Abstract
The actomyosin cortex is a thin film, containing actin filaments and myosin molecular motors, located beneath the plasma-membrane of eukaryotic cells. Active processes, driven by ATP hydrolysis, can generate mechanical forces in the cortex. Coordinated force-generation drives large-scale mechanical flows and orientation patterns. These flows can pattern proteins coupled to the cortex leading to the emergence of active mechanochemical patterns. In this review, we discuss physical approaches to understand force-generation and the concomitant patterns observed in the actomyosin cortex. We briefly outline the hydrodynamic theory of active gels as applicable to the cortex and discuss its consequences. We speculate on the role of the actomyosin cortex in sculpting large-scale tissues and end with an outlook for open problems.
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References
Howard J (2001) Mechanics of motor proteins and the cytoskeleton. Sinauer Associates, Sunderland, Massachusetts
Boal D (2012) Mechanics of the cell, 2nd edn. Cambridge University Press, Cambridge
Kolomeisky AB (2015) Motor proteins and molecular motors. CRC Press, Boca Raton
Shaevitz J, Nørrelykke S (2010) The cytoskeleton: I-beams of the cell. Phys Today 63:60
Fletcher DA, Mullins RD (2010) Cell mechanics and the cytoskeleton. Nature 463:485
Kraning-Rush CM, Carey SP, Califano JP, Smith BN, Reinhart-King C (2011) The role of the cytoskeleton in cellular force generation in 2D and 3D environments. Phys Biol 8:015009
Huber F, Schnauß J, Rönicke S, Rauch P, Müller K, Fütterer C, Käs J (2013) Emergent complexity of the cytoskeleton: from single filaments to tissue. Adv Phys 62:1
Harris AR, Jreij P, Fletcher DA (2018) Mechanotransduction by the actin cytoskeleton: converting mechanical stimuli into biochemical signals. Annu Rev Biophys 47:617
Huber F, Boire A, López MP, Koenderink GH (2015) Cytoskeletal crosstalk: when three different personalities team up. Curr Opin Cell Biol Cell Archit 32:39
Herrmann H, Bär H, Kreplak L, Strelkov SV, Aebi U (2007) Intermediate filaments: from cell architecture to nanomechanics. Nat Rev Mol Cell Biol 8:562
Block J, Schroeder V, Pawelzyk P, Willenbacher N, Köster S (2015) Physical properties of cytoplasmic intermediate filaments. Biochimica et Biophysica Acta (BBA) Mol Cell Res Mechanobiol 1853:3053
Wickstead B, Gull K (2011) The evolution of the cytoskeleton. J Cell Biol 194:513
Pollard TD, Goldman RD (2018) Overview of the cytoskeleton from an evolutionary perspective. Cold Spring Harbor Perspect Biol 10:a030288
Mast SO (1926) Structure, movement, locomotion, and stimulation in amoeba. J Morphol 41:347
White JG, Borisy GG (1983) On the mechanisms of cytokinesis in animal cells. J Theor Biol 101:289
Bray D, Heath J, Moss D (1986) The membrane-associated ‘Cortex’ of animal cells: its structure and mechanical properties. J Cell Sci 1986:71
Bray D, White J (1988) Cortical flow in animal cells. Science 239:883
Levayer R, Lecuit T (2012) Biomechanical regulation of contractility: spatial control and dynamics. Trends Cell Biol 22:61
Salbreux G, Charras G, Paluch E (2012) Actin cortex mechanics and cellular morphogenesis. Trends Cell Biol 22:536
Clark AG, Wartlick O, Salbreux G, Paluch EK (2014) Stresses at the cell surface during animal cell morphogenesis. Curr Biol 24:R484
Chalut KJ, Paluch EK (2016) The actin cortex: a bridge between cell shape and function. Dev Cell 38:571
