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• W2887287789 abstract "The orientation of cell division is crucial in many developmental processes. In this issue of Developmental Cell, Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar report a myosin-dependent mechanism whereby physical contact between cells influences the division plane, with implications for understanding how cell division orientation is regulated in multicellular contexts. The orientation of cell division is crucial in many developmental processes. In this issue of Developmental Cell, Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar report a myosin-dependent mechanism whereby physical contact between cells influences the division plane, with implications for understanding how cell division orientation is regulated in multicellular contexts. The orientation of the cell division plane is critical for development (di Pietro et al., 2016di Pietro F. Echard A. Morin X. Regulation of mitotic spindle orientation: an integrated view.EMBO Rep. 2016; 17: 1106-1130Crossref PubMed Scopus (176) Google Scholar, Morin and Bellaïche, 2011Morin X. Bellaïche Y. Mitotic spindle orientation in asymmetric and symmetric cell divisions during animal development.Dev. Cell. 2011; 21: 102-119Abstract Full Text Full Text PDF PubMed Scopus (312) Google Scholar, Rose and Gonczy, 2014Rose L. Gonczy P. Polarity establishment, asymmetric division and segregation of fate determinants in early C. elegans embryos.WormBook. 2014; https://doi.org/10.1895/wormbook.1.30.2Crossref Scopus (118) Google Scholar, Werts and Goldstein, 2011Werts A.D. Goldstein B. How signaling between cells can orient a mitotic spindle.Semin. Cell Dev. Biol. 2011; 22: 842-849Crossref PubMed Scopus (21) Google Scholar). During asymmetric divisions, the division plane must be coordinated with cytoplasmic asymmetry, so that molecules are differentially segregated to daughter cells to specify fates. Of equal import, the division plane determines daughter cell position, thus contributing to cell signaling events that affect fate or proliferation. In the current issue of Developmental Cell, Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar identify a mechanism in which simple physical contact—with a cell or even an inert bead—causes a twisting movement of the entire cell, thus altering the final division plane. Further, they show that this effect is modulated by cell signaling. The findings have important implications for understanding how multiple cues, both physical and molecular, must be coordinated in multicellular contexts to achieve proper division patterns. In animal cells, the mitotic spindle specifies the site of the cleavage furrow, and thus, many studies of division orientation have focused on spindle positioning. Most mechanisms involve the microtubule motor dynein, which is tethered to the cortex or membrane and exerts pulling forces on the astral microtubules of the spindle (di Pietro et al., 2016di Pietro F. Echard A. Morin X. Regulation of mitotic spindle orientation: an integrated view.EMBO Rep. 2016; 17: 1106-1130Crossref PubMed Scopus (176) Google Scholar, Morin and Bellaïche, 2011Morin X. Bellaïche Y. Mitotic spindle orientation in asymmetric and symmetric cell divisions during animal development.Dev. Cell. 2011; 21: 102-119Abstract Full Text Full Text PDF PubMed Scopus (312) Google Scholar, Rose and Gonczy, 2014Rose L. Gonczy P. Polarity establishment, asymmetric division and segregation of fate determinants in early C. elegans embryos.WormBook. 2014; https://doi.org/10.1895/wormbook.1.30.2Crossref Scopus (118) Google Scholar, Werts and Goldstein, 2011Werts A.D. Goldstein B. How signaling between cells can orient a mitotic spindle.Semin. Cell Dev. Biol. 2011; 22: 842-849Crossref PubMed Scopus (21) Google Scholar). Intrinsic polarity cues, such as the PAR proteins, and extrinsic signals, such as Wnt, can regulate the force machinery to orient the spindle on a specific cell or tissue axis (see Figure 1 for examples). Despite progress in elucidating the details of microtubule-based pathways, much remains to be learned about how division patterns are specified during development. Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar now explore division patterning further in the C. elegans embryo, whose division pattern is invariant (Figure 1A). Asymmetric divisions of the P cells are dynein dependent, while the molecular basis for the orientation of the symmetrically dividing AB cell is not understood. The overall pattern of cell arrangements in the embryo has been assumed to result from spindle positioning combined with constraints of the eggshell (Figure 1A) (Rose and Gonczy, 2014Rose L. Gonczy P. Polarity establishment, asymmetric division and segregation of fate determinants in early C. elegans embryos.WormBook. 2014; https://doi.org/10.1895/wormbook.1.30.2Crossref Scopus (118) Google Scholar). Recently, however, the ABa and ABp cells were found to exhibit chiral flows of cortical non-muscle myosin (movements oriented perpendicular to the cell’s long axis), which produces a late tilt of the division axis; this tilt changes cell contacts and specifies all ensuing left-right asymmetries in the worm (Figure 1A) (Naganathan et al., 2014Naganathan S.R. Fürthauer S. Nishikawa M. Jülicher F. Grill S.W. Active torque generation by the actomyosin cell cortex drives left-right symmetry breaking.Elife. 