An Interdisciplinary Perspective on the Origin of Maize

Mary W. Eubanks

Latin American Antiquity, Vol. 12, No. 1. (Mar., 2001), pp. 91-98.

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 AN INTERDISCIPLINARY PERSPECTIVE ON THE ORIGIN OF MAIZE

Mary W. Eubanks

This paper addresses objections raised in an article by Bennetzen et al. (2000) in response to MacNeish and Euhanks (2000).
Bennetzen et al. interpret the findings reported by MacNeish ar~dEuhanks as opposition to the teosinte hypothesis for the ori-
gin of maize. However, by demonstrating a mutagenic mechanism that could have generated the genetic diversity essential for
the transition from teosinte to maize, and the subsequent explosive evolution of maize in the archaeological record, the Trip-
sacum-diploperennis introgression derivatives confirm that teosinte is a progenitor of maize. Although Bennetzen et al. claim
that the Tripsacum- diploperennis crosses are not credible, DNA fingerprinting verified that the hybrids contain genes from
their teosinte and Tripsacum parents. Archaeohotanical remains of teosinte, Tripsacum, and hybrid specimens have been
reported from Tamaulipas and Oa,~aca.One of the "hybrid" specimens from Tamaulipas is virtually identical to an e-~peri-
mental Tripsacum-diploperennis segregate. The ability to experimentally reproduce forms that closely resemble archaeologi-
cal specimens lends compelling support to the hypothesis that intergeneric hyhridzation gave rise to the mutations that, through
human selection, transformed teosinte into domesticated maize.

Este ar-ticulo discute las objeciones en el articulo por Bennetzen et al. (2000)en el cual responden a MacNeish y Eubanks (2000).
Bennetzen et al. mantienen que 10s hallazgos de MacNeish y Euhanks no apoyan a la hipdtesis que considera teosinte como el
progenitor del maiz. Per-o, a1 demostrar un mecanismo de 10s pasos de transicidn delprogeizitor teosinte a1 maiz domesticado, 10s
hibridos Tripsacum-diploperennis confirman yue teosinte es un antepasado del maiz. Aunque Bennetzen et al. creen que 10s hibri-
dos Tripsacum-diploperennis no son creihles, la comprobacidn de DNA comprueba que 10s hihridos tienen 10s genes de 10s padres,
teosirite y Tripsacum. Los restos arqueohotcinicos de teosinte, Tripsacum, y 10s hibridos (que son virtualmente ide'nticos a algu-
rzas segregariones experimentales de Tripsacum-diplopervnni.~F2) apoyatz a la hipdtesis que sugier-e que una hibridacidn entre
teosinte y Tripsacum producid las mutaciones que, con la ayuda de 10s humanos, transformaron el teosinte a1 maiz domesticado.
Esta evidencia e.rperimenta1 nueva apoya a la hipdtesis que mantiene que teosinte fue un antepasado del maiz domesticado.

T
his paper is a response to criticism of Mac-
Neish and Eubanks (2000) raised in a rejoin-
der by Bennetzen et al. (this issue) concerning
the genetic evidence for the origin of maize. Although
Bennetzen et al. claim that the Tripsacum-diplop-
erennis experimental crosses upon which MacNeish
and Eubanks base their argument for involvement of
Tripsacurn in the origin of maize are not credible,
the authenticity of these crosses has been verified by
DNA testing. For descriptions and documentation of
Tripsacurn-diploperennis hybrid plants, as well as a
method for employing these crosses as a genetic
bridge to move beneficial genes from Tripsacum into
maize, seeEubanks (l989,1992,1994,1996a, 1998,
1999a). The reasons that references cited by Ben-
netzen et al. do not reveal genetic evidence for Trip-
sacum ancestry in the origin of maize are: (1) those
studies did not include many, if any, of the 16 Trip-
sacum species that exist in the Americas, and (2) the
authors ignore the literature that does signal a pos-
sible role for Tripsacurn in the history of maize
(Belousova 1970; Bethaud et al. 1997; Blakey 1993;
Brink and de Wet 1983; Chaganti 1965; de Wet et
al. 1983; Dewald et al. 1987; Eubanks 199913; Far-
quharson 1954; Galinat 1973; Galinat et al. 1964;
James 1979; Kindiger and Beckett 1990; Leblanc et
al. 1995; Lin et al. 1985; Maguire 1962; Randolph
1950; Rao and Galinat 1976; Sehgal 1963). For an
extensive, up-to-date review of the genetic evidence,
see Eubanks (2001). Specific issues raised by Ben-
netzen et al. are addressed below. Then the genetic
evidence is integrated with the evidence from the
archaeological record to bring the big picture of
maize domestication into sharper focus.

