in India since the ancient times.
Vedic Indians realized the importance of plants for
subsistence as well as for medicinal use (herbs) that
attracted them to undertake exhaustive research on theimportance of wild plants in domestic and medicinal use andas a result, the Science of Indian medicine, the Ayurveda,
developed.
Archaeological evidences from the Indian subcontinent
indicate that the domestication of animals and plants beganaround 10,000–7000 cal years BP in the northwestern partof India, in the region watered by the Indus River and its
tributaries11. Paleomonsoon records from marine5,15,22–26
and land sequences14,16 indicate that the early Holocene(~ 10,000–7000 years BP) was an interval of warmer andwetter conditions with intensified southwest (SW) monsoon(Figure 1). Ganga, Brahmaputra as well as Indus Rivers
were flowing in their full strength during this time14,27–29.
Mehrgarh (now in Pakistan) with lush green vegetation
was suited to be the centre for early civilization in the
Indian subcontinent and for the transformation from hunting and gathering communities to communities with settled agriculture and domesticated animals11. The Bolan
valley with abundant vegetation including wild cereals,presence of wild animals and water availability might haveattracted the earlier man, even before the evolution ofagriculture. The early settlement at Mehrgarh dates backto ~ 9000 cal years BP30–31, which presents the oldest evidence so far for the beginning of agriculture and domestication of animals in the Indus system, coinciding withthe humid phase of the SW monsoon15,16. Mehrgarh provides an important evidence for the change from hunting,gathering and pastoralism to a subsistence economy, centered around settled agriculture and the domestication ofwild animals11. Agriculture might have allowed people tobecome sedentary, establish permanent villages and towns
and become aware of the social system of the societies.
Wheat and barley were cultivated from the very beginning of agriculture and among the fruits, jujube, stones ofdate palm and grape seeds were first used32. The spread of
agriculture thus was essentially the expansion of the highly
successful pattern of wheat and barley production anddomestication of cattle, sheep and goat that emerged at
Mehrgarh area11,33. The increase in rainfall in the Indian
subcontinent must have created conditions favourable for57

REVIEW ARTICLES
the expansion of agriculture throughout the Indus Basinand other areas as well. Thus, the early Holocene interval(10,000–7000 cal yrs BP) was marked by a subsistence
economy, based on the cultivation of wheat and barley andsettled civilization.
It is likely that the summer monsoon season was longer
in the early Holocene34 with excess moisture content thanwhat was required for food production. This might haveprovided enough moisture for the winter monsoon rainsneeded for winter crops. Since both wheat and barley are
winter (Rabi) crops, it seems probable that grain production would initially have depended mainly upon winter rains
and subsequently, people had developed methods of irrigation and rainwater harvesting as an adaptation to monsoon variability2,35. The cultivation of rice was probably
introduced during the early Holocene humid phase in the
Ganga–Yamuna Doab (interfluve)11. The excess summer
monsoon rain in the early Holocene probably did not allow
early farmers of the Indus region to grow Kharif (rainyseason) crops like maize, millet (jawar, bajra) and a variety of lentils, which were probably introduced later withthe beginning of arid phase about 4000 cal years BP25,36–
37. In summary, the strong monsoon of the early Holocene
appears to have been wet enough for agriculture during thewinter season, while the late Holocene (last 4000 years) was
drier, coinciding with the appearance of rainy season crops.
Aridity too had considerable influence on the subsequent
diversification of crops and their cultivars. The cultivationof millet in southern India may have been in response toaridity. There is evidence that small millets Brachiaria
ramosa and Setaria glauca were domesticated in India. A
study by Kimata et al.38 suggests that domestication process may have passed through three distinct phases: (i) art
associated mimic weed with upland rice and some millets; (ii) a secondary crop mixed with little millet or kodo
millet; and (iii) as an independent crop. Indeed, this phased process has been promoted as a means of adaptation
to aridity and gave rise to further diversification of thecrop pattern.

Evidence from China
China provides significant evidence of climatic effects onhumans and their farming systems during the Holocene.
Several human–induced changes in vegetation have occurred during the Late Quaternary in the Middle Yangtze Rivervalley, where one stock of rice was domesticated39. Fossil
rice phytoliths identified from the late glacial to Holocene
sediments in the East China Sea, were probably transported by the Yangtze River. The rice phytoliths are foundfirst in the sequence datable to ~ 13,900 cal yr BP and disappeared during the period of 13,000–10,000 cal yr BP,
coinciding with the last glaciation, the so-called ‘Younger
Dryas’40. The coincident disappearance of domesticatedrice phytoliths with cold climate conditions suggests animmense influence of climate on human actions during that58

time. Warmer and wetter conditions during the period 13,900
to 13,000 cal yr BP and after 10,000 cal yr BP are most likely
to have favoured rice domestication in the area41.
Climate thus has influenced vegetation and societies
throughout the tropics and other regions, leading to thedevelopment of different civilizations. The abrupt and dramatic changes in climate during the Holocene caused humanmigrations in many areas, whereas some societies adapted
other means of sustenance. In India’s Thar Desert, people
used rainwater harvesting as an adaptation to monsoon
failures. The rising global temperature due to increase ingreenhouse gases could be a cause of worry for climate
modelers, as well as policy planners, because, as the temperature rises, the climate undergoes more dramatic changes
with droughts in some areas and floods in other areas.
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ACKNOWLEDGEMENTS. I thank D. N. Pandey for valuable comments of an earlier version of this manuscript. Support by R. K. Singhand Moumita Das in making plots is appreciated.
Received 23 January 2004; revised accepted 28 April 2004

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