Ray Rappaport chronology: Twenty-five years of seminal papers on cytokinesis in the Journal of Experimental Zoology

Between 1960 and 1993 Ray Rappaport (Fig. 1) published a series of seminal articles in the Journal of Experimental Zoology that defined the fundamental rules for how the mitotic apparatus determines the position of the cleavage furrow. This lasting achievement is recognized in every cell biology textbook and his story is told in every cell biology class. His legendary experiments remain fresh and his papers still make exciting reading. This appeal comes in part from Ray’s gift as a storyteller, but it’s mainly the events in these riveting stories that hold one’s attention. The work is so important that his strategies and ideas strongly influence current research on cell division. We still debate the meaning of his experiments. This remarkable accomplishment was achieved in a manner that has been all but lost in contemporary biological research. Ray did virtually all of his experiments with his own two hands during summers at the Mt. Desert Island Biological Laboratory in Salisbury Cove Maine. During the rest of the year he taught at Union College in Schenectady New York. The Mt. Desert Island Biological Laboratory is a rustic collection of wooden buildings, a world apart from the pretentious palaces designed by signature architects that house many biological research labs today. Ray’s equipment was as modest as the lab in Maine. It consisted of a Unitron inverted microscope, a deFonbrune microforge, a dissecting microscope, micromanipulators, a simple centrifuge, and various observation chambers that he fabricated himself from materials available at a hardware store. Ray used a rowboat to collect sand dollars from the local waters for his experiments. He never used an electron microscope or a fluorescence microscope. The work was done without large research grants, graduate students, postdoctoral fellows, or research assistants. A couple of undergraduates helped with two early papers. Ray’s wife Barbara was a constant source of support, helped with writing, and co-authored some later papers. Although Ray never worked with a gene, protein, or antibody, he defined the rules for how the mitotic apparatus stimulates the cortex to form a furrow. He showed that the asters of the mitotic spindle are the source of this positive signal (at least in echinoderm eggs), measured the rate that the stimulus moves from the mitotic asters to the cortex, identified the time required for the stimulus to make a lasting impression on the cortex, and discovered a latent period between receipt of the signal and the formation of a visible