Koenderink GH, Paluch EK (2018) Architecture shapes contractility in actomyosin networks. Curr Opin Cell Biol 50:79
Chugh P, Paluch EK (2018) The actin cortex at a glance. J Cell Sci 131:jcs186254
Agarwal P, Zaidel-Bar R (2019) Principles of actomyosin regulation in vivo. Trends Cell Biol 29:150
Illukkumbura R, Bland T, Goehring NW (2020) Patterning and polarization of cells by intracellular flows. Curr Opin Cell Biol 62:123
Kelkar M, Bohec P, Charras G (2020) Mechanics of the cellular actin cortex: from signalling to shape change. Curr Opin Cell Biol 66:69
Svitkina TM, Verkhovsky AB, McQuade KM, Borisy GG (1997) Analysis of the actin-myosin II system in fish epidermal keratocytes: mechanism of cell body translocation. J Cell Biol 139:397
Verkhovsky AB, Svitkina TM, Borisy GG (1999) Self-polarization and directional motility of cytoplasm. Curr Biol 9:11
Eghiaian F, Rigato A, Scheuring S (2015) Structural, mechanical, and dynamical variability of the actin cortex in living cells. Biophys J 108:1330
Clark AG, Dierkes K, Paluch EK (2013) Monitoring actin cortex thickness in live cells. Biophys J 105:570
Fritzsche M, Erlenkämper C, Moeendarbary E, Charras G, Kruse K (2016) Actin kinetics shapes cortical network structure and mechanics. Sci Adv 2:e1501337
Saha A, Nishikawa M, Behrndt M, Heisenberg C-P, Jülicher F, Grill SW (2016) Determining physical properties of the cell cortex. Biophys J 110:1421
Jülicher F, Ajdari A, Prost J (1997) Modeling molecular motors. Rev Mod Phys 69:1269
Sweeney HL, Hammers DW (2018) Muscle contraction. Cold Spring Harb Perspect Biol 10:a023200
Murrell MP, Gardel ML (2012) F-actin buckling coordinates contractility and severing in a biomimetic actomyosin cortex. Proc Natl Acad Sci 109:20820
Lenz M, Gardel ML, Dinner AR (2012a) Requirements for contractility in disordered cytoskeletal bundles. New J Phys 14:033037
Lenz M, Thoresen T, Gardel ML, Dinner AR (2012b) Contractile units in disordered actomyosin bundles arise from F-actin buckling. Phys Rev Lett 108:238107
Lenz M (2014) Geometrical origins of contractility in disordered actomyosin networks. Phys Rev X 4:041002
Murrell M, Oakes PW, Lenz M, Gardel ML (2015) Forcing cells into shape: the mechanics of actomyosin contractility. Nat Rev Mol Cell Biol 16:486
Chen S, Markovich T, MacKintosh FC (2020) Motor-free contractility in active gels. Phys Rev Lett 125:208101
Chugh P, Clark AG, Smith MB, Cassani DAD, Dierkes K, Ragab A, Roux PP, Charras G, Salbreux G, Paluch EK (2017) Actin cortex architecture regulates cell surface tension. Nat Cell Biol 19:689
Ennomani H, Letort G, Guérin C, Martiel J-L, Cao W, Nédélec F, De La Cruz EM, Théry M, Blanchoin L (2016) Architecture and connectivity govern actin network contractility. Curr Biol 26:616
Dasanayake NL, Michalski PJ, Carlsson AE (2011) General mechanism of actomyosin contractility. Phys Rev Lett 107:118101
Wang S, Wolynes PG (2012) Active contractility in actomyosin networks. Proc Natl Acad Sci 109:6446
Ditlev JA, Mayer BJ, Loew LM (2013) There is more than one way to model an elephant. Experiment-driven modeling of the actin cytoskeleton. Biophys J 104:520
Hawkins RJ, Liverpool TB (2014) Stress reorganization and response in active solids. Phys Rev Lett 113:028102
Hiraiwa T, Salbreux G (2016) Role of turnover in active stress generation in a filament network. Phys Rev Lett 116:188101
Ronceray P, Broedersz CP, Lenz M (2016) Fiber networks amplify active stress. Proc Natl Acad Sci 113:2827
