Evidence for a functional role of the cytoskeleton in determination of the dorsoventral axis in Xenopus laevis eggs.

A normal table of events of the first cleavage period in the fertilized egg (cf. Gerhart, 1980) has been completed (cf. Table I) by studying external and internal features. Through a cytological study of eggs fixed after video time-lapse observation such features can directly be correlated and it has been shown that the first postfertilization wave (PFW) reflects spermaster growth, which causes rearrangements of animal yolk material. This may, in conjunction with the interaction of the spermaster rays with the cortex, define, in time as well as in space, the asymmetric cortical contraction which we suppose to evoke asymmetry in the animal hemisphere by formation of the vitelline wall (Pasteels, 1964) and in the vegetal hemisphere by formation of the Vegetal Dorsalising Centre (Kirschner et al. 1981). Neither prick-activated eggs nor fertilized eggs incubated in vinblastine develop a spermaster. Under these conditions abnormal cytoplasmic segregation may be directed by gravity alone. For normal development the activated egg must in some way, for instance through the sperm centriole, organize microtubule assembly into a monaster. The centriole acts as a microtubule-organizing centre in structuring the egg's cytoskeleton, and through this directs localization of the various yolk components, in time as well as in space. In egg rotation experiments performed under appropriate conditions, the cytoskeleton is disturbed and yolk rearranges under gravity till a new equilibrium is established which determines a new dorsoventral polarity. Such experiments also show that neither the dorsal cytoplasm nor the grey crescent cortex act as the ultimate dorsal determinants, since their localization is unaltered upon rotation, whereas the overall yolk distribution is significantly changed.