| Independent Domestication of Beans and Cotton
Sam Cox December, 2000
Domestication origins are often difficult to locate, since most domestications
of modern crops took place several thousand years ago, and archaeological
evidence is difficult to come by, or difficult to interpret when found.
In addition, crops have undergone constant change since domestication,
thus, an ancient domesticated crop is often difficult to distinguish from
its wild progenitor in fossil records until long after domestication, when
the crop has attained significant morphological uniqueness. Archaeological
remains of crops are infrequent, and strongly associated with certain climates
which preserve plant material better. For these reasons, it is difficult
to conclude definitively that any crop had more than one domestication
center since the alternative, one domestication with subsequent diffusion,
is difficult to disprove. Nevertheless, strong evidence for multiple domestication
of some crops exists, and one can conclude with only minor doubt that some
crops did indeed undergo domestication in different areas at different
times.
BEAN
Strong evidence supports the theory that beans have been independently
domesticated one or more times. Four species of beans have been domesticated
form the thirty or so wild species of South and Central America. These
are Phaseolus vulgaris, P. lunatis, P. acutifolius and P. coccineus,
the first two being the prime suspects of multiple domestications (Kaplan,
1965; Harlan, 1971; Heiser, 1965; Evans, 1976)).
Examinations of the wild Phaseolus species form which domestications
likely occurred revealed that these wild populations were often morphologically
distinct, geographically isolated, and in some cases non-interfertile,
as in beans from Mesoamerica and South America (Gepts, 1993). Wild beans
are members of the species P. aborigineus. Grouping all these populations
under one species is technically questionable, but has been long the standard,
and is unlikely to change under new evidence (Sauer, 1993).
Strong evidence for multiple domestication of wild progenitors involves
a seed storage protein called phaseolin, which has eight different genetically
controlled types among wild populations (Gepts, 1993). It is significant
that among domesticates, six of these protein types are present. Some take
this as a sign of multiple domestication events (Harlan, 1992). It is unlikely,
given the geographical isolation of the wild populations, that one domestication
incorporated that many protein types, and is only slightly less unlikely
that one people domesticated multiple types. The dominant presence of some
of these protein types largely rules out introgression from wild species
since introgressed genes would be present only in isolated instances instead
of widely in the general population, as in the rare Columbian B-type protein
(Blumler, 1992). Additionally, beans only rarely cross pollinate, so introgression
from wild species should not be a significant source of gene flow. The
majority of bean types contain either the S-type, found in beans from Mesoamerica,
or the T-type, found in beans from South America (Sauer, 1993).
Mulitple domestication theory of beans received a boost in 1996 with
the discovery of differential presence of the fin gene between accessions
from South America and Mesoamerica. The fin gene causes a dwarf growth
habit, as opposed to typical climbing habit. It reduces the number of nodes,
time to maturity, and height of the plant by transforming apical vegetative
meristems into terminal reproductive meristems (Koinange, et al. 1996).
Traditional bean cultivation in Mexico involved beans climbing up corn
stalks. The absence of corn, at least early on, in Peruvian culture would
make a bush bean preferable for cultivation.
Arhcaeological evidence also supports at least two independent domestications
of beans. P. vulgaris was present in both Peru and Mexico by 5000
BC. P. lunatis was present in Peru by 6500 BC and in Mexico by 800
AD. The presence of P. vulgaris in both locations at the same time
strongly supports independent domestication. The later appearance of P.
lunatis in Mexico is not as convincing for independant domestication,
since certainly 6000 years is long enough for diffusion to take place.
However, beans found in archaeological sites from South America have consistently
larger seeds than those found in Mexico. This difference in seed size suggests
different domesticates.
Isozyme and morphological traits also support independent gene pools
stemming from imdependent domestications (Gepts, 1993).
In summary, at least two independent domestications of beans have taken
place; one in South America and one in Central America. Possibly more than
one domestication took place in South America. The evidence for this includes
differences in isozymes, phaseolin protein types, morphology, geography
of wild ancestors and the presence of the fin gene. Archaeological remains
also support independent domestication.
COTTON
An important fiber crop for several civilizations, cotton was undoubtedly
domesticated multiple times. Very little argument on this point can be
made since domesticated forms were found on both sides of the Atlantic
much earlier than any known trans-Atlantic travel. However, independent
domestication has even been suggested in the New World, based on several
lines of evidence (Pickersgill, 1977). Altogether, cotton has likely been
domesticated four times around the world (Sauer, 1993).
There are four domesticated species of cotton. Prior to the relatively
recent expansion of cotton production into temperate zones, all cultivars
were perrenial shrubs or trees (Harlan, 1992). Only species possessing
the A genome produce filamentous fibers used for textiles.
East of the Atlantic, the diploid (AA) Gossypium herbaceum was
most likely domesticated in Ethiopia. The exact date is not known, but
Theophrastus recorded seeing a "wool bearing tree", which was probably
cotton, in 350 B.C. G. arboreum, another AA diploid, seems to have been
domesticated in India, where the current center of diversity is. This cotton
was more widely used than G. herbaceum in the Old World prior to
trans-Atlantic travel.
The New World has four wild tetraploid species (AADD), two of which
have been domesticated. Molecular evidence shows that all of the tetraploid
cottons contain the same A genome, suggesting that the A genome was incorporated
into cotton a very long time ago, perhaps 1-2 million years ago, with subsequent
speciation into four distinct groups (Gepts, 1993). Diploid New World cottons
do not contain the A genome (Harlan, 1992). The A genome is present, in
more or less the same form, on both sides of the Atlantic (Sauer, 1993).
This has caused some consternation about the source of the A genome, and
possible transport across the Atlantic. The most plausible explanation
is that the A genome is very ancient, and probably occupied a range on
the ancient continent of Pangea over much of current South America, Africa
and India (Harlan, 1992). The immediate ancestor of either cotton species
is not definitively known, and may be extinct (Gepts, 1993)
Evidence of multiple New World domestication is primarily archeaological.
G. barbadense was domesticated in South America. Remains of this
cotton were found in a site dated to 2500 B.C. G. hirsutem was probably
domesticated in Mexico, since definitive remains of domesticated cotton
have been found at a site dated to 3500 B.C. Both domestication centers
are centers of diversity for their respective species (Gepts, 1993). The
early presence of both domesticated species in geographically isolated
areas strongly suggests independent domestication. Although the two species
are morphologically similar, they are easily distinguished by isozyme alleles
(Gepts, 1993).
Given the evidence, it can safely be concluded that cotton was domesticated
four times in India, Africa, Mexico and South America.
References
Blumler MA. 1992. Independent inventionism and recent genetic evidence
on plant domestication. Economic Botany. 46(1):98-111.
Evans AM. 1976. Beans. In: Evolution of Crop Plants. Edited by: NW Simmonds,
et al. Longman; London. pp 168-172
Gepts P. The use of molecular and biochemical markers in crop evolution
studies. In: Evolutionary Biology, V27.1993 Edited by: MK Hecht, et al.
Plenum Press, NY, NY.
Harlan JR. 1992. Crops and Man, 2ed. Ameican Society of Agronomy, Inc.;
Madison, WI.
Harlan JR. 1971. Agricultural origins: Centers and Noncenters. Science
174:468-474.
Heiser CB. 1965. Cultivated plants and cultural diffusion in nuclear
America. Am. Anthropl. 67:930-949.
Kaplan L. 1965. Archaeology and domestication in American Phaseolus.
Econ. Bot. 19:358-368.
Koinange EMK, Singh SP, Gepts P. 1996. Genetic control of the domestication
syndrome in common bean. Crop Sci.36:1037-1045.
Sauer JD. 1993. Historical Geography of Crop Plants. CRC Press; Boca
Raton, FL. pp73-80.
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