Mechanisms of cell polarization

Robinson, 1975, Polarizing fucoid eggs drive a calcium current through themselves, Science, 187, 70, 10.1126/science.1167318 Csikász-Nagy, 2013, Projecting cell polarity into the next decade, Phil Trans R Soc B, 368, 20130001, 10.1098/rstb.2013.0001 Onsum, 2007, A mathematical model for neutrophil gradient sensing and polarization, PLoS Comput Biol, 3, e36, 10.1371/journal.pcbi.0030036 Marée, 2006, Polarization and movement of keratocytes: a multiscale modelling approach, Bull Math Biol, 68, 1169, 10.1007/s11538-006-9131-7 Marée, 2012, How cells integrate complex stimuli: the effect of feedback from phosphoinositides and cell shape on cell polarization and motility, PLoS Comput Biol, 8, e1002402, 10.1371/journal.pcbi.1002402 Wolgemuth, 2010, The moving boundary node method: a level set-based, finite volume algorithm with application to cell motility, J Comput Phys, 229, 7287, 10.1016/j.jcp.2010.06.014 Shi, 2013, Interaction of motility, directional sensing, and polarity modules recreates the behaviors of chemotaxing cells, PLoS Comput Biol, 9, e1003122, 10.1371/journal.pcbi.1003122 Shao, 2010, Computational model for cell morphodynamics, Phys Rev Lett, 105, 108104, 10.1103/PhysRevLett.105.108104 Shao, 2012, Coupling actin flow, adhesion, and morphology in a computational cell motility model, Proc Natl Acad Sci U S A, 109, 6851, 10.1073/pnas.1203252109 Bourne, 2002, A chemical compass, Nature, 419, 21, 10.1038/419021a Ridley, 2003, Cell migration: integrating signals from front to back, Science, 302, 1704, 10.1126/science.1092053 Levchenko, 2002, Models of eukaryotic gradient sensing: application to chemotaxis of amoebae and neutrophils, Biophys J, 82, 50, 10.1016/S0006-3495(02)75373-3 Schneider, 2006, Mechanisms of gradient sensing and chemotaxis: conserved pathways, diverse regulation, Cell Cycle, 5, 1130, 10.4161/cc.5.11.2770 King, 2009, Chemotaxis: finding the way forward with Dictyostelium, Trends Cell Biol, 19, 523, 10.1016/j.tcb.2009.07.004 Skoge, 2014, Cellular memory in eukaryotic chemotaxis, Proc Natl Acad Sci U S A, 111, 14448, 10.1073/pnas.1412197111 Song, 2006, Dictyostelium discoideum chemotaxis: threshold for directed motion, Eur J Cell Biol, 85, 981, 10.1016/j.ejcb.2006.01.012 Lan, 2015, A biomechanical model for cell polarization and intercalation during Drosophila germband extension, Phys Biol, 12, 56011, 10.1088/1478-3975/12/5/056011 Bryant, 2014, A molecular switch for the orientation of epithelial cell polarization, 31, 171 Gord, 2014, Computational modelling of epidermal stratification highlights the importance of asymmetric cell division for predictable and robust layer formation, J R Soc Interface, 11, 10.1098/rsif.2014.0631 Tarle, 2015, Modeling the finger instability in an expanding cell monolayer, Integr Biol, 7, 1218, 10.1039/C5IB00092K Camley, 2016, Collective signal processing in cluster chemotaxis: roles of adaptation, amplification, and co-attraction in collective guidance, PLoS Comput Biol, 12, 10.1371/journal.pcbi.1005008 Halaoui, 2014, Rewiring cell polarity signaling in cancer, Oncogene, 34, 1 Byrne, 2016, Bistability in the Rac1, PAK and RhoA signalling network is a feature of cell motility, Cell Syst, 2, 38, 10.1016/j.cels.2016.01.003 Parri, 2010, Rac and Rho GTPases in cancer cell motility control, Cell Commun Signal, 8, 23, 10.1186/1478-811X-8-23 Franca-Koh, 2004, Moving forward: mechanisms of chemoattractant gradient sensing, Physiology, 19, 300, 10.1152/physiol.00017.2004 Comer, 2006, Phosphoinositide 3-kinase activity controls the chemoattractant-mediated activation and adaptation of adenylyl cyclase, Mol Biol Cell, 17, 357, 