Unequal cell division regulated by the contents of germinal vesicles

Fertilization occurs during meiosis in many animals, when maternal centrosomes participate in the formation of spindles at the animal pole, which results in polar body formation. Paternal centrosomes do not participate in cell division during oocyte maturation. After meiosis, they form the spindles while the maternal centrosomes are discarded. It is unknown why paternal centrosomes do not form spindles during meiosis. Here, we show that the artificial incorporation of sperm at the animal pole of immature starfish oocytes causes unequal cell division and the formation of polar body-like fragments. The removal of germinal vesicles from the animal pole blocks the formation of polar body-like fragments. Furthermore, translocation of germinal vesicles to the vegetal pole by centrifugation induces the extrusion of polar body-like fragments from the vegetal pole, where sperm penetration is prerequisite. After germinal vesicle breakdown, cyclin B is localized in the maternal and paternal asters and spindles near the germinal vesicle. These results suggest that germinal vesicle components such as the cdc2-cyclin B complex interact with asters and spindles and can induce unequal cell division. During normal fertilization, paternal centrosomes are likely kept away from the germinal vesicle components, resulting in the inhibition of unequal paternal centrosome-dependent cell division.