Ca²⁺ Signalling and Membrane Dynamics During Cytokinesis in Animal Cells.

Interest in the role of Ca²⁺ signalling as a possible regulator of the combinatorial processes that result in the separation of the daughter cells during cytokinesis, extend back almost a 100 years. One of the key processes required for the successful completion of cytokinesis in animal cells (especially in the large holoblastically and meroblastically dividing embryonic cells of a number of amphibian and fish species), is the dynamic remodelling of the plasma membrane. Ca²⁺ signalling was subsequently demonstrated to regulate various different aspects of cytokinesis in animal cells, and so here we focus specifically on the role of Ca²⁺ signalling in the remodelling of the plasma membrane. We begin by providing a brief history of the animal models used and the research accomplished by the early twentieth century investigators, with regards to this aspect of animal cell cytokinesis. We then review the most recent progress made (i.e., in the last 10 years), which has significantly advanced our current understanding on the role of cytokinetic Ca²⁺ signalling in membrane remodelling. To this end, we initially summarize what is currently known about the Ca²⁺ transients generated during animal cell cytokinesis, and then we describe the latest findings regarding the source of Ca²⁺ generating these transients. Finally, we review the current evidence about the possible targets of the different cytokinetic Ca²⁺ transients with a particular emphasis on those that either directly or indirectly affect plasma membrane dynamics. With regards to the latter, we discuss the possible role of the early Ca²⁺ signalling events in the deformation of the plasma membrane at the start of cytokinesis (i.e., during furrow positioning), as well as the role of the subsequent Ca²⁺ signals in the trafficking and fusion of vesicles, which help to remodel the plasma membrane during the final stages of cell division. As it is becoming clear that each of the cytokinetic Ca²⁺ transients might have multiple, integrated targets, deciphering the precise role of each transient represents a significant (and ongoing) challenge.