10.1091/mbc.e05-08-0781 Leslie, 2008, Understanding PTEN regulation: PIP2, polarity and protein stability, Oncogene, 27, 5464, 10.1038/onc.2008.243 Park, 2007, Central roles of small GTPases in the development of cell polarity in yeast and beyond, Microbiol Mol Biol Rev, 71, 48, 10.1128/MMBR.00028-06 Etienne-Manneville, 2008, Polarity proteins in migration and invasion, Oncogene, 27, 6970, 10.1038/onc.2008.347 Jilkine, 2011, A comparison of mathematical models for polarization of single eukaryotic cells in response to guided cues, PLoS Comput Biol, 7, e1001121, 10.1371/journal.pcbi.1001121 Levine, 2013, The physics of eukaryotic chemotaxis, Phys Today, 66, 24, 10.1063/PT.3.1884 Tsujita, 2015, Feedback regulation between plasma membrane tension and membrane-bending proteins organizes cell polarity during leading edge formation, Nat Cell Biol, 17, 749, 10.1038/ncb3162 Houk, 2012, Membrane tension maintains cell polarity by confining signals to the leading edge during neutrophil migration, Cell, 148, 175, 10.1016/j.cell.2011.10.050 Yam, 2007, Actin–myosin network reorganization breaks symmetry at the cell rear to spontaneously initiate polarized cell motility, J Cell Biol, 178, 1207, 10.1083/jcb.200706012 Dalous, 2008, Reversal of cell polarity and actin-myosin cytoskeleton reorganization under mechanical and chemical stimulation, Biophys J, 94, 1063, 10.1529/biophysj.107.114702 Weber, 2012, A mechanoresponsive cadherin-keratin complex directs polarized protrusive behavior and collective cell migration, Dev Cell, 22, 104, 10.1016/j.devcel.2011.10.013 Allen, 2013, Electrophoresis of cellular membrane components creates the directional cue guiding keratocyte galvanotaxis, Curr Biol, 23, 560, 10.1016/j.cub.2013.02.047 Shanley, 2006, Influx of extracellular Ca2+ is necessary for electrotaxis in Dictyostelium, J Cell Sci, 119, 4741, 10.1242/jcs.03248 Zhao, 2006, Electrical signals control wound healing through phosphatidylinositol-3-OH kinase-gamma and PTEN, Nature, 442, 457, 10.1038/nature04925 Lo, 2000, Cell movement is guided by the rigidity of the substrate, Biophys J, 79, 144, 10.1016/S0006-3495(00)76279-5 Van Norman, 2016, Asymmetry and cell polarity in root development, Dev Biol, 419, 165, 10.1016/j.ydbio.2016.07.009 Grieneisen, 2007, Auxin transport is sufficient to generate a maximum and gradient guiding root growth, Nature, 449, 1008, 10.1038/nature06215 Abley, 2013, An intracellular partitioning-based framework for tissue cell polarity in plants and animals, Development, 140, 2061, 10.1242/dev.062984 Andrew, 2007, Chemotaxis in shallow gradients is mediated independently of PtdIns 3-kinase by biased choices between random protrusions, Nat Cell Biol, 9, 193, 10.1038/ncb1536 Gierer, 1972, A theory of biological pattern formation, Kybernetik, 12, 30, 10.1007/BF00289234 Turing, 1952, The chemical theory of morphogenesis, Phil Trans Roy Soc, 237, 37 Meinhardt, 1999, Orientation of chemotactic cells and growth cones: models and mechanisms, J Cell Sci, 112, 2867, 10.1242/jcs.112.17.2867 Parent, 1999, A cell's sense of direction, Science, 284, 765, 10.1126/science.284.5415.765 Ma, 2004, Two complementary, local excitation, global inhibition mechanisms acting in parallel can explain the chemoattractant-induced regulation of PI(3,4,5)P-3 response in Dictyostelium cells, Biophys J, 87, 3764, 10.1529/biophysj.104.045484 Iglesias, 2008, Navigating through models of chemotaxis, Curr Opin Cell Biol, 20, 35, 10.1016/j.ceb.2007.11.011 Iglesias, 2011, Biased excitable networks: how cells direct motion in response to gradients, Curr Opin Cell Biol, 24, 245, 10.1016/j.ceb.2011.11.009 Xiong, 2010, Cells navigate with a