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World centers of primary diversity of domesticated plants

WORLD CENTRES OF PRIMARY DIVERSITY OF DOMESTICATED PLANTS


Origin of crop plants

The history of crops and man reveals that the evolution of man and the evolution of crop plants went hand in hand. Man has become so utterly dependent on plants such that the plants have 'domesticated' him (Harlan, 1975). The origins of crop plants and their history reveal that the evolution of crop plants is a gradual and slow process rather than sudden and rapid. As such, some cultivated plants differ very little from their original wild relatives, while others differ enormously from their progenitors.

To know precisely about the present day crop plants there needs a thorough understanding of what kinds of plants did man eat before today's crops were available? What were the conditions that prompted him to domesticate the plants? How was his life before agriculture?

To have a rough idea about these questions from the hunter man to the present day civilized agricultural society. Lee and Devore, (1968) cf. Harlan, (1975) put it as “cultural man has been on earth for some 2 million years, for over 99 percent of this period he has lived as a hunter gatherer. Only in the last 10,000 years man has begun to domesticate plants and animals, to use metals and to harness energy sources etc.”

Hunter man was too ignorant or unintelligent to understand the life cycles of plants and they were intellectually insensitive and incapable of 'improvement'.

Gatherers used to collect grass seeds, legumes, roots and tubers, oilseeds, fruits and nuts, vegetables and spices for their food, totaling about 1,400 species. The list suggests that
(i)                 many more species have been gathered from the wild than have ever been domesticated,
(ii)              even after agriculture is fully developed, gathering wild plant foods is still a worthwhile effort, and
(iii)            wild plant resources are of the same general kinds as domesticated plant resources (Harlan, 1975).

GRASS SEEDS like wild rice (Zizania aquatica, Oryza barthii, Oryza /ongistaminata, O. subu/ata, Panicum, Sporobolus, Digitaria, Avena barbata and A. fatua, etc.

LEGUMES include Acacia spp., Canavalia, Vigna, Tephrosia, Phaseolus, Dolichos, etc.

ROOTS AND TUBERS include Dioscorea, Allium, Ipomoea, Cyperus rotundus, Eleocharis, Nymphaea etc.

OIL PLANTS fruits of Arecaceae, E!aeis guineensis, Cocos nucifera; Pistacia, Aleurites, Theobroma (cocoa), Olea, Butyrospermum (Shea butter tree), Sesamum and Lilium.

FRUITS AND NUTS Walnut (Juglans), hickory (Carya), Chestnut (Castanea), Oak (Quercus), pine-nuts (Pinus); Vitis, Ficus, Citrus, Musa, Artocarpus, Annona and Carica etc.

VEGETABLES AND SPICES species of wild Capsicum, Lycopersicon, Nicotiana, Cucumis, Momordica, Luffa, Lagenaria etc.

Thus, the gatherers exploited a wide range of plants wherever they found them. As a result of this practice, independent domestications of different species of the same genus occur and if the genus is wide spread, the different domesticates originate in different continents of the world. Some examples of such wide spread domestications include, Amaranthus, Annona, Cucurbita, Gossypium, Phaseolus, Digitaria, Oryza, Solanum, Dolichos, Ipomoea, Panicum etc.

From the ethaographic evidence (scientific description of races of mankind) it is inferred that gatherers had some knowledge of planting seeds and sowed seeds of wild plants like Chenopodium, Oryzopsis, Eleocharis etc. They also had the knowledge of seed-bed preparation and irrigation to increase production.

Gatherers had the knowledge of how to make use of poisonous foods safe, like seeds of Zamia (a Cycad); certain leguminous and Solanaceous fruits; Dioscorea spp.; Aroid tubers. They also know about drugs, narcotics, medicines, fish poisons, arrow poisons, gums, resins, glues, dyes and paints, bark cloth, woods for spears, arrows, bows, shields, fire sticks and canoes. They had the knowledge of spinn1ng, weaving, basket-making and construction of house hold utensils, fish traps and ceremonial objects.

The Australian aborigins used tobacco for chewing, and another masticatory, Duboisia hopwoodi which contains hyoscyamine, norhyoscyamine and scopolamine.

Agriculture, in mythologies of all civilizations reveal that it is a divine gift to humans as can be seen in Egypt by Goddess Isis, Demeter in Greece and Ceres in Rome. According to the Chinese mythology P’an Ku separated the heaven and the earth, created the sun, moon and stars, and produced plants and animals on earth.

