Formation and dynamics of cytoplasmic domains and their genetic regulation during the zebrafish oocyte-to-embryo transition.

Establishment and movement of cytoplasmic domains is of great importance for the emergence of cell polarity, germline segregation, embryonic axis specification and correct sorting of organelles and macromolecules into different embryonic cells. The zebrafish oocyte, egg and zygote are valuable material for the study of cytoplasmic domains formation and dynamics during development. In this review we examined how cytoplasmic domains form and are relocated during zebrafish early embryogenesis. Distinct cortical cytoplasmic domains (also referred to as ectoplasm domains) form first during early oogenesis by the localization of mRNAs to the vegetal or animal poles of the oocyte or radially throughout the cortex. Cytoplasmic segregation in the late oocyte relocates non-cortical cytoplasm (endoplasm) into the preblastodisc and yolk cell. The preblastodisc is a precursor to the blastodisc, which gives rise to the blastoderm and most the future embryo. After egg activation, the blastodisc enlarges by transport of cytoplasm from the yolk cell to the animal pole, along defined pathways or streamers that include a complex cytoskeletal meshwork and cytoplasmic movement at different speeds. A powerful actin ring, assembled at the margin of the blastodisc, appears to drive the massive streaming of cytoplasm. The fact that the mechanism(s) leading to the formation and relocation of cytoplasmic domains are affected in maternal-effect mutants indicates that these processes are under maternal control. Here, we also discuss why these mutants represent outstanding genetic entry points to investigate the genetic basis of cytoplasmic segregation. Functional studies, combined with the analysis of zebrafish mutants, generated by forward and reverse genetic strategies, are expected to decipher the molecular mechanism(s) by which the maternal factors regulate cytoplasmic movements during early vertebrate development.