Liman J, Bueno C, Eliaz Y, Schafer NP, Waxham MN, Wolynes PG, Levine H, Cheung MS (2020) The role of the Arp2/3 complex in shaping the dynamics and structures of branched actomyosin networks. Proc Natl Acad Sci 117:10825
Chaikin PM, Lubensky TC (1995) Principles of condensed matter physics. Cambridge University Press, Cambridge
Fang X, Kruse K, Lu T, Wang J (2019) Nonequilibrium physics in biology. Rev Mod Phys 91:045004
de Groot SR, Mazur P (1962) Non-equilibrium thermodynamics. Wiley, New York
Ramaswamy S (2017) Active matter. J Stat Mech Theory Exp 2017:054002
Kruse K, Joanny JF, Jülicher F, Prost J, Sekimoto K (2005) Generic theory of active polar gels: a paradigm for cytoskeletal dynamics. Eur Phys J E 16:5
Jülicher F, Kruse K, Prost J, Joanny JF (2007) Active behavior of the cytoskeleton. Phys Rep 449:3
Joanny J-F, Prost J (2009) Active gels as a description of the actin-myosin cytoskeleton. HFSP J 3:94
Ramaswamy S (2010) The mechanics and statistics of active matter. Annu Rev Cond Matter Phys 1:323
Marchetti MC, Joanny JF, Ramaswamy S, Liverpool TB, Prost J, Rao M, Simha RA (2013) Hydrodynamics of soft active matter. Rev Mod Phys 85:1143
Prost J, Jülicher F, Joanny J-F (2015) Active gel physics. Nat Phys 11:111
Jülicher F, Grill SW, Salbreux G (2018) Hydrodynamic theory of active matter. Rep Prog Phys 81:076601
Doostmohammadi A, Ignés-Mullol J, Yeomans JM, Sagués F (2018) Active nematics. Nat Commun 9:3246
Ramaswamy S (2019) Active fluids. Nat Rev Phys 1:640
de Gennes PG, Prost J (1993) The physics of liquid crystals, 2nd edn. Clarendon Press, Oxford
Fürthauer S, Strempel M, Grill SW, Jülicher F (2012) Active chiral fluids. Eur Phys J E 35:1
Fürthauer S, Strempel M, Grill SW, Jülicher F (2013) Active chiral processes in thin films. Phys Rev Lett 110:048103
Markovich T, Tjhung E, Cates ME (2019) Chiral active matter: microscopic ‘torque dipoles’ have more than one hydrodynamic description. New J Phys 21:112001
Naganathan SR, Fürthauer S, Nishikawa M, Jülicher F, Grill SW (2014) Active torque generation by the actomyosin cell cortex drives left-right symmetry breaking. eLife 3:e04165
Tee YH, Shemesh T, Thiagarajan V, Hariadi RF, Anderson KL, Page C, Volkmann N, Hanein D, Sivaramakrishnan S, Kozlov MM, Bershadsky AD (2015) Cellular chirality arising from the self-organization of the actin cytoskeleton. Nat Cell Biol 17:445
Pimpale LG, Middelkoop TC, Mietke A, Grill SW (2020) Cell lineage-dependent chiral actomyosin flows drive cellular rearrangements in early Caenorhabditis elegans development. eLife 9:e54930
Naganathan SR, Middelkoop TC, Fürthauer S, Grill SW (2016) Actomyosin-driven left-right asymmetry: from molecular torques to chiral self organization. Curr Opin Cell Biol 38:24
Inaki M, Liu J, Matsuno K (2016) Cell chirality: its origin and roles in left- right asymmetric development. Philos Trans R Soc B 371:20150403
Banerjee DS, Munjal A, Lecuit T, Rao M (2017) Actomyosin pulsation and flows in an active elastomer with turnover and network remodeling. Nat Commun 8:1121
Garzon-Coral C, Fantana HA, Howard J (2016) A force-generating machinery maintains the spindle at the cell center during mitosis. Science 352:1124
Mayer M, Depken M, Bois JS, Jülicher F, Grill SW (2010) Anisotropies in cortical tension reveal the physical basis of polarizing cortical flows. Nature 467:617
Nishikawa M, Naganathan SR, Jülicher F, Grill SW (2017) Controlling contractile instabilities in the actomyosin cortex. eLife 6:e19595
Doubrovinski K, Swan M, Polyakov O, Wieschaus EF (2017) Measurement of cortical elasticity in Drosophila melanogaster embryos using ferrofluids. Proc Natl Acad Sci 114:1051