local-excitation, global-inhibition-biased excitable network, Proc Natl Acad Sci U S A, 107, 17079, 10.1073/pnas.1011271107 Levine, 2006, Directional sensing in eukaryotic chemotaxis: a balanced inactivation model, PNAS, 103, 9761, 10.1073/pnas.0601302103 Xu, 2003, Divergent signals and cytoskeletal assemblies regulate self-organizing polarity in neutrophils, Cell, 114, 201, 10.1016/S0092-8674(03)00555-5 Wedlich-Soldner, 2003, Spontaneous cell polarization through actomyosin-based delivery of the Cdc42 GTPase, Science, 299, 1231, 10.1126/science.1080944 Otsuji, 2007, A mass conserved reaction-diffusion system captures properties of cell polarity, PLoS Comput Biol, 3, e108, 10.1371/journal.pcbi.0030108 Lin, 2012, Synthetic spatially graded Rac activation drives cell polarization and movement, Proc Natl Acad Sci U S A, 109, E3668, 10.1073/pnas.1210295109 Dawes, 2007, Phosphoinositides and Rho proteins spatially regulate actin polymerization to initiate and maintain directed movement in a one-dimensional model of a motile cell, Biophys J, 92, 744, 10.1529/biophysj.106.090514 Jilkine, 2007, Mathematical model for spatial segregation of the Rho-family GTPases based on inhibitory crosstalk, Bull Math Biol, 69, 1943, 10.1007/s11538-007-9200-6 Goryachev, 2008, Dynamics of Cdc42 network embodies a Turing-type mechanism of yeast cell polarity, FEBS Lett, 582, 1437, 10.1016/j.febslet.2008.03.029 Holmes, 2016, Analysis of a minimal Rho-GTPase circuit regulating cell shape, Phys Biol, 13, 10.1088/1478-3975/13/4/046001 Guilluy, 2011, Rho protein crosstalk: another social network?, Trends Cell Biol, 21, 718, 10.1016/j.tcb.2011.08.002 Bosgraaf, 2009, Navigation of chemotactic cells by parallel signaling to pseudopod persistence and orientation, PLoS One, 4, e6842, 10.1371/journal.pone.0006842 Hoeller, 2007, Chemotaxis in the absence of PIP3 gradients, Curr Biol, 17, 813, 10.1016/j.cub.2007.04.004 Veltman, 2014, PIP(3)-dependent macropinocytosis is incompatible with chemotaxis, J Cell Biol, 204, 497, 10.1083/jcb.201309081 Skoge, 2010, Gradient sensing in defined chemotactic fields, Integr Biol, 2, 659, 10.1039/c0ib00033g Dertinger, 2001, Generation of gradients having complex shapes using microfluidic networks, Anal Chem, 73, 1240, 10.1021/ac001132d Irimia, 2006, Microfluidic system for measuring neutrophil migratory responses to fast switches of chemical gradients, Lab Chip, 6, 191, 10.1039/B511877H Lockley, 2014, Image based validation of dynamical models for cell reorientation, Cytom A, 87, 471, 10.1002/cyto.a.22600 Loomis, 2012, Innate non-specific cell substratum adhesion, PLoS One, 7, e42033, 10.1371/journal.pone.0042033 Fuller, 2010, External and internal constraints on eukaryotic chemotaxis, Proc Natl Acad Sci U S A, 107, 9656, 10.1073/pnas.0911178107 Prentice-Mott, 2016, Directional memory arises from long-lived cytoskeletal asymmetries in polarized chemotactic cells, Proc Natl Acad Sci U S A, 113, 1267, 10.1073/pnas.1513289113 Doyle, 2009, One-dimensional topography underlies three-dimensional fibrillar cell migration, J Cell Biol, 184, 481, 10.1083/jcb.200810041 Lin, 2015, Interplay between chemotaxis and contact inhibition of locomotion determines exploratory cell migration, Nat Commun, 6, 6619, 10.1038/ncomms7619 Carmona-Fontaine, 2008, Contact inhibition of locomotion in vivo controls neural crest directional migration, Nature, 456, 957, 10.1038/nature07441 Abercrombie, 1953, Observations on the social behaviour of cells in tissue culture. I. Speed of movement of chick heart fibroblasts in relation to their mutual contacts, Exp Cell Res, 5, 111, 10.1016/0014-4827(53)90098-6 Theveneau, 2010, Integrating chemotaxis and