CROP SCIENCE, VOL. 53, JANUARY– FEBRUARY 2013 WWW.CROPS.ORG 189
RESEARCH
F
or millions of people in developing countries, plant products
with dietary provitamin A carotenoids such as b -carotene,
b -cryptoxanthin, and a-carotene are the major source of vitamin
A (West and Darnton-Hill, 2008). Within these countries, insuf-
icient provitamin A or vitamin A content in diets is a major cause
Genetic Analysis of Visually Scored Orange
Kernel Color in Maize
Kristin Chandler, Alexander E. Lipka, Brenda F. Owens, Huihui Li, Edward S. Buckler,
Torbert Rocheford,* and Michael A. Gore*
ABSTRACT
Increasing levels of provitamin A carotenoids in
maize (Zea mays L.) grain through plant breed-
ing has potential to help humans suffering from
vitamin A deiciency. In parts of Africa where this
deiciency is prevalent, there is a consumer pref-
erence for white maize grain and an avoidance of
yellow maize grain. White grain has minimal lev-
els of carotenoids whereas yellow grain can have
appreciable levels of carotenoids. There is a new
effort to introduce orange maize that contains
high levels of provitamin A, which appears to be
a more acceptable color than yellow to consum-
ers in Africa. The implementation of this program
requires backcross selection to convert African-
adapted germplasm with white grain to orange.
We conducted a study to assess the heritability
of visual scores for relative intensity of orange
kernel color and identify genetic markers asso-
ciated with orange color across and within 10
families of the maize nested association map-
ping population. We found visually scored ker-
nel color to have a moderately high heritability
and identiied ive common quantitative trait loci
(QTL) and six rare QTL for intensity of orange
color. Notably, half of them coincided with carot-
enoid biosynthetic genes. Our results indicate
that breeders in Africa, Asia, and throughout the
world would have lexibility to select for orange
kernel color visually and/or with gene-speciic
markers. Such selection can be combined with
marker-assisted selection efforts to increase
provitamin A levels in maize grain.
K. Chandler, B. Owens, and T. Rocheford, Department of Agronomy,
Purdue University, West Lafayette, IN; A.E. Lipka, USDA-ARS,
Robert W. Holley Center for Agriculture and Health, Ithaca, NY; H.
Li, Institute of Crop Science, The National Key Facility for Crop Gene
Resources and Genetic Improvement, Chinese Academy of Agricultural
Sciences, Beijing, China; E.S. Buckler, USDA-ARS, Robert W. Holley
Center for Agriculture and Health, Institute for Genomic Diversity,
Department of Plant Breeding and Genetics, Cornell University, Ithaca,
NY; M.A. Gore, USDA-ARS, U.S. Arid Land Agricultural Research
Center, Maricopa, AZ; Current Address: K. Chandler, Pioneer,
Hi-Bred International, Inc., 4200 West Service Road, Unit 4, Evans,
CO 80620. K. Chandler and A.E. Lipka contributed equally to this
work. *Corresponding authors (Torbert@purdue.edu; Michael.Gore@
ars.usda.gov).
Abbreviations: AR1, irst-order autoregressive; BLUP, best lin-
ear unbiased predictor; CCD, carotenoid cleavage dioxygenase; FDR,
false discovery rate;
2
ˆ
C
h , heritability across the entire experiment;
2
ˆ
l
h ,
heritability on a line-mean basis across all nested association mapping
families;
2
ˆ
p
lw
h , heritability on a line-mean basis within only the pth nested
association mapping family;
2
ˆ
p
h , heritability on an individual plot basis
across all nested association mapping families; MAS, marker-assisted
selection; NAM, nested association mapping; PVE, phenotypic variance
explained; Q×E, quantitative trait loci × environment; QTL, quantita-
tive trait loci/locus; RIL, recombinant inbred line; SNP, single nucleo-
tide polymorphism; ZEP, zeaxanthin epoxidase.
Published in Crop Sci. 53:189–200 (2013).
doi: 10.2135/cropsci2012.02.0129
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