Patterning and lineage specification in the amphibian embryo.

Xenopus has been widely used to study early embryogenesis because the embryos allow for efficient functional assays of gene products by the overexpression of RNA. The first asymmetry of the embryo is initiated during oogenesis and is manifested by the darkly pigmented animal hemisphere and lightly pigmented vegetal hemisphere. Upon fertilization a second asymmetry, the dorsal-ventral asymmetry, is established, with the sperm entry site defining the prospective ventral region. During the cleavage stage, a vegetal cortical cytoplasm (VCC)/beta-catenin signaling pathway is differentially activated on the prospective dorsal side of the embryo. The overlapping of the VCC/beta-catenin and transforming growth factor beta (TGF-beta) pathways in the dorsal vegetal quadrant specifies dorsal-vental axis formation by regulating formation of the Spemann organizer, including the anterior endomesoderm. The organizer initiates gastrulation to form a triploblastic embryo in which the mesoderm layer is located between the ectoderm layer and the endoderm layer. The interplay between maternal and zygotic TGF-beta s and the T-box transcription factors in the vegetal hemisphere initiates the specification of germ-layer lineages. TGF-beta signaling originating from the vegetal region induces mesoderm in the equatorial region, and initiates endoderm differentiation directly in the vegetal region. The ectoderm develops from the animal region, which does not come into contact with the vegetal TGF-beta signals. A large number of the downstream components and transcriptional targets of early developmental pathways have been identified and characterized. This review gives an overview of recent advances in the understanding of the functional roles and interactions of the molecular players important for axis determination and germ-layer specification during early Xenopus embryogenesis.