Migration and division of cleavage nuclei in the gall midge,Wachtliella persicariae

Abstract

The control of nuclear division and migration was studied in time-lapse films of the multinucleate egg cell of a gall midge by experimental alterations of the mitotic pattern. During each cleavage cycle, a wave of randomly oriented saltations of yolk particles (WROS) is seen to travel through the ooplasm. This wave proved to be an indispensable prerequisite for the accompanying anaphase wave and for the activation of the nuclear migration cytasters: WROS cycles can occur autonomously without cleavage nuclei being present, but there is no anaphase without a WROS passing the dividing nucleus. WROSs and mitotic waves can be inverted, and the WROS cycles and the cleavage cycles can be desynchronized by temperature grandients or by locally impaired gas exchange. If a nucleus is not ready for anaphase when met by a WROS, it will only divide in the course of the next WROS. WROSs thus indicate autonomous anaphase-triggering waves governing the cleavage divisions. Rhythmic ooplasmic movements continue even if the WROSs as well as the nuclear divisions are inhibited by colchinine. The characteristics of the WROSs support the hypothesis that each of them is the visible effect of a wave of calcium release (similar to that established in vertebrate eggs) which acts locally on the microtubular system and may continue even if the WROSs are suppressed. The correlations between a possible calcium release, WROS activity, microtubule disassembly and nuclear cycle are discussed.