contact-inhibition during collective cell migration: small GTPases at work, Small GTPases, 1, 113, 10.4161/sgtp.1.2.13673 Theveneau, 2010, Collective chemotaxis requires contact-dependent cell polarity, Dev Cell, 19, 39, 10.1016/j.devcel.2010.06.012 Nagel, 2014, Geometry-driven polarity in motile amoeboid cells, PLoS One, 9, e113382, 10.1371/journal.pone.0113382 Irimia, 2009, Spontaneous migration of cancer cells under conditions of mechanical confinement, Integr Biol, 1, 506, 10.1039/b908595e Boneschansker, 2014, Microfluidic platform for the quantitative analysis of leukocyte migration signatures, Nat Commun, 5, 4787, 10.1038/ncomms5787 Hamza, 2014, Retrotaxis of human neutrophils during mechanical confinement inside microfluidic channels, Integr Biol, 6, 175, 10.1039/C3IB40175H Boyden, 2005, Millisecond-timescale, genetically targeted optical control of neural activity, Nat Neurosci, 8, 1263, 10.1038/nn1525 Levskaya, 2009, Spatiotemporal control of cell signalling using a light-switchable protein interaction, Nature, 461, 997, 10.1038/nature08446 Wu, 2009, A genetically encoded photoactivatable Rac controls the motility of living cells, Nature, 461, 104, 10.1038/nature08241 Toettcher, 2013, Using optogenetics to interrogate the dynamic control of signal transmission by the Ras/Erk module, Cell, 155, 1422, 10.1016/j.cell.2013.11.004 Wang, 2010, Light-mediated activation reveals a key role for Rac in collective guidance of cell movement in vivo, Nat Cell Biol, 12, 591, 10.1038/ncb2061 Machacek, 2009, Coordination of Rho GTPase activities during cell protrusion, Nature, 461, 99, 10.1038/nature08242 Jost, 2015, Probing yeast polarity with acute, reversible, optogenetic inhibition of protein function, ACS Synth Biol, 4, 1077, 10.1021/acssynbio.5b00053 Lee, 2014, Reversible protein inactivation by optogenetic trapping in cells, Nature, 11, 633 Mori, 2008, Wave-pinning and cell polarity from a bistable reaction-diffusion system, Biophys J, 94, 3684, 10.1529/biophysj.107.120824 Mori, 2011, Asymptotic and bifurcation analysis of wave-pinning in a reaction-diffusion model of cell polarization, SIAM J Appl Math, 71, 1401, 10.1137/10079118X Walther, 2012, Deterministic versus stochastic cell polarisation through wave-pinning, Bull Math Biol, 74, 2570 Holmes, 2015, Local Perturbation analysis: a computational tool for biophysical reaction-diffusion models, Biophys J, 108, 230, 10.1016/j.bpj.2014.11.3457 Nguyen, 2015, DYVIPAC: an integrated analysis and visualisation framework to probe multi-dimensional biological networks, Sci Rep, 5, 12569, 10.1038/srep12569 Neilson, 2011, Chemotaxis: a feedback-based computational model robustly predicts multiple aspects of real cell behaviour, PLoS Biol, 9, e1000618, 10.1371/journal.pbio.1000618 Graner, 1992, Simulation of biological cell sorting using a two-dimensional extended Potts model, Phys Rev Lett, 69, 2013, 10.1103/PhysRevLett.69.2013 Rappel, 1999, Self-organized vortex state in two-dimensional Dictyostelium dynamics, Phys Rev Lett, 83, 1247, 10.1103/PhysRevLett.83.1247 Szabó, 2010, Collective cell motion in endothelial monolayers, Phys Biol, 7, 46007, 10.1088/1478-3975/7/4/046007 Kabla, 2012, Collective cell migration: leadership, invasion and segregation, J R Soc Interface, 9, 3268, 10.1098/rsif.2012.0448 Niculescu, 2015, Crawling and gliding: a computational model for shape-driven cell migration, PLoS Comput Biol, 11, e1004280, 10.1371/journal.pcbi.1004280 Karma, 1996, Phase-field method for computationally efficient modeling of solidification with arbitrary interface kinetics, Phys Rev E Stat Phys Plasmas Fluids Relat Interdiscip Top, 53, R3017 Karma, 2001, Phase-field