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WUSCHEL-related homeobox (WOX) gene is a plant-specific clade of homeobox transcription factors. Increasing evidences reveal that WOXs play critical roles in early embryogenesis, which involves zygote development, initiation of zygote division, and apical or basal cell lineage establishment. However, how  WOXs  regulate these developmental events remains largely unknown, and even detailed expression pattern in gametes and early proembryos is not yet available. Here, 13  WOX  family genes were identified in  Nicotiana tabacum  genome. Comparative analysis of 13  WOX  family genes with their homologs in  Arabidopsis thaliana  reveals relatively conserved expression pattern of  WUS  and  WOX5  in shoot/root apical meristem. Whereas variations were also found, e.g., lacking homolog of  WOX8  (a marker for suspensor cell) in tobacco genome and the expression of  WOX2 / WOX9  in both apical cell and basal cell. Transient transcriptional activity analysis revealed that WOXs in WUS clade have repressive activities for their target's transcription, whereas WOXs in ancient and intermediate clade have activation activities, giving a molecular basis for the phylogenetic classification of tobacco WOXs into three major clades. Expression pattern analysis revealed that some  WOXs  (e.g.,  WOX 13a ) expressed in both male and female gametes and some  WOXs  (e.g.,  WOX 11  and  WOX 13b ) displayed the characteristics of parent-of-origin genes. Interestingly, some  WOXs  (e.g.,  WOX2  and  WOX9 ), which are essential for early embryo patterning, were  de novo  transcribed in zygote, indicating relevant mechanism for embryo pattern formation is only established in zygote right after fertilization and not carried in by gametes. We also found that most  WOXs  displayed a stage-specific and cell type-specific expression pattern. Taken together, this work provides a detailed landscape of  WOXs  in tobacco during fertilization and early embryogenesis, which will facilitate the understanding of their specific roles in these critical developmental processes of embryogenesis.

frontiers in plant science

Comparative Analysis of WUSCHEL-Related Homeobox Genes Revealed Their Parent-of-Origin and Cell Type-Specific Expression Pattern During Early Embryogenesis in Tobacco