Mary W. Eubanks . Senior Research Scientist, Department of Biology, Duke University, Durham, NC 27708-0338

Latin American Antiquity, 12( I), 2001, pp. 9 1-98

Copyright0 2001 by the Society for American Archaeology

 LATIN AMERICAN ANTIQUITY [Vol. 12, No. 1,20011

A Long or Short Way to Maize? by substituting perennial teosinte for Tripsacurn.

Beadle (1939, 1980), on the other hand, favored
Rather than challenging teosinte's role in the ori- the hypothesis that maize arose from teosinte through
gin of maize, the Tripsacurn- diploperennis crosses human selection for mutations in the teosinte spike
confirm teosinte as a progenitor of maize. By exper- because it was the most parsimonious explanation,
imentally producing a prototype of ancient maize and and it accounted for the close relationship between
demonstrating how the teosinte spike can be rapidly maize and teosinte. Although Beadle acknowledged
transformed into the maize ear within a few gener- the possibility that teosinte was secondarily derived
ations, the Tripsacurn-teosinte hybrids resolve dis- from an extinct or undiscovered wild prototype, his
crepanciesbetween the biological and archaeological position was: "Any hypothesis involving an extinct
data and unify the total evidence. Because the ques- or unknown ancestor must remain unsatisfactory
tion of the origin of maize has had a long history of until tangible evidence for the existence of such a
controversy with vitriolic barbs still directed at plant is forthcoming" (1939:245).
researchers who challenge popular dogma (Dold Evidence for a long, progressive evolution from
1997), the basic principles upon which this debate teosinte to maize is absent from the archaeological
was founded are reviewed to put the defensive pos- record. The oldest archaeological remains of maize,
ture of Bennetzen et al. into historical perspective. excavated in the Valley of Tehuacin in southern Mex-
The primary advocates of the two famous compet- ico, suddenly appear in the archaeological record
ing hypotheses who argued back and forth in the lit- around 5500 B.C. (Mangelsdorf et al. 1967a), and
erature for many years were Paul Mangelsdorf and these ancient maize cobs show all the morphologi-
George Beadle. cal traits that distinguish it from its wild relatives.
Mangelsdorf and Reeves (193 1)were the first sci- However, specimens showing intermediate steps in
entists to successfully produce hybrids between Zea this transformation process are rare (Galinat 1985).
and Tripsacurn. Edgar Anderson then proposed that Although morphological characteristics of ears of F,
maize originated from a cross betweenzea and Trip- Tripsacurn-diploperennishybrids provide the miss-
sacurn (MangelsdorfandReeves 1939:212),and this ing link in the evolutionary transition to domestica-
stimulated the extensive study of maize-Tripsacurn tion (Eubanks 1995, 1997), which was the basis for
hybrids upon which Mangelsdorf and Reeves based proposing that hybridization between teosinte and
their tripartite theory for the origin of maize: (1) Tripsacurn could have given rise to the genetic diver-
extinct maize was postulated to have hybridized with sity needed for a transformation of teosinte into
Tripsacurn; (2) the annual teosintes and domesti- maize, backcrosses of the hybrids to teosinte and
cated maize evolved from such a wide cross mating; Tripsacurn mask those maize key-trait genes and the
(3)introgressionand recombination between domes- backcross progeny resemble their wild parents. How
ticated maize and the teosintes then gave rise to the such F, hybrids between teosinte and Tripsacurn
rapid and explosive evolution of maize. could produce an early maize prototype was not
When the exciting discovery of diploid perennial revealed until I studied a segregating population of
teosinte, Zea diploperennis, was reported by Iltis et ten F2 Tripsacurn-diploperennis hybrids from inter-
al. in 1979, Wilkes (1979) hypothesized that the crosses between plants that had Tripsacurn as the
annual teosintes arose from crosses between peren- maternal parent throughout the lineage. The plant that
nial teosinte and maize in the early stages of domes- produced ears like those of ancient maize from
tication. Mangelsdorf came out of retirement and Tehuacin (MacNeish and Eubanks 2000: 15, Figure
conducted breeding experiments to test the Wilkes 3) provided the "tangible evidence" Beadle required
hypothesis (Mangelsdorf et al. 1981). In a segregat- and was the basis for the article by MacNeish and
ing F, population of experimental crosses between Eubanks.
diploperennis and a primitive Mexican popcorn,
numerous phenotypes with all of the characteristics Verification of Tripsacum-Diploperennis