Plant domestication

According to Childe (1952, Cf. Harlan, 1975) the process of plant domestication, by man went through a series of steps like hunter, herder and then a cultivator, the so called Neolithic revolution i.e., a shift from hunting and gathering to food production.

The origin of domestication of plants by man seems to be that the gatherer took up, agriculture or farming since it invo1ves more energy to obtain food by intensive gathering and thought that cultivation is more advantageous than gathering.

World centres of origin of domesticated plants

Milestone work : Two great works, historically, by Alphonse de Candolle and N.I. Vavilov are worth mentioning in any discussion of the origins of cultivated plants.

De Candolle was a French botanist and systematist who produced a voluminous work called the 'Prodromus Systematis Naturalis' Regni Vegetabilis 'Origin of Cultivated Plants (l886) concerning the geography of plants, distributions of wild relatives, history, names, linguistic derivatives, archeology, variation patterns etc. on the cultivated plants.

N. I. Vavilov, a Russian geneticist and agronomist at the National Institute of Plant Industry, USSR, relates to Studies on the Origin of Cultivated Plants published in 1926 and 1951. Vavilov launched an ambitious plant breeding programme that was ever attempted to collect and assemble all of the useful germplasm of all crops. A vigorous, world wide plant exploration, programme was launched and a systematic survey for genetic resources of crop plants was started by him. He was interested in the genetic diversity of crop plants and their centres of origin.

The centre of origin could be determined by an analysis of patterns of variation, and according to Vavilov, the geographic region in which one found the greatest genetic diversity was the center of origin for that particular crop.

From, his extensive studies, Vavilov, proposed eight centers of origin for most of the cultivated plants of the world (Fig.1.1) :

Figure 1.1. Centres of origin for crop plants according to Vavilov (1951). 1. Chinese Centre. 2. Indian Centre. 3. Central Asiatic Centre. 4. Near Eastern Centre. 5. Mediterranian Centre. 6. Abyssinian Centre. 7. South-Mexican and Central American Centre. 8. South American Centre.


On the basis of a large amount of supplementary data we have been enabled to locate more exactly the regions of the origin of cultivated plants. 

Southwestern Asia including Transcaucasia and the northwestern portion of India originated soft wheats and rye as well as many grain Leguminosae, alfalfa, Persian clover, etc. Here, especially in the western part of this area, is the home of the most important fruit trees.

India is the native country of rice, sugar cane and many tropical plants.

The mountains and foothills of Eastern China are the home of many fruit trees, truck crops and the soybean. The vast regions of Central Asia, investigated by us in detail in 1929, have proved alien to the primary process of form origination. In spite of some former botanical suppositions, Central Asia and Siberia have had no influence upon the origin of cultivated plants.

Abyssinia, though economically a country of no particular importance with its cultivated area of only several million acres, shows a striking concentration of the diversity of the genes of wheat, barley and many leguminous grain crops.

Certain countries bordering the Mediterranean are the home of the olive tree, the carol tree, a series of original forage plants and Egyptian clover.

The 7th & 8th centers must be sought in America. In the New World the primary process of form origination is narrowly localized; the regions showing a striking species and varietal diversity occupy comparatively small territories concentrated in Southern Mexico and Central America as well as in Peru and Bolivia. The home of corn and of the upland cotton in all probability is Mexico and Central America, whereas that of the potato is in Peru and Bolivia.

These centers have developed on the basis of an extremely rich wild flora. Here we find conditions especially favorable to the development of species and varietal diversity. These regions have proved equally favorable to civilizations and of course it is no accident that the map showing the distribution of the chief sources of food plants essentially coincides with that of the distribution of the first agricultural civilizations.

The mountain and foothill regions in the subtropics are the most remarkable places for comprehending the evolution of cultivated plants as well as of many wild species.

In these regions the beginnings of the evolutionary process manifest themselves in a salient way especially when we compare the evolution of different species and genera. The existence of such group evolution of different species and genera facilitates greatly an understanding of the evolutionary process.

But it soon became apparent that the pattern is much more complex than what Vavilov has thought of, since some crops do not have centers of diversity. (Fig. 1.2).

According to Harlan (1971) there are three independent systems each with a center and a non center. According to him no single model will explain agricultural origins and he recognizes a humanistic no-model, model to explain the origins of domesticated plants, which is mainly a humanistic problem ‘Man took the initiative in modifying his environment, and plants responded genetically to his activities’. Plant domestication is an evolutionary process operating under the influence of human activities.