Peukes J, Betz T (2014) Direct measurement of the cortical tension during the growth of membrane blebs. Biophys J 107:1810
Fischer-Friedrich E, Toyoda Y, Cattin CJ, Müller DJ, Hyman AA, Jülicher F (2016) Rheology of the active cell cortex in mitosis. Biophys J 111:589
eSilva MS, Stuhrmann B, Betz T, Koenderink GH (2014) Time-resolved microrheology of actively remodeling actomyosin networks. New J Phys 16:075010
Bois JS, Jülicher F, Grill S (2011) Pattern formation in active fluids. Phys Rev Lett 106:1
Kumar KV, Bois JS, Jülicher F, Grill SW (2014) Pulsatory patterns in active fluids. Phys Rev Lett 112:208101
Howard J, Grill SW, Bois JS (2011) Turing’s next steps: the mechanochemical basis of morphogenesis. Nat Rev Mol Cell Biol 12:392
Husain K, Rao M (2017) Emergent structures in an active polar fluid: dynamics of shape, scattering, and merger. Phys Rev Lett 118:078104
Das A, Polley A, Rao M (2016) Phase segregation of passive advective particles in an active medium. Phys Rev Lett 116:068306
Goswami D, Gowrishankar K, Bilgrami S, Ghosh S, Raghupathy R, Chadda R, Vishwakarma R, Rao M, Mayor S (2008) Nanoclusters of GPI-anchored proteins are formed by cortical actin-driven activity. Cell 135:1085
Chaudhuri A, Bhattacharya B, Gowrishankar K, Mayor S, Rao M (2011) Spatiotemporal regulation of chemical reactions by active cytoskeletal remodeling. Proc Natl Acad Sci 108:14825
Gowrishankar K, Ghosh S, Saha S, Rumamol C, Mayor S, Rao M, (2012) Active remodeling of cortical actin regulates spatiotemporal organization of cell surface molecules. Cell 149:1353
Salbreux G, Prost J, Joanny JF (2009) Hydrodynamics of cellular cortical flows and the formation of contractile rings. Phys Rev Lett 103:058102
Reymann A-C, Staniscia F, Erzberger A, Salbreux G, Grill SW (2016) Cortical flow aligns actin filaments to form a furrow. eLife 5:e17807
Mietke A, Jemseena V, Kumar KV, Sbalzarini IF, Jülicher F (2019a) Minimal model of cellular symmetry breaking. Phys Rev Lett 123:188101
Menon VV, Soumya SS, Agarwal A, Naganathan SR, Inamdar MM, Sain A (2017) Asymmetric flows in the intercellular membrane during cytokinesis. Biophys J 113:2787
Grill SW, Howard J, Schäffer E, Stelzer EHK, Hyman AA (2003) The distribution of active force generators controls mitotic spindle position. Science 301:518
Grill S, Kruse K, Jülicher F (2005) Theory of mitotic spindle oscillations. Phys Rev Lett 94:108104
Pollard TD (2017) Nine unanswered questions about cytokinesis. J Cell Biol 216:3007
Pollard TD, O’Shaughnessy B (2019) Molecular mechanism of cytokinesis. Annu Rev Biochem 88:661
Pollard TD (2014) The value of mechanistic biophysical information for systems-level understanding of complex biological processes such as cytokinesis. Biophys J 107:2499
Hird SN, White JG (1993) Cortical and cytoplasmic flow polarity in early embryonic cells of Caenorhabditis elegans. J Cell Biol 121:1343
Munro E, Nance J, Priess JR (2004) Cortical flows powered by asymmetrical contraction transport PAR proteins to establish and maintain anterior-posterior polarity in the early C. elegans embryo. Dev Cell 7:413
Goehring NW, Trong PK, Bois JS, Chowdhury D, Nicola EM, Hyman A, Grill SW (2011) Polarization of PAR proteins by advective triggering of a pattern-forming system. Science 334:1137
Gross P, Kumar KV, Goehring NW, Bois JS, Hoege C, Jülicher F, Grill SW (2019) Guiding self-organized pattern formation in cell polarity establishment. Nat Phys 15:293
