Laboratory of Genetics, University of Wisconsin, 425-G Henry Mall, Madison, WI, 53706, USA.
Department of Biochemistry, University of Wisconsin, 433 Babcock Dr., Madison, WI, 53706, USA.
Plant J. 2018 Mar;93(5):871-882. doi: 10.1111/tpj.13815. Epub 2018 Feb 7.
Many plants require prolonged exposure to cold to acquire the competence to flower. The process by which cold exposure results in competence is known as vernalization. In Arabidopsis thaliana, vernalization leads to the stable repression of the floral repressor FLOWERING LOCUS C via chromatin modification, including an increase of trimethylation on lysine 27 of histone H3 (H3K27me3) by Polycomb Repressive Complex 2 (PRC2). Vernalization in pooids is associated with the stable induction of a floral promoter, VERNALIZATION 1 (VRN1). From a screen for mutants with a reduced vernalization requirement in the model grass Brachypodium distachyon, we identified two recessive alleles of ENHANCER OF ZESTE-LIKE 1 (EZL1). EZL1 is orthologous to A. thaliana CURLY LEAF 1, a gene that encodes the catalytic subunit of PRC2. B. distachyon ezl1 mutants flower rapidly without vernalization in long-day (LD) photoperiods; thus, EZL1 is required for the proper maintenance of the vegetative state prior to vernalization. Transcriptomic studies in ezl1 revealed mis-regulation of thousands of genes, including ectopic expression of several floral homeotic genes in leaves. Loss of EZL1 results in the global reduction of H3K27me3 and H3K27me2, consistent with this gene making a major contribution to PRC2 activity in B. distachyon. Furthermore, in ezl1 mutants, the flowering genes VRN1 and AGAMOUS (AG) are ectopically expressed and have reduced H3K27me3. Artificial microRNA knock-down of either VRN1 or AG in ezl1-1 mutants partially restores wild-type flowering behavior in non-vernalized plants, suggesting that ectopic expression in ezl1 mutants may contribute to the rapid-flowering phenotype.
许多植物需要长时间暴露在低温下才能获得开花的能力。低温暴露导致开花能力的过程称为春化作用。在拟南芥中,春化作用导致花抑制因子 FLOWERING LOCUS C 通过染色质修饰稳定抑制,包括多梳抑制复合物 2 (PRC2) 导致组蛋白 H3 赖氨酸 27 三甲基化 (H3K27me3) 的增加。禾本科植物中的春化作用与花启动子 VERNALIZATION 1 (VRN1) 的稳定诱导有关。在模式禾本科植物柳枝稷中,我们通过筛选对春化作用要求降低的突变体,鉴定出两个隐性增强子的 ZESTE 样 1 (EZL1) 等位基因。EZL1 与拟南芥卷曲叶 1 同源,卷曲叶 1 基因编码 PRC2 的催化亚基。B. distachyon ezl1 突变体在长日照 (LD) 光周期中无需春化作用即可快速开花;因此,EZL1 是在春化作用之前维持营养状态所必需的。ezl1 中的转录组研究表明,数千个基因的表达失调,包括叶中几个花同源基因的异位表达。EZL1 的缺失导致 H3K27me3 和 H3K27me2 的全局减少,这与该基因在 B. distachyon 中对 PRC2 活性的主要贡献一致。此外,在 ezl1 突变体中,开花基因 VRN1 和 AGAMOUS (AG) 异位表达且 H3K27me3 减少。在 ezl1-1 突变体中,人工 microRNA 敲低 VRN1 或 AG 部分恢复了非春化植物的野生型开花行为,表明 ezl1 突变体中的异位表达可能导致快速开花表型。