Bread wheat (Triticum aestivum) is a hexaploid species with A, B, and D ancestral genomes. Most bread wheat genes are present in the genome as triplicated homoeologous genes (homoeologs) derived from the ancestral species. Here, we report that both genetic and epigenetic alterations have occurred in the homoeologs of a wheat class E MADS box gene. Two class E genes are identified in wheat, wheat SEPALLATA (WSEP) and wheat LEAFY HULL STERILE1 (WLHS1), which are homologs of Os MADS45 and Os MADS1 in rice (Oryza sativa), respectively. The three wheat homoeologs of WSEP showed similar genomic structures and expression profiles. By contrast, the three homoeologs of WLHS1 showed genetic and epigenetic alterations. The A genome WLHS1 homoeolog (WLHS1-A) had a structural alteration that contained a large novel sequence in place of the K domain sequence. A yeast two-hybrid analysis and a transgenic experiment indicated that the WLHS1-A protein had no apparent function. The B and D genome homoeologs, WLHS1-B and WLHS1-D, respectively, had an intact MADS box gene structure, but WLHS1-B was predominantly silenced by cytosine methylation. Consequently, of the three WLHS1 homoeologs, only WLHS1-D functions in hexaploid wheat. This is a situation where three homoeologs are differentially regulated by genetic and epigenetic mechanisms.
In wheat plants at the vegetative growth stage, the shoot apical meristem (SAM) produces leaf primordia. When reproductive growth is initiated, the SAM forms an inflorescence meristem (IM) that differentiates a series of spikelet meristem (SM) as the branch. The SM then produces a series of floret meristem (FM) as the branch. To identify the mechanisms that regulate formation of the reproductive meristems in wheat, we have investigated a leaf initiation mutant, fushi-darake (fdk) which was developed by ion beam mutagenesis. The morphological traits were compared in wild type (WT) and fdk mutant plants grown in the experimental field. WT plants initiated leaves from SAM at regular intervals in spiral phyllotaxy, while fdk plants had 1/2 alternate phyllotaxy with rapid leaf emergence. The fdk plants have increased numbers of nodes and leaves compared with WT plants. The time interval between successive leaf initiation events (plastochron) was measured in plants grown in a growth chamber. The fdk plants clearly show the rapid leaf emergence, indicating a shortened plastochron. Each tiller in fdk plants branches at the upper part of the culm. The fine structure of organ formation in meristems of fdk plants was examined by scanning electron microscopy (SEM). The SEM analysis indicated that fdk plants show transformation of spikelet meristems into vegetative shoot meristems. In conclusion, the fdk mutant has a heterochronic nature, i.e., both reproductive and vegetative programs were simultaneously in operation during the reproductive phase, resulting in a shortened plastochron and transformation of reproductive spikelets into vegetative shoots.
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