of races of annual teosinte were recovered among Crosses

the segregates. These findings supported Wilkes's Bennetzen et al. propose that the Tripsacurn-diplop-
idea for the origin of annual teosintes. Mangelsdorf erennis plants are actually maize-teosinte hybrids
(1983, 1986) then revised the tripartite hypothesis because the plants have the same chromosome num-
 COMMENTS 93

ber of maize and teosinte and look llke maize-teosinte fingerprinting by two different laboratories, Linkage
hybrids. The haploid chromosome number of maize Genetics and Biogenetics, Inc., has verified inheri-
and teosinte is 10, and the haploid number of Trip- tance of private alleles from each diploperennis and
sacum is 18. Although most maize-Tripsacum Tripsacum parent in all the Tripsacum-diploperen-
hybrids that have been reported contain 28 or 46 nis crosses. This documents the Tripsacum parent-
chromosomes and have a high degree of sterility, 20 age of these crosses and eliminates the putative Zea
chromosome diploids from maize-Tripsacum intro- contamination proposed by Bennetzen et al. Fur-
gression derivatives are occasionally recovered that thermore, the molecular evidence reveals that the
have up to four Tripsacum substitutionchromosomes crosses produced are extraordinary. Major genomic
and are female fertile (James 1979). Like those reorganization is required for viable progeny to be
maize-Tripsacum hybrids, the recombinant progeny formed. Recombinant progeny contain a suite of
of diploperennial teosinte and Tripsacum have 20 mutations, i.e., de novo alleles not observed in the
chromosomes. They also have greater than 90 per- polymorphism profiles of either parent (Eubanks
cent fertility. The chromosome architecture of 1999a).These mutations provide a rich store of novel
diploperennis that differs markedly from maize and genes and genetic diversity for speciation under
annual teosinte is similar to Tripsacum in terminal selection. If the Tripsacum-diploperennis recombi-
knob patterns and total length of the chromosomes nants represent a reconstructed prototype of ancient
(Eubanks 1987, 2001). This architectural similarity maize, the mutated phenotypes that would not have
enhances pairing between the chromosomes of these survived in the wild, could have been preserved
two genera and that occasionally results in viable through artificial selection by humans, and they
recombinant progeny. Cytological examination and would have provided the primordial genetic mater-
DNA fingerprinting show that precise chromosomal ial that gave rise to domesticated maize.
rearrangements consistently recur in all hybrids
recovered (Eubanks 1999a). Translocations and Molecular Evidence for the Evolutionary