model of mode III dynamic fracture, Phys Rev Lett, 87, 45501, 10.1103/PhysRevLett.87.045501 Fenton, 2005, Modeling wave propagation in realistic heart geometries using the phase-field method, Chaos, 15, 10.1063/1.1840311 Folch, 1999, Phase-field model for Hele-Shaw flows with arbitrary viscosity contrast. I. Theoretical approach, Phys Rev E, 60, 1724, 10.1103/PhysRevE.60.1724 Kockelkoren, 2003, Computational approach for modeling intra- and extracellular dynamics, Phys Rev E Stat Nonlin Soft Matter Phys, 68, 37702, 10.1103/PhysRevE.68.037702 Li, 2009, Solving PDEs in complex geometries: a diffuse domain approach, Commun Math Sci, 7, 81, 10.4310/CMS.2009.v7.n1.a4 Camley, 2014, Polarity mechanisms such as contact inhibition of locomotion regulate persistent rotational motion of mammalian cells on micropatterns, Proc Natl Acad Sci U S A, 111, 14770, 10.1073/pnas.1414498111 Kulawiak, 2016, Modeling contact inhibition of locomotion of colliding cells migrating on micropatterned substrates, PLoS Comput Biol, 12, e1005239, 10.1371/journal.pcbi.1005239 Nonomura, 2012, Study on multicellular systems using a phase field model, PLoS One, 7, e33501, 10.1371/journal.pone.0033501 Marth, 2016, Collective migration under hydrodynamic interactions–a computational approach, Interface Focus, 6, 20160037, 10.1098/rsfs.2016.0037 Edelstein-Keshet, 2013, From simple to detailed models for cell polarization, Phil Trans R Soc B, 368, 20130003, 10.1098/rstb.2013.0003 Delile, 2017, A cell-based computational model of early embryogenesis coupling mechanical behaviour and gene regulation, Nat Commun, 8, 13929, 10.1038/ncomms13929 Hecht, 2011, “Self-assisted” amoeboid navigation in complex environments, PLoS One, 6, e21955, 10.1371/journal.pone.0021955 Dalle Nogare, 2014, Leading and trailing cells cooperate in collective migration of the zebrafish posterior lateral line primordium, Development, 141, 3188, 10.1242/dev.106690 Tweedy, 2016, Self-generated chemoattractant gradients: attractant depletion extends the range and robustness of chemotaxis, PLoS Biol, 14, e1002404, 10.1371/journal.pbio.1002404 Venkiteswaran, 2013, Generation and dynamics of an endogenous, self-generated signaling gradient across a migrating tissue, Cell, 155, 674, 10.1016/j.cell.2013.09.046 Streichan, 2011, Collective cell migration guided by dynamically maintained gradients, Phys Biol, 8, 45004, 10.1088/1478-3975/8/4/045004 Poujade, 2007, Collective migration of an epithelial monolayer in response to a model wound, Proc Natl Acad Sci, 104, 15988, 10.1073/pnas.0705062104 Yamaguchi, 2015, Leader cells regulate collective cell migration via Rac activation in the downstream signaling of integrin β1 and PI3K, Sci Rep, 5, 7656, 10.1038/srep07656 Revenu, 2014, Quantitative cell polarity imaging defines leader-to-follower transitions during collective migration and the key role of microtubule-dependent adherens junction formation, Development, 141, 1282, 10.1242/dev.101675 Stramer, 2016, Mechanisms and in vivo functions of contact inhibition of locomotion, Nat Rev Mol Cell Biol, 18, 43, 10.1038/nrm.2016.118 Camley, 2014, Velocity alignment leads to high persistence in confined cells, Phys Rev E Stat Nonlin Soft Matter Phys, 89, 62705, 10.1103/PhysRevE.89.062705 Zimmermann, 2016, Contact inhibition of locomotion determines cell-cell and cell-substrate forces in tissues, Proc Natl Acad Sci U S A, 113, 2660, 10.1073/pnas.1522330113 Devenport, 2014, The cell biology of planar cell polarity, J Cell Biol, 207, 171, 10.1083/jcb.201408039 Zallen, 2007, Planar polarity and tissue morphogenesis, Cell, 129, 1051, 10.1016/j.cell.2007.05.050 Dawes, 2011, PAR-3 oligomerization