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Analyses of Ectodermal Transcription Factors and Identification of Cis-regulatory Elements for the Left-Sided Nodal Gene Expression during Embryogenesis of the Ascidian, Halocynthia roretzi

施, 禹 大阪大学 DOI:10.18910/82030

2021.03.24

概要

In developing embryos, appropriate gene expression mediates cell fate decision, differentiation, and morphogenesis. The spatiotemporal expression of zygotic genes is regulated by transcription factors. Investigation of the expression patterns and the transcriptional regulatory relationships is thus essential to understand embryonic development. The ascidian, Halocynthia roretzi, is a tunicate characterized by fast and invariant embryogenesis. Taking advantages of ascidian embryogenesis, gene regulatory networks for ectodermal development and left-right determination were analyzed.

First, spatiotemporal expression patterns and developmental roles of three transcription factors that are expressed in ectodermal cells of the early embryos were studied. Staged RNA- seq of the ascidian H. roretzi has previously suggested that these genes encoding transcription factors are transiently expressed at the blastula stage, which is the stage at which cell fates are specified and differentiation starts. In the present study, the expression patterns and functions of the transcription factors, FoxJ-r, SoxF, and SP8/9 were studied. The results showed that all three genes were expressed in the animal hemisphere as early as the 16-cell stage. Functional analyses using FoxJ-r morphants showed that they resulted in the disruption of laterality and the absence of epidermal mono-cilia, suggesting its functions in cilia formation and, consequently, in the generation of left-right asymmetry. SoxF knockdown resulted in incomplete epiboly by the ectoderm during gastrulation, while SP8/9 knockdown showed no phenotype until the tailbud stage in the present study, although it was expressed during blastula stages. These results indicate that transcription factor genes expressed at the cleavage stages play roles in diverse functions, and their roles are not limited to cell fate specification.

Second, the cis-regulatory elements of nodal gene were analyzed. The nodal signaling pathway plays an important role in establishment of the left-right axis during embryogenesis in many animals. Previous studies have shown that the left-sided expression of nodal gene in H. roretzi is triggered by the contact of left epidermis with and sensing vitelline membrane proteins. However, due to aggregation of the vitelline membrane proteins, it was difficult to identify the responsible proteins by biochemical methods. Here, using comparative genomics approach and reporter gene assay, I tried to identify the cis-regulatory elements of nodal gene. Phylogenetic footprinting analyses between two ascidian species suggested the putative regulatory regions locate both in the upstream region and the first intron. The reporter gene assay showed that -249 to -156 upstream region and the first intron both play crucial role for the left-sided nodal expression. In silico analysis provided a list of candidate receptor proteins that could be activated by the vitelline membrane signal for specification of the left-right asymmetry via the nodal expression.

The above results provide fundamental information for further studies on transcriptional regulatory relationships during embryogenesis, and shed a light on our understanding of embryonic development.

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