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Review
. 2016 Aug;73(16):3115-36.
doi: 10.1007/s00018-016-2220-3. Epub 2016 Apr 16.

The final cut: cell polarity meets cytokinesis at the bud neck in S. cerevisiae

Maria Angeles Juanes  1   2 , Simonetta Piatti  3
Affiliations
Review

The final cut: cell polarity meets cytokinesis at the bud neck in S. cerevisiae

Maria Angeles Juanes et al. Cell Mol Life Sci. 2016 Aug.

Abstract

Cell division is a fundamental but complex process that gives rise to two daughter cells. It includes an ordered set of events, altogether called "the cell cycle", that culminate with cytokinesis, the final stage of mitosis leading to the physical separation of the two daughter cells. Symmetric cell division equally partitions cellular components between the two daughter cells, which are therefore identical to one another and often share the same fate. In many cases, however, cell division is asymmetrical and generates two daughter cells that differ in specific protein inheritance, cell size, or developmental potential. The budding yeast Saccharomyces cerevisiae has proven to be an excellent system to investigate the molecular mechanisms governing asymmetric cell division and cytokinesis. Budding yeast is highly polarized during the cell cycle and divides asymmetrically, producing two cells with distinct sizes and fates. Many components of the machinery establishing cell polarization during budding are relocalized to the division site (i.e., the bud neck) for cytokinesis. In this review we recapitulate how budding yeast cells undergo polarized processes at the bud neck for cell division.

Keywords: Actomyosin ring; Budding yeast; Cytokinesis; Formins; Mitotic exit network; Septins.

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Figures

Fig. 1
Fig. 1
Spindle positioning relative to the polarity axis determines the outcome of cell division. The cartoon depicts a polarized cell, where a gradient of polarized factors increases from left to right (gray shadow). Depending on spindle positioning, which dictates the position of the cleavage furrow, cell division will be asymmetric or symmetric
Fig. 2
Fig. 2
Organization of the actin cytoskeleton during budding yeast cell cycle. The cartoon illustrates budding yeast cells at different cell cycle stages and the distribution of actin structures (patches, cables and contractile F-ring) therein. Depending on whether actin organization is polarized, cell growth can be apical (directed towards the tip of the bud) or isotropic (with the bud expanding in all directions). After a transient depolarization of actin patches and cables in mitosis, the actin cytoskeleton repolarizes in telophase to bring about cytokinesis. See text for details
Fig. 3
Fig. 3
Structural organization of the yeast formins Bni1 and Bnr1. Formins form a doughnut-shaped dimer that encircles the nascent actin filament during its elongation. The main interactors and regulators of each formin are depicted. See text for details. GBD GTPase-binding domain, DID diaphanous inhibitory domain, DD dimerization domain, CC coiled coil, SBD Spa2-binding domain, FH1/2 formin homology domain, DAD diaphanous auto-regulatory domain, BBD Bud6-binding domain
Fig. 4
Fig. 4
Main steps in budding yeast cytokinesis. See text for details
Fig. 5
Fig. 5
The septin ring during budding yeast cell cycle. Sequential stages of septin organization during the cell cycle of budding yeast
Fig. 6
Fig. 6
The mitotic exit network (MEN) and its regulation by the spindle position checkpoint (SPOC). a MEN signaling takes place mostly at SPBs, where the GTPase Tem1 in its active GTP-bound state promotes recruitment and activation of the Cdc15 protein kinase, which in turn recruits the Dbf2-Mob1 kinase complex that ultimately activates the Cdc14 phosphatase, thereby triggering mitotic exit and cytokinesis. Tem1 is kept inactive by the GTPase-activating protein Bfa1-Bub2 that can be inhibited by the polo kinase Cdc5, whose activity is counteracted by the kinase Kin4 in the mother cell. In turn, the Lte1 protein, which is localized specifically in the bud, restrains Kin4 in the mother compartment. b The MEN inhibitor Kin4 and the MEN activator Lte1 are spatially segregated in the mother and bud compartment, respectively (Kin4 red, Lte1 green). As long as an SPB has not moved into the bud, Tem1 and MEN are kept inactive, thereby coupling spindle positioning and nuclear division with mitotic exit
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