Pattern formation in zebrafish--fruitful liaisons between embryology and genetics.

Vertebrate embryos, despite quite diverse early morphologies, appear to employ similar cellular strategies and conserved biochemical pathways in their development (Eyal-Giladi, 1997). In the past decade, a small tropical teleost, zebrafish (Danio rerio), became an important model system in which to study development (Streisinger et al., 1981). By combining embryology with molecular and classical genetic methods, our understanding of early inductive and morphogenetic events during vertebrate embryogenesis significantly advanced. In zebrafish, dorsal-ventral polarity is established during early cleavage and is dependent on microtubular transport of determinants from the vegetal pole to the blastomeres positioned on top of the yolk cell. The syncytium forming from these marginal blastomeres in the early blastula exhibits dorsal-ventral asymmetry with beta-catenin localized to the nuclei on the presumptive dorsal side of the syncytium. The yolk cell is a source of signals that induce and pattern overlying blastoderm. Therefore, the dorsal yolk syncytial layer is equivalent to the Nieuwkoop center of the amphibian embryo. The embryonic shield, a thickening of the dorsal blastoderm margin, exhibits properties similar to the amphibian Spemann organizer. However, certain inductive and patterning signals from the organizer might be produced before the shield forms or might originate outside of the shield. Similar to the amphibian embryo, the key patterning functions of the fish dorsal organizer (i.e., dorsalization of mesoderm, ectoderm, and coordination of gastrulation movements) are performed by secreted molecules that antagonize the ventralizing activity of the swil (zbmp-2) and zbmp-4 gene products expressed on the ventral side of the embryo. These functions of the dorsal organizer require the activity of the chordino gene (a zebrafish homologue of chordin), bozozok, mercedes and ogon loci.