Vavilov thought that areas of maximum genetic diversity, represented centers of origin and that the origin of a crop could be identified by the simple procedure of analysing variation patterns and plotting regions where diversity was concentrated. It turned out that centers of diversity are not the same as centers of origin, yet many crops do exhibit centers of diversity. 



FIG. 1.2. Origin and evolution of plants according to Harlan.


The reasons for the origin of secondary centers of crop plants might be due to (Harlan, 1975)
(i)                 A long history of continuous cultivation,
(ii)              Ecological diversity,
(iii)            Human diversity, different tribes are attracted to different races of a crop,
(iv)             Introgression with wild or weedy relatives or between different races of the crop, which leads to hybridization, segregation and selection, and
(v)               The deliberate introduction of certain exotic plants by mail from one continent to another during history.

Germplasm collections of world crop plants are made continuously at the following institutes in the world
§   Food and Agricultural Organizations of the United Nations;
§   The RockfelIer Foundation;
§   The 'Ford Foundation;
§   The Consultative Group;
§   Eucarpia;
§   The United States Development and Agriculture;
§   The Vavilov Institute of the Soviet Union;
§   CSIRO of Australia;
§   The Kihara Institute of Japan;
§   The National Bureau of Plant Genetic Resources of India etc.

Seed storage facilities are available in large quantities at Fort Collins in the United States of America at the National Seed Storage Laboratory. Other places for storage of seed include : Bari (Italy); Braunschweig (West Germany); Izamin- (Turkey); Japan, Bulgaria, Poland, United Kingdom and Australia.

Some of the most important crops in the world are the following: 
(1) Wheat,                               (2) Rice,                                  (3) Maize,
(4) Potato,                               (5) Barley,                               (6) Manioc,    
(7) Oats,                                  (8) Sorghum,                           (9) Soybean,
(10) Cane sugar,                      (11) Beet sugar,                      (12) Citrus,
(13) Cotton fiber,                    (14) Cotton seed,                    (15) Bean, Pea, Chickpea,
(16) Rye,                                 (17) Banana,                           (18) Tomato,              
(19) Millets,                            (20) Sesame,                           (21) Palm oil,
(22) Pea-nut,                           (23) Sweet potato and yams, (24) Coffee,
(25) Tobacco,                          (26) Rubber                             (27) Cocoa and          
(28) Tea.
Process of domestication of crop plants from their wild progenitors

Domestication of cultivated plants is an evolutionary process through human intervention and the process involves a slow and gradual progression from the wild state to incipient domesticated forms or species. But, the process differs from crop to crop, some species evolve directly and some indirectly by a series of steps in their evolution.

Harvesting wild grass seeds was the beginning of domestication by the gatherers. It is always the selection that is associated with harvesting, which causes domestication. Most of the seeds that do not shatter are harvested and most of the seeds that shatter escape the harvest. The shattering character in cereals is simple, one or two gene controlled. Domestication introduces into the crop plants a non-shattering, annual habit with lack of seed dormancy from a wild, shattering, perennial habit and with seed dormancy. This has been well exemplified in the evolution of rice crop (Oryza sativa L.).

There has been a trend in all cereals called the ‘Sunflower effect’ i.e., from many small inflorescences to a few or a single large inflorescence, which is usually accompanied by an increase in the seed size. The head of a commercial cultivar of sunflower, an ear of maize, or a head of modern grain-sorghum or grain-type, pearl millet are strikingly different from their wild progenitors. (Harlan, 1975).

Domestication also introduces a trend towards lower protein and higher carbohydrate content of cereals; increasing seed size and hence endosperm. The embryo is richer in protein and oil but does not increase in the same proportion as the endosperm. This type of selection results in increased seedling vigour.

Cultivated plants have the capacity to evolve rapidly. Rapid evolution is possible only through some variation on the theme of the differentiation-hybridization cycle in which variability already accumulated can be exploited. Mutations play an important role as the sources of variability in crop plants. The crop-weed interaction is the only system by which differentiation-hybridization cycles can be set up in cultivated plants. This enhances variability and broadens the base for plant selection.

Several isolating barriers are known to exist which fragment the populations and are kept genetically apart whereby differentiation occurs. They include: geographic and ecological separation, difference in time of blooming, self-fertilization, and translocation races, polyploid races, gametophytic factors, cryptic chromosomal differences and meiotic irregularities. Differentiation again largely depends upon genetic buffering i.e., the amount of redundancy of genetic information.

Under domestication, changes occur, until the end products are radically different in appearance from their wild progenitors. Thus, domestication results in great   morphological changes without substantial change in the genetic background.



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