may provide an actin-independent mechanism to maintain distinct par protein domains in the early Caenorhabditis elegans embryo, Biophys J, 101, 1412, 10.1016/j.bpj.2011.07.030 Goehring, 2011, PAR proteins diffuse freely across the anterior–posterior boundary in polarized C. elegans embryos, J Cell Biol, 193, 583, 10.1083/jcb.201011094 Meinhardt, 2007, Computational modelling of epithelial patterning, Curr Opin Genet Dev, 17, 272, 10.1016/j.gde.2007.05.004 Amonlirdviman, 2005, Mathematical modeling of planar cell polarity to understand domineering nonautonomy, Science, 307, 423, 10.1126/science.1105471 Axelrod, 2013, Mathematical modeling of planar cell polarity signaling, 27 Gao, 2013, Multiscale modeling and analysis of planar cell polarity in the Drosophila wing, IEEE/ACM Trans Comput Biol Bioinforma, 10, 337, 10.1109/TCBB.2012.101 Iden, 2008, Crosstalk between small GTPases and polarity proteins in cell polarization, 9, 846 Nishimura, 2005, PAR-6–PAR-3 mediates Cdc42-induced Rac activation through the Rac GEFs STEF/Tiam1, Nat Cell Biol, 7, 270, 10.1038/ncb1227 Hetmanski, 2016, A MAPK-Driven feedback loop suppresses Rac activity to promote RhoA-driven cancer cell invasion, PLoS Comput Biol, 12, e1004909, 10.1371/journal.pcbi.1004909 Kim, 2015, Network-based identification of feedback modules that control RhoA activity and cell migration, J Mol Cell Biol, 7, 242 Chau, 2012, Designing synthetic regulatory networks capable of self-organizing cell polarization, Cell, 151, 320, 10.1016/j.cell.2012.08.040 Guetta-Terrier, 2015, Protrusive waves guide 3D cell migration along nanofibers, J Cell Biol, 211, 683, 10.1083/jcb.201501106 Giese, 2015, Influence of cell shape, inhomogeneities and diffusion barriers in cell polarization models, Phys Biol, 12, 66014, 10.1088/1478-3975/12/6/066014 Holmes, 2012, A comparison of computational models for eukaryotic cell shape and motility, PLoS Comput Biol, 8, e1002793, 10.1371/journal.pcbi.1002793 Camley, 2017, Crawling and turning in a minimal reaction-diffusion cell motility model: coupling cell shape and biochemistry, Phys Rev E, 95, 012401, 10.1103/PhysRevE.95.012401 Tuncer, 2017, Projected finite elements for systems of reaction-diffusion equations on closed evolving spheroidal surfaces, Commun Comput Phys, 21, 718, 10.4208/cicp.OA-2016-0029 Sailem, 2014, Cross-talk between Rho and Rac GTPases drives deterministic exploration of cellular shape space and morphological heterogeneity, Open Biol, 4, 130132, 10.1098/rsob.130132 Cooper, 2015, Apolar and polar transitions drive the conversion between amoeboid and mesenchymal shapes in melanoma cells, Mol Biol Cell, 26, 4163, 10.1091/mbc.E15-06-0382 Boureux, 2007, Evolution of the Rho family of ras-like GTPases in eukaryotes, Mol Biol Evol, 24, 203, 10.1093/molbev/msl145 Takeda, 2012, Incoherent feedforward control governs adaptation of activated ras in a eukaryotic chemotaxis pathway, Sci Signal, 5, ra2, 10.1126/scisignal.2002413 Nakajima, 2014, Rectified directional sensing in long-range cell migration, Nat Commun, 5, 5367, 10.1038/ncomms6367 Zihni, 2015, RhoGTPase signalling at epithelial tight junctions: bridging the GAP between polarity and cancer, Int J Biochem Cell Biol, 64, 120, 10.1016/j.biocel.2015.02.020 McCormack, 2013, Cycling around cell-cell adhesion with Rho GTPase regulators, J Cell Sci, 126, 379, 10.1242/jcs.097923 Vega, 2015, The Rho GTPase RhoB regulates cadherin expression and epithelial cell-cell interaction, Cell Commun Signal, 13, 1, 10.1186/s12964-015-0085-y Reffay, 2014, Interplay of RhoA and mechanical forces in collective cellmigration driven by leader cells, Nat Cell Biol, 16, 217, 10.1038/ncb2917