Title : SKIP, a splicing factor, links alternative splicing to SWR1 chromatin remodeling complex and controls flowering time in Arabidopsis
Abstract:
S imilar to other eukaryotes, splicing is emerging as an important process affecting development and stress tolerance in plants. Ski-interacting protein (SKIP), a splicing factor, is essential for circadian clock function and abiotic stress tolerance; however, the mechanisms whereby it regulates flowering time are unknown. In this study, we found that mutation of SKIP, skip-1, confers early flowering phenotype under long day (LD) and shot day (SD) conditions. SKIP is required for the splicing of serrated leaves and early flowering (SEF) pre-mRNA through directly binding, which encodes a component of the ATP-dependent SWR1 chromatin remodeling complex (SWR1-C). It has been reported that the SWR1-C exchanges histone H2A for H2A.Z, producing variant nucleosomes. The SWR1-C is required for H2A.Z deposition at flowering locus c (FLC), MADS affecting flowering (MAF) 4, and MAF5 chromatin (which encode central flowering-time suppressors), and promotes transcription at these loci. The skip-1 mutation caused the elevated accumulation of alternatively spliced SEF isoforms, including those showing intron retention. However, the level of mature SEF mRNA encoding functional SEF protein was reduced by skip-1. Defects in the splicing of SEF pre-mRNA reduced H2A.Z enrichment at the chromatin of FLC, MAF4, and MAF5, suppressed the expression of these genes; activated the expression of suppressor of constans 1 (SOC1), flowering locus T (FT), and twin sister of FT (TSF) (which encode flowering-time integrators); produced an early flowering phenotype in skip-1 plants. Our findings indicate that SKIP regulates SWR1-C function via alternative splicing to control the floral transition in Arabidopsis thaliana