fusions between the ends of some of the Tripsacum History of Zea

chromosomes, along with possible partial elimina- The molecular evidence for maize evolution includes
tion of some genetic material, are responsible for the isozyme analyses, DNA studies of nuclear and cyto-
2n = 20 diploid chromosome number of the Trip- plasmic genes, and transposable elements. For a
sacum-diploperennis recombinant offspring. Such review of these studies, see Eubanks (2001). There
change in chromosome number is a well-docu- are incongruities in different datasets; thus, inter-
mented phenomenon among wide cross hybrids in pretations about their significance for maize evolu-
experimental plant-breeding studies (Chetelat et al. tion vary. The molecular studies, which include
1989; Davies et al. 1990; Jenkins and White 1990; samples of many of the Zea taxa but few if any Trip-
Jenkins et al. 1988; John and Freeman 1975; Linde- sacum spp., consistently show a close relationship
Laursen and von Bothmer 1988; McClintock 1984; between annual teosinte (Z. m. pawiglumis and Z.
Mikklesen et al. 1996; Singh 1993; Wagner et al. m. mexicana) and maize, and this undergirds the
1993), and its occurrence in nature has been docu- rationale for the hypothesis that one or more of the
mented (Lord and Richards 1977). See Eubanks Mexican annual teosintes is the progenitor of maize.
(2001) for karyotypes of maize, annual teosinte, Phylogenetic analysis (Buckler and Holtsford 1996),
perennial teosinte, Tripsacum, and Tripsacum X however, indicates that these same teosintes are sis-
diploperennis hybrids, and for a review of the liter- ter taxa of maize descended from a common ances-
ature on change in chromosome number in wide tor rather than being progenitor to maize.
cross hybrids. Tripsacum is a large, diverse genus with as many
It is important to note that maize-teosinte hybrids as 16 species that range throughout the Americas
closely resemble maize-Tripsacum hybrids (Man- (Brink and de Wet 1983; de Wet et al. 1976, 1983).
gelsdorf and Reeves 1939, Figure 84). Therefore, It is divided into two sections: Tripsacum and Fas-
teosinte and Tripsacum have genes with the same ciculata. Section Fasciculata is of particular interest
effects, and Tripsacum-diploperenniscrosses would because its distinguishing feature is the upper mem-
therefore be expected to look like both maize-teosinte .ber of the pair of male flowers is pedicellate, and this
and maize-Tripsacum hybrids, which they do. DNA is a key feature of Zea. Most studies cited by Ben-
94 LATIN AMERICAN ANTIQUITY
netzen et al. do not include any data for Tripsacum,
or include only T. dactyloides from the Tripsacum
section which is the least Zea-like of all the Trip-
sacums. Virtually no molecular data on Tripsacum
spp. from Latin America, particularly ones in sec-
tion Fasciculata, have been published. Before we can
conclude how many genes are shared between Zea
and Tripsacum, all Tripsacum species need to be
sampled employing methods that can reveal evidence
of hybridization. If the basic operating assumption
of the method is that descent is monophyletic from
a single ancestor, it will not be possible to discern a
reticulate evolutionary pathway involving hybridiza-
tion and introgression (McDade 1992). It is also
important to consider the fact that data from extant
taxa do not adequately reflect ancient gene pools of
7,000 to 10,000years ago when the environment and
biogeographical distributions of the various species
were different from what they are today.
Archaeological Evidence for Tripsacum-
plants in that population produced a range of inflo-
Teosinte Recombinant Progeny
rescence phenotypes, some of which produced flow-
The question of the origin of maize involves biology
and archaeology, and as Bennetzen et al. point out,
"There are difficulties for those of us in one of these
fields to judge the evidence from the other." Although
a multidiscipline approach can remove the blinders
that result from the tunnel vision of discipline spe-
cialization (Eubanks 1996b), communicating across
divergent disciplines is difficult and often impedes
the problem-solving orientation of interdisciplinary
studies (Roeske 1983). In order to be meaningful,
the findings of experimental research must be con-
sistent with the archaeological record documenting
maize evolution. Equally well trained in archaeol-
ogy and biology, with extensive experience study-
ing maize-teosinte and Tripsacum- teosinte crosses,
my evidential focus on the evolutionary biology pic-
ture is through the archaeological lens.
There is no evidence in the archaeological record
for a long, gradual evolution in which the mutations
that transformed teosinte into maize could have accu-
mulated. Reexamination of the record in light of the
evidence from segregating experimental Tripsacum-
diploperennis crosses reveals that the transition from
teosinte to maize may have required only a few gen-
erations and could have happened very rapidly. Ears
produced by F, Tripsacum-diploperennis plants sim-
ulated a missing link in the evolutionary history of
maize (Eubanks 1995, 1997). Like maize, they had
[Vol. 12, No. 1, 20011