Niwayama R, Shinohara K, Kimura A (2011) Hydrodynamic property of the cytoplasm is sufficient to mediate cytoplasmic streaming in the Caenorhabditis elegans embryo. Proc Natl Acad Sci 108:11900
Mogilner A, Manhart A (2018) Intracellular fluid mechanics: coupling cytoplasmic flow with active cytoskeletal gel. Annu Rev Fluid Mech 50:347
Mittasch M, Gross P, Nestler M, Fritsch AW, Iserman C, Kar M, Munder M, Voigt A, Alberti S, Grill SW, Kreysing M (2018) Non-invasive perturbations of intracellular flow reveal physical principles of cell organization. Nat Cell Biol 20:344
Kruse K, Joanny JF, Jülicher F, Prost J (2006) Contractility and retrograde flow in lamellipodium motion. Phys Biol 3:130
Hawkins RJ, Poincloux R, Bénichou O, Piel M, Chavrier P, Voituriez R (2011) Spontaneous contractility-mediated cortical flow generates cell migration in three-dimensional environments. Biophys J 101:1041
Álvarez-González B, Meili R, Bastounis E, Firtel RA, Lasheras JC, del Álamo JC (2015) Three-dimensional balance of cortical tension and axial contractility enables fast amoeboid migration. Biophys J 108:821
Callan-Jones AC, Voituriez R (2013) Active gel model of amoeboid cell motility. New J Phys 15:025022
Charras G, Paluch E (2008) Blebs lead the way: how to migrate without lamellipodia. Nat Rev Mol Cell Biol 9:730
Paluch EK, Raz E (2013) The role and regulation of blebs in cell migration. Curr Opin Cell Biol 25:582
Lim FY, Chiam K-H, Mahadevan L (2012) The size, shape, and dynamics of cellular blebs. Europhys Lett 100:28004
Alert R, Casademunt J (2016) Bleb nucleation through membrane peeling. Phys Rev Lett 116:068101
Manakova K, Yan H, Lowengrub J, Allard J (2016) Cell surface mechanochemistry and the determinants of bleb formation, healing, and travel velocity. Biophys J 110:1636
Mizuno D, Tardin C, Schmidt CF, MacKintosh FC (2007) Nonequilibrium mechanics of active cytoskeletal networks. Science 315:370
Soares e Silva M, Depken M, Stuhrmann B, Korsten M, MacKintosh FC, Koenderink GH, (2011) Active multistage coarsening of actin networks driven by myosin motors. Proc Natl Acad Sci 108:9408
Shah EA, Keren K (2014) Symmetry breaking in reconstituted actin cortices. eLife 3:e01433
Malik-Garbi M, Ierushalmi N, Jansen S, Abu-Shah E, Goode BL, Mogilner A, Keren K (2019) Scaling behaviour in steady-state contracting actomyosin networks. Nat Phys 15:509
Ierushalmi N, Malik-Garbi M, Manhart A, Shah EA, Goode BL, Mogilner A, Keren K (2020) Centering and symmetry breaking in confined contracting actomyosin networks. eLife 9:e55368
Tan TH, Malik-Garbi M, Abu-Shah E, Li J, Sharma A, MacKintosh FC, Keren K, Schmidt CF, Fakhri N (2018) Self-organized stress patterns drive state transitions in actin cortices. Sci Adv 4:eaar2847
Schaller V, Weber C, Semmrich C, Frey E, Bausch AR (2010) Polar patterns of driven filaments. Nature 467:73
Huber L, Suzuki R, Krüger T, Frey E, Bausch AR (2018) Emergence of coexisting ordered states in active matter systems. Science 361:255
Pontani L-L, van der Gucht J, Salbreux G, Heuvingh J, Joanny J-F, Sykes C (2009) Reconstitution of an actin cortex inside a liposome. Biophys J 96:192
Köster DV, Husain K, Iljazi E, Bhat A, Bieling P, Mullins RD, Rao M, Mayor S (2016) Actomyosin dynamics drive local membrane component organization in an in vitro active composite layer. Proc Natl Acad Sci 113:E1645
Thompson DW (1992) On growth and form: the complete, Revised edn. Dover Publications, New York
