Mature oocytes of the annelidan worm Pseudopotamilla occelata have a wide perivitelline space between the oocyte surface and the vitelline envelope and are arrested at the first metaphase (MI). We found a novel twostep Ca 2+ increase in normally fertilized oocytes. The first Ca 2+ increase originated at a cortex situated underneath a fertilizing sperm on the vitelline envelope, but failed to propagate beyond the center of the oocyte. The first localized Ca 2+ increase was then followed by a larger Ca 2+ increase starting from the whole oocyte cortex and spreading inwardly to the center. The first localized Ca 2+ increase at fertilization was suppressed by the phospholipase C inhibitor U73122, and a similar Ca 2+ change was induced by inositol 1,4,5-trisphosphate (IP 3 2+ increase induces the formation of large cytoplasmic protrusion, which helps the fertilizing sperm to enter the oocyte, whereas the following global Ca 2+ increase is a prerequisite for the retraction of the cytoplasmic protrusion and the resumption of meiosis from MI.