Plant Biotechnology Journal (2003) 1, pp. 59–70
© 2003 Blackwell Publishing Ltd 59
Blackwell Science, Ltd
Transcriptional control of nutrient partitioning during rice
grain filling
Tong Zhu*, Paul Budworth, Wenqiong Chen, Nicholas Provart
1
, Hur-Song Chang, Sonia Guimil, Wenpei Su,
Bram Estes, Guangzhou Zou
2
and Xun Wang
Torrey Mesa Research Institute, Syngenta Research and Technology, 3115 Merryfield Row, San Diego, CA 92121, USA
Summary
Cereal grains accumulate carbohydrates, storage proteins and fatty acids via different
pathways during their development. Many genes that participate in nutrient partitioning
during grain filling and that affect starch quality have been identified. To understand how
the expression of these genes is coordinated during grain development, a genomic
approach to surveying the participation and interactions of all the pathways is necessary.
Using recently published rice genome information, we designed a rice GeneChip
microarray that covers half the rice genome. By monitoring the expression of 21 000
genes in parallel, we identified genes involved in the grain filling process and found that
the expression of genes involved in different pathways is coordinately controlled in a
synchronized fashion during grain filling. Interestingly, a known promoter element in
genes encoding seed storage proteins, AACA, is statistically over-represented among the
269 genes in different pathways with diverse functions that are significantly up-regulated
during grain filling. By expression pattern matching, a group of transcription factors that
have the potential to interact with this element was identified. We also found that most
genes in the starch biosynthetic pathway show multiple distinct spatial and temporal
expression patterns, suggesting that different isoforms of a given enzyme are expressed
in different tissues and at different developmental stages. Our results reveal key
regulatory machinery and provide an opportunity for modifying multiple pathways by
manipulating key regulatory elements for improving grain quality and quantity.
Received 6 September 2002; revised 23
September 2002; accepted 1 October 2002.
* Correspondence (fax +1 858 812 1097;
e-mail tong.zhu@syngenta.com)
Present addresses:
1
Department of Botany,
University of Toronto, 25 Willcocks Street,
Toronto, Ontario, M5S 3B2, Canada;
2
ActivX
Biosciences, 11025 N. Torrey Pines Road, La
Jolla, CA 92037, USA .
Keywords: gene expression, grain
development, microarray, Oryza sativa ,
regulatory elements.
Introduction
The most important economic characteristics of agricultural
grain crops are their yield, nutritional characteristics and culi-
nary quality. Yield and nutritional value are mostly deter-
mined by the synthesis and storage of carbohydrates,
proteins and minerals during grain filling, and culinary
quality is affected by the interaction of various enzymes to
produce the final structure of the starch at the molecular
and granule levels. The manipulation of these pathways
can result in significant improvements in nutritional value
(Mazur et al ., 1999; Ye et al ., 2000). For example, reducing
the amount of even one enzyme in the starch biosynthetic
pathway (granule-bound starch synthase) can dramatically
affect the culinary quality (Singh et al ., 2000; Umemoto
et al ., 1995), resulting in softer, less sticky cooked rice. Many
genes that participate in nutrient partitioning during grain
filling and that affect starch quality have been identified
in rice and other cereals (e.g. Umemoto et al ., 1995). How-
ever, these genes and their transcriptional controls are
poorly understood, especially in cereal grains.
Major biological processes such as cereal grain filling are
believed to require a close coordination of gene expression
among many important pathways. However, direct experi-
mental evidence for this hypothesis has been lacking due to
the absence of genomic sequence information and func-
tional genomic technologies. To prove this hypothesis and
to identify key regulators in grain development, a parallel
examination of gene expression on a genome scale is
necessary. Recent studies have examined gene expression