Lecuit T, Lenne P-F, Munro E (2011) Force generation, transmission, and integration during cell and tissue morphogenesis. Annu Rev Cell Dev Biol 27:157
Munjal A, Lecuit T (2014) Actomyosin networks and tissue morphogenesis. Development 141:1789
Gross P, Kumar KV, Grill SW (2017) How active mechanics and regulatory biochemistry combine to form patterns in development. Annu Rev Biophys 46:337
Ranft J, Basan M, Elgeti J, Joanny J-F, Prost J, Jülicher F (2010) Fluidization of tissues by cell division and apoptosis. Proc Natl Acad Sci USA 107:20863
Hannezo E, Prost J, Joanny J-F (2011) Instabilities of monolayered epithelia: shape and structure of villi and crypts. Phys Rev Lett 107:1
Hannezo E, Dong B, Recho P, Joanny J-F, Hayashi S (2015) Cortical instability drives periodic supracellular actin pattern formation in epithelial tubes. Proc Natl Acad Sci 112:8620
Maître J-L, Berthoumieux H, Krens SFG, Salbreux G, Jülicher F, Paluch E, Heisenberg C-P (2012) Adhesion functions in cell sorting by mechanically coupling the cortices of adhering cells. Science 338:253
Rauzi M, Lenne P-F, Lecuit T (2010) Planar polarized actomyosin contractile flows control epithelial junction remodelling. Nature 468:1110
Collinet C, Rauzi M, Lenne P-F, Lecuit T (2012) Local and tissue-scale forces drive oriented junction growth during tissue extension. Nat Cell Biol 17:1247
Munjal A, Philippe J-M, Munro E, Lecuit T (2015) A self-organized biomechanical network drives shape changes during tissue morphogenesis. Nature 524:351
Etournay R, Popović M, Merkel M, Nandi A, Blasse C, Aigouy B, Brandl H, Myers G, Salbreux G, Jülicher F, Eaton S (2015) Interplay of cell dynamics and epithelial tension during morphogenesis of the Drosophila pupal wing. eLife 4:e07090
Popović M, Nandi A, Merkel M, Etournay R, Eaton S, Jülicher F, Salbreux G (2017) Active dynamics of tissue shear flow. New J Phys 19:033006
Behrndt M, Salbreux G, Campinho P, Hauschild R, Oswald F, Roensch J, Grill SW, Heisenberg C-P (2012) Forces driving epithelial spreading in zebrafish gastrulation. Science 338:257
Begnaud S, Chen T, Delacour D, Mège R-M, Ladoux B (2016) Mechanics of epithelial tissues during gap closure. Curr Opin Cell Biol Cell Dyn 42:52
Srivastava P, Shlomovitz R, Gov NS, Rao M (2013) Patterning of polar active filaments on a tense cylindrical membrane. Phys Rev Lett 110:168104
Münster S, Jain A, Mietke A, Pavlopoulos A, Grill SW, Tomancak P (2019) Attachment of the blastoderm to the vitelline envelope affects gastrulation of insects. Nature 568:395
Jain A, Ulman V, Mukherjee A, Prakash M, Cuenca MB, Pimpale LG, Münster S, Haase R, Panfilio KA, Jug F, Grill SW, Tomancak P, Pavlopoulos A (2020) Regionalized tissue fluidization is required for epithelial gap closure during insect gastrulation. Nat Commun 11:5604
Brugués A, Anon E, Conte V, Veldhuis JH, Gupta M, Colombelli J, Muñoz JJ, Brodland GW, Ladoux B, Trepat X (2014) Forces driving epithelial wound healing. Nat Phys 10:683
Alert R, Trepat X (2020) Physical models of collective cell migration. Annu Rev Cond Matter Phys 11:77
Boocock D, Hino N, Ruzickova N, Hirashima T, Hannezo E (2020) Theory of mechanochemical patterning and optimal migration in cell monolayers. Nat Phys. https://doi.org/10.1038/s41567-020-01037-7
Pérez-González C, Alert R, Blanch-Mercader C, Gómez-González M, Kolodziej T, Bazellieres E, Casademunt J, Trepat X (2018) Active wetting of epithelial tissues. Nat Phys 15:79
Farhadifar R, Röper J-C, Aigouy B, Eaton S, Jülicher F (2007) The influence of cell mechanics, cell–cell interactions, