Mechanisms of gastrulation and tail formation in ascidians.

Ascidian embryos are useful for examining how events that occur during fertilization and cleavage affect gastrulation because they gastrulate early in development, during the seventh cleavage. In ascidians, both dorsal-ventral and anterior-posterior axes are determined before first cleavage. The dorsal-ventral axis is fixed along the animal-vegetal axis of the fertilized egg following the first phase of ooplasmic segregation, perhaps due to determinants moved to the vegetal pole in concert with the myoplasm and plasma membrane components. The first ooplasmic movements appear to be driven by the actin network in the cortical myoplasm. The anterior-posterior axis becomes apparent after the second phase of ooplasmic segregation, when the cortical myoplasm becomes detached from the egg membrane, and moves to the posterior pole of the embryo. This movement is dependent on microtubules and has been attributed to the formation and movement of the sperm aster. A major component of the cortical myoplasm, p58, is co-localized along the microtubules emanating from the sperm aster. Gastrulation begins during the seventh cleavage with the invagination of the large endodermal cells at the vegetal pole of the embryo. The neural plate appears as a thickening of the epidermis on the dorsal side of the larva during the ninth cleavage; then the neural folds are formed, join, and close, elaborating the neural tube. Following neurulation, the tail is elongated as the neural tube and notochord cells intercalate at the midline of the embryo. Investigations using anural (tailless) ascidian larvae suggest that some of the processes underlying elongation can be restored by the zygotic genome. Although ascidian larvae contain fewer cells and cell types than vertebrate embryos, ascidian gastrulation and morphogenesis appear to employ similar mechanisms to those in vertebrate embryos. The extent of our current knowledge about the mechanisms involved in gastrulation and tail formation is summarized, and further experiments are suggested to explore the molecular mechanisms underlying these processes.