2014; 3: e04165Crossref PubMed Scopus (133) Google Scholar, Pohl and Bao, 2010Pohl C. Bao Z. Chiral forces organize left-right patterning in C. elegans by uncoupling midline and anteroposterior axis.Dev. Cell. 2010; 19: 402-412Abstract Full Text Full Text PDF PubMed Scopus (78) Google Scholar). This finding, and knowledge that myosin flows can generate force, prompted Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar to examine a role for myosin in the AB division. The authors first showed that loss of microtubules had little effect on the final cleavage plane of AB relative to it neighboring cell P1 and confirmed that a component of the dynein force machine was not required. They then isolated the AB cell and cultured it alone or in contact with P1, using fluorescently tagged proteins to follow the mitotic spindle and cortical myosin. The initial spindle position and cleavage plane of AB and P1 were random relative to each other. However, as division continued, the entire AB cell rotated to align parallel with the P1 cell. Even more amazing, this behavior could be induced by an inert bead (Figure 1B). Using the bead assay, the authors went on to dissect the mechanism involved. Bead-induced division orientation required myosin, but not microtubules. Analysis of cortical myosin showed that the AB cell exhibits chiral myosin flows, as well as flow toward the equator (where myosin accumulates for cleavage). Significantly, cells with or without bead contact exhibited similar chiral flows, but only AB-bead combinations exhibited asymmetric equatorial flows (Figure 1B). Further, bead contact correlated with reduced levels of active myosin at the adjacent cortex. These results indicate that contact can regulate myosin, independent of cell signaling. Based on their findings, the authors propose that unequal myosin cortical forces result in a torque that rotates the entire cell, changing the final cleavage plane (Figure 1B). The AB cell also exhibited asymmetric myosin flows in vivo, but there the AB spindle is already aligned in parallel (Figure 1A). Thus, rather than being the main mechanism for specifying the division axis, torque may compensate for perturbations to spindle position, ensuring a robust division pattern. However, myosin flows may play a major role when spindle positioning is less regulated. Significantly, the authors showed that mouse embryo cells also react to cell contact. Initial spindle positions in mouse embryos are random, but the four-cell arrangement is reproducible (Figure 1C), leaving open the question of how the pattern is specified (Chen et al., 2018Chen Q. Shi J. Tao Y. Zernicka-Goetz M. Tracing the origin of heterogeneity and symmetry breaking in the early mammalian embryo.Nat. Commun. 2018; 9: 1819Crossref PubMed Scopus (51) Google Scholar). Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar showed that the mouse AB cell divides in parallel with its sister or a bead, even in the absence of the zona covering; contact downregulated myosin in these cells also. Although mammalian embryos do not undergo early asymmetric divisions, the four-cell arrangement appears important for gene expression (Chen et al., 2018Chen Q. Shi J. Tao Y. Zernicka-Goetz M. Tracing the origin of heterogeneity and symmetry breaking in the early mammalian embryo.Nat. Commun. 2018; 9: 1819Crossref PubMed Scopus (51) Google Scholar). Another key finding of the study is that cell signaling can block the effects of myosin forces, at least in the EMS cell. In wild-type C. elegans, EMS divides asymmetrically along the anterior-posterior axis (Figure 1A); spindle positioning is dynein dependent, and Wnt signaling has been shown to be instructive in cell isolation experiments (Goldstein et al., 2006Goldstein B. Takeshita H. Mizumoto K. Sawa H. Wnt signals can function as positional cues in establishing cell polarity.Dev. Cell. 2006; 10: 391-396Abstract Full Text Full Text PDF PubMed Scopus (122) Google Scholar, Rose and Gonczy, 2014Rose L. Gonczy P. Polarity establishment, asymmetric division and segregation of fate determinants in early C. elegans embryos.WormBook. 2014; https://doi.org/10.1895/wormbook.1.30.2Crossref Scopus (118) Google Scholar, Werts and Goldstein, 2011Werts A.D. Goldstein B. How signaling between cells can orient a mitotic spindle.Semin. Cell Dev. Biol. 2011; 22: 842-849Crossref PubMed Scopus (21) Google Scholar). In the current study, the authors showed that either a bead or an AB cell induces EMS to divide in parallel, indicating that EMS has the myosin rotation machinery. However, when a wild-type P2 cell (but not a Wnt-deficient P2) was present (Figure 1B), the EMS division instead oriented toward P2. These results raise the intriguing possibility that in vivo, the Wnt pathway functions both to induce dynein-dependent spindle positioning and to inhibit cell torque caused by cell contacts. In summary, Sugioka and Bowerman, 2018Sugioka K. Bowerman B. Combinatorial contact cues specify cell division orientation by directing myosin flows.Dev. Cell. 2018; 46 (this issue): 257-270Abstract Full Text Full Text PDF PubMed Scopus (31) Google Scholar have identified a contact-based mechanism that can change the cell division plane and thus affect cell positioning. This mechanism may work together with, or in opposition to, microtubule-based spindle positioning to diversify division patterns. Major challenges for the future are to identify the pathway that transduces physical contact cues to the myosin machinery and to understand how this mechanism is coordinated with others in vivo to produce proper division patterning during development. Combinatorial Contact Cues Specify Cell Division Orientation by Directing Cortical Myosin FlowsSugioka et al.Developmental CellJuly 19, 2018In BriefAnimal morphogenesis requires diverse orientations of cell division. However, how this division axis diversity is achieved remains to be elucidated. Sugioka et al. document cell contact-dependent mechanisms that diversify cell division axes by modulating cortical myosin flow and show that these mechanisms function in both C. elegans and mouse embryos. Full-Text PDF Open Archive" @default.
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• W2887287789 title "Cell Contact Gives a Twist to Cell Division" @default.
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