Abstract

BackgroundMany genes involved in responses to photoperiod and vernalization have been characterized or predicted in Arabidopsis (Arabidopsis thaliana), Brachypodium (Brachypodium distachyon), wheat (Triticum aestivum) and barley (Hordeum vulgare). However, little is known about the transcription regulation of these genes, especially in the large, complex genomes of wheat and barley.ResultsWe identified 68, 60, 195 and 61 genes that are known or postulated to control pathways of photoperiod (PH), vernalization (VE) and pathway integration (PI) in Arabidopsis, Brachypodium, wheat and barley for predicting transcription factor binding sites (TFBSs) in the promoters of these genes using the FIMO motif search tool of the MEME Suite. The initial predicted TFBSs were filtered to confirm the final numbers of predicted TFBSs to be 1066, 1379, 1528, and 789 in Arabidopsis, Brachypodium, wheat and barley, respectively. These TFBSs were mapped onto the PH, VE and PI pathways to infer about the regulation of gene expression in Arabidopsis and cereal species. The GC contents in promoters, untranslated regions (UTRs), coding sequences and introns were higher in the three cereal species than those in Arabidopsis. The predicted TFBSs were most abundant for two transcription factor (TF) families: MADS-box and CSD (cold shock domain). The analysis of publicly available gene expression data showed that genes with similar numbers of MADS-box and CSD TFBSs exhibited similar expression patterns across several different tissues and developmental stages. The intra-specific Tajima D-statistics of TFBS motif diversity showed different binding specificity among different TF families. The inter-specific Tajima D-statistics suggested faster TFBS divergence in TFBSs than in coding sequences and introns. Mapping TFBSs onto the PH, VE and PI pathways showed the predominance of MADS-box and CSD TFBSs in most genes of the four species, and the difference in the pathway regulations between Arabidopsis and the three cereal species.ConclusionOur approach to associating the key flowering genes with their potential TFs through prediction of putative TFBSs provides a framework to explore regulatory mechanisms of photoperiod and vernalization responses in flowering plants. The predicted TFBSs in the promoters of the flowering genes provide a basis for molecular characterization of transcription regulation in the large, complex genomes of important crop species, wheat and barley.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-016-2916-7) contains supplementary material, which is available to authorized users.

Highlights

  • Many genes involved in responses to photoperiod and vernalization have been characterized or predicted in Arabidopsis (Arabidopsis thaliana), Brachypodium (Brachypodium distachyon), wheat (Triticum aestivum) and barley (Hordeum vulgare)

  • The lack of promoter sequences in the two Arabidopsis genes, one in PH and the other in VE, is due to their overlap with upstream adjacent genes. These overlaps are: a protein-coding gene AT2G18915 [ADAGIO2 (ADO2)/LOV KELCH PROTEIN 2 (LKP2)] on the reverse strand overlaps with a noncoding RNA gene AT2G18917 on the forward strand, whereas AT2G18880 [VERNALIZATION5/VIN3-LIKE 2 (VEL2)/VIN3-LIKE 3 (VIL3)] overlaps with AT2G18876 encoding a microtubuleassociated protein; both genes are on the forward strand

  • Using the FIMO motif discovery tool in MEMPpE [28], we predicted a large number of putative transcription factor binding sites (TFBSs) in the promoters of the genes related to the PH, VE and pathway integration (PI) pathways in Arabidopsis, Brachypodium, wheat and barley

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Summary

Introduction

Many genes involved in responses to photoperiod and vernalization have been characterized or predicted in Arabidopsis (Arabidopsis thaliana), Brachypodium (Brachypodium distachyon), wheat (Triticum aestivum) and barley (Hordeum vulgare). The genetic basis of flowering time control has been studied extensively in the model plant Arabidopsis (Arabidopsis thaliana) with over 200 putative floweringrelated genes being identified [1, 2] These genes have served as the reference for genome-wide prediction of flowering gene homologs in other plants including a cereal model species Brachypodium (Brachypodium distachyon), and two important cereal crops in shortseason cropping regions, wheat (Triticum aestivum) and barley (Hordeum vulgare) [3, 4]. A recent study reported that two-thirds of the key circadian clock components are conserved in Arabidopsis and barley [11], including critical photoperiod genes like CONSTANS (CO), EARLY FLOWERING 4 (ELF4), and PSEUDO RESPONSE REGULATORs (PRRs) Another major flowering-related pathway controlling the vernalization response pathway is reported to be less conserved between Arabidopsis and monocots [5,6,7,8,9]

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