a pair of kernels in a single cupule instead of a sin-
gle kernel in a cupulate fruitcase as in teosinte. Also,
like maize, the cupulate segments did not break apart
easily as they do in teosinte and Tripsacum. Even so,
the F, specimens were missing important features of
maize. Although the kernels were slightly exposed
at the tips of the glumes, they were still partially
encased in hard glumes that is a characteristic of
teosinte and Tripsacum. Also, the rachis of the spike
was two-ranked as in teosinte and Tripsacum, rather
than a multirowed structure as in the maize cob. I did
not recover arecombinant phenotype with all the fea-
tures of early maize until I grew an F, population
from segregating Tripsacum-teosinte crosses in
which Tripsacum was the maternal parent of the lin-
eage. One of the segregants from that population
produced ears that are a match for the reconstruction
of ancient Tehucan maize drawn by Galinat (com-
pare Mangelsdorf et al. 1967b:200, Figure 124 to
MacNeish and Eubanks 2000:15, Figure 3). Other
ering spikes that closely resemble archaeobotanical
specimens from Oaxaca and Tamaulipas.
Four primitive maize specimens were found just
above Zone B1 in Guil6 Naquitz rockshelter in the
Valley of Oaxaca (Flannery 1986:8). The Oaxaca
specimens were examined by Richard I. Ford and
George Beadle. Both experts agreed they were either
(1) "maize-teosinte hybrids," or (2) "primitive maize
that shows strong teosinte influence in its ancestry."
One of those unusual ears bears striking morpho-
logical resemblance to ears produced by an F, Trip-
sacum-teosinte derivative in which Tripsacum was
the maternal parent of its progenitor F, hybrid (com-
pare Figure 1 to Flannery 1986:8, Figure 1.2e). Both
specimens have single spikelets in triangular fruit-
cases that are borne in yolks. Yolking is a precursor
feature in the transition from the two-ranked, shat-
tering fruitcases of the wild relatives to the many-
ranked, fused rachis that forms the maize cob
(Galinat 1970). Another Oaxaca specimen (Flannery
198623,Figure 2. lc) is of a two-rowed cob, and sim-
ilar two-rowed cobs were produced by plants among
the same Tripsacum-teosinte segregants. In view of
the AMS dates (Smith 1997) and conventional radio-
carbon dates for Zone B at Guila Naquitz (Flannery
1986), it is possible these Oaxaca specimens could
date to as early as ca. 6000 B.P. Since pollen with
the morphological attributes of modern examples of
 COMMENTS
Figure 1. F, Tripsacum-teosinte derivative with spikelets in
triangular fruitcases borne in yolks. Yolking is a precursor
in the transition from teosinte's two-ranked shattering spike
to the many-ranked maize cob. This inflorescence type bears
resembalnce to a preceramic "hybrid" maize specimen from
Oaxaca (see Flannery 1986:8, Figure 1.2~).
Tripsacum was reported in Zones B1, B2 and C,
maize in Zones B 1 and B2, and teosinte in Zone B2,
it is possible that these specimens represent a segre-
gating population of Tripsacum-teosinte hybrids that
were either being gathered in the wild, or were in the
initial stages of maize domestication. This fits the
explanation proposed by Beadle and Ford that the
specimens represent "primitive maize that shows
strong teosinte influence in its ancestry."
Archaeobotanical remains of Tripsacum have
been found along with teosinte in Mesoamerica in
the Infiernillo Canyon caves in southwestern
Tamaulipas (Mangelsdorf et al. 1967b). This is
highly significant because the biogeographic range
of teosinte today is west of the Sierra Madre Orien-
tal mountain range. Along with the Tripsacum and
teosinte remains are specimens that were described
in the report as "maize-teosinte" hybrids. These spec-
imens were identified as maize-teosinte hybrids
95
Figure 2. Tripsacum-teosinte segregant on the left compared
to a prehistoric "hybrid" from Romero's Cave on the right.
Note the single kernel per cupule instead of the paired ker-
nels found in maize-teosinte hybrids, the soft papery glumes
instead of the hard lower glume of teosinte, and the non-
shattering rachis. The archaeobotanical specimen is in the
Tamaulipas collection at Harvard University.
because they had thicker stalks than teosinte and the
rachis did not break apart like the shattering fruit-
cases of teosinte. However, there must have been
some question because labels accompanying the
specimens read "maize-teosinte hybrid?" A con-
founding feature of these specimensis that they have
a single kernel per cupule instead of the paired ker-
nels found in maize-teosinte hybrids, soft papery
glumes instead of the hard lower glume of teosinte,
and nonshattering rachises. This was why the authors
were not certain whether the putative maize-teosinte
hybrids were F, hybrids or segregates appearing in
subsequent generations,and they entertained the pos-
sibility that the specimens represented the ancestral
form of maize. Among these hybrid specimens at
Harvard University is a virtual match for an inflo-
rescence phenotype produced by one of the Trip-
sacum-teosinte segregants (Figure 2). This supports
the idea that the hybrids represent segregatesof sub-
96 LATIN AMERICAN ANTIQUITY [Vol. 12, No. 1,2001]

sequent generations of the ancestral form of maize, introgression between Tripsacurn and a primitive
but they are probably Tripsacurn-teosinte derivatives teosinte. Once humans began selecting and cultivat-
rather than maize-teosinte hybrids. ing plants with recombinant genomes from these
Other archaeobotanical evidence for Tripsacurn wild grasses, the evolution of domesticated maize
use is large quantities of seed recovered from rock- could have progressed quite rapidly. The AMS dates
shelters in eastern North America (Gilmore 1931; obtained by Dolores Piperno on the preceramic
Jones 1936). AMS dates on Chenopodiurn seeds maize from GuilLi Naquitz rockshelter in Oaxaca
from Newt Kash Hollow in Kentucky, a site where promise to shed more light on the question of where
Tripsacurn was found, are ca. 3400 B.P. (Smith and when such a "hopeful [recombinant] monster"
1992).Although the Tripsacurn grain is contained in (Gould 1984)bearing the primordial genes of maize
a hard cupulate fruitcase like teosinte, the edge of first appeared.
the outer glume does not overlap the inner glume like
it does in teosinte, a trait that makes it virtually impos- Acknowledgments. A special thank you to Susan Rossi-Wilcox
sible to extract the teosinte kernel for food. This dis- and Malinda Bluestain for facilitating my study of the
Tamaulipas specimens at Harvard University and the Tehuacin
tinctive feature of the Tripsacurn fruitcase makes it materials at the Robert S. Peabody Museum, respectively. I am
relatively easy to extract the grain. Since Tripsacurn grateful to Richard S. MacNeish, Walton C. Galinat, Margaret
is nutritious with three times the protein content of Houston, James Schoenwetter, James Knutson, and Laura
maize and teosinte (Jackson 1980),is delicious, and Dunn for their helpful comments on earlier drafts of this paper.
the kernels are as large as popcorn, prehistoric Research support was provided by the National Science
Foundation grant nos. 9660146,9801386, and DEB-94-15541.
hunters and foragers were probably exploiting the
grain for food in addition to other uses such as mate- References Cited
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An Interdisciplinary Perspective on the Origin of Maize
Mary W. Eubanks
Latin American Antiquity, Vol. 12, No. 1. (Mar., 2001), pp. 91-98.
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