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as our understanding of plant response to stress
increases.
Identifying drought tolerance traits using
major-effect drought yield QTL lines presents
a unique opportunity to explore the traits that are
affecting plant response to drought, with con-
fidence that the differences in traits between
+
ular in respective drought-prone rice-growing
regions. Pyramiding of multiple DTY QTLs,
together with pathogen-resistance genes, is the
next step for meeting the goal of appreciable (
1
tha 1 ) yield advantages under drought stress that
could be adopted by farmers.
>
QTL lines have a high proba-
bility of being responsible for differences in
yield. Working with material that is known to
have improved yield under drought and being
able to compare genetically similar germplasm
(
QTL and
References
Asia Rice Foundation. 2005. http://www.asiarice.org/
sections/learnrice/riceislife.html.
Atlin G.N., Lafitte H.R., 2002. Marker-assisted breeding ver-
sus direct selection for drought tolerance in rice. In:
Saxena N.P., O'Toole J.C. (Eds.), Field Screening for
Drought Tolerance in Crop Plants with Emphasis on
Rice. Proceedings of an International Workshop on Field
Screening for Drought Tolerance in Rice , 11-14 Dec
2000, ICRISAT, Patancheru, India, p. 208.
Bernardo R., Charcosset A. 2006. Usefulness of gene infor-
mation in marker-assisted recurrent selection: A simula-
tion appraisal. Crop Science 46:614-621.
Bernardo R., Yu J. 2007. Prospects of genome wide selection
for quantitative traits in maize. Crop Science 47:1082-
1090.
Bernier J., Atlin G.N., Serraj R., Kumar A., Spaner D. 2008.
Breeding upland rice for drought resistance. Journal of
the Science of Food and Agriculture 88:927-939.
Bernier J., Kumar A., Venuprasad R., Spaner D., Atlin
G.N. 2007. A large-effect QTL for grain yield under
reproductive-stage drought stress in upland rice. Crop
Science 47:507-516.
Bernier J., Serraj R., Kumar A., Venuprasad R., Impa S.,
Gowda R.P.V., Oane R., Spaner D., Atlin G. 2009. The
large-effect drought-resistance QTL qtl12.1 increases
water
QTL lines) facilitates more rapid
pinpointing of the traits behind the QTLs. Fur-
thermore, this approach allows us to be open-
minded in our understanding of the mecha-
nisms behind drought tolerance; selecting for
yield under drought, as opposed to selecting for
specific traits, is confirming that many mecha-
nisms - some unexpected - can contribute to
grain yield under drought.
+
QTL and
Summary
Recent progress in improving rice yield under
drought through direct selection for yield is ben-
efiting from the efficiency and high-throughput
capacity of molecular strategies. Advancement
of generations while simultaneously applying
drought stress and the use of large (
300 lines)
populations have been key components of this
progress. Major-effect drought-yield QTLs that
are effective in multiple backgrounds across a
wide range of environments have been identified.
Some of these QTLs are also seen to be con-
tributed from multiple drought-tolerance donor
genotypes. The drought response of physiologi-
cal traits associated with the drought-yield QTLs
has agreed with the yield response to increas-
ing drought stress severity and has pointed
to some novel drought traits. Marker-assisted
backcrossing and the development of BIL and
NIL populations of the drought yield QTLs in
multiple susceptible backgrounds is an efficient
way of improving the drought tolerance of local
varieties whose grain and plant types are pop-
>
uptake
in
upland
rice.
Field
Crops
Research
110:139-146.
Blum A. 1988. Plant Breeding for Stress Environments . CRC
Press, Boca Raton, FL.
Blum A. 2011. Drought resistance - is it really a complex
trait? Functional Plant Biology 38:753-757.
Collard B.C.Y., Jahufer M.Z.Z., Brouwer J.B., Pang E.C.K.
2005. An introduction to markers, quantitative trait loci
(QTL) mapping and marker-assisted selection for crop
improvement: The basic concepts. Euphytica 142:169-
196.
Collins N.C., Tardieu F., Tuberosa R. 2008. Quantitative trait
loci and crop performance under abiotic stress: Where do
we stand? Plant Physiology 147:469-486.
Courtois B., Shen L., Petalcorin, Carandang S., Mauleon
R., Li Z.K. 2003. Locating QTLs controlling constitu-
tive root traits in the rice population IAC 165 x Co39.
Euphytica 134:335-345.
Courtois B., Lafitte R. 1999. Improving rice for drought-
prone upland environments. In: Ito O., O'Toole J., Hardy
B. (Eds.) Genetic Improvement of Rice for Water-limited
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