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doi:10.1534/genetics.106.056580
A more recent version of this article appeared on June 1, 2006.
REGULAR RESEARCH PAPERS |
Non-additive regulation of FRI and FLC loci mediates flowering-time variation in Arabidopsis allopolyploids
Jianlin Wang 1, Lu Tian 2, Hyeon-Se Lee 1 and Z. Jeffrey Chen 2*
1 Texas A&M University
2 University of Texas at Austin
* To whom correspondence should be addressed. E-mail: zjchen{at}mail.utexas.edu.
Submitted on January 31, 2006
Revised on March 5, 2006
Accepted on 14 March 2006
Allopolyploidy is formed by combining two or more divergent genomes and occurs throughout the evolutionary history of many plants and some animals. Transcriptome analysis indicates that many genes in various biological pathways including flowering time are expressed non-additively (different from the mid-parent value). However, the mechanisms for non-additive gene regulation in a biological pathway are unknown. Natural variation of flowering time is largely controlled by two epistatically acting loci, namely, FRIGIDA (FRI) and FLOWERING LOCUS C (FLC). FRI upregulates FLC expression that represses flowering in Arabidopsis. Synthetic Arabidopsis allotetraploids contain two sets of FLC and FRI genes originating from A. thaliana and A. arenosa, respectively and flower late. Inhibition of early flowering is caused by upregulation of A. thaliana FLC (AtFLC) that is trans-activated by A. arenosa FRI (AaFRI). Two duplicate FLCs (AaFLC1 and AaFLC2) originating from A. arenosa are expressed in some allotetraploids but silenced in other lines. The deletions in the promoter regions and first introns of A. arenosa FLCs are associated with expression variation in the allotetraploids. The strong AtFLC and AaFLC loci are maintained in natural Arabidopsis allotetraploids, leading to extremely late flowering. Furthermore, FLC expression correlates positively with histone H3-Lys4 methylation and H3-Lys9 acetylation and negatively with H3-Lys9 methylation, epigenetic marks for gene activation and silencing. We provide evidence for interactive roles of sequence evolution, chromatin modification, and trans-acting effects in natural selection of orthologous FLC loci, which determines the fate of duplicate genes during allopolyploid evolution.
Key Words: FLC, FRI, epigenetic regulation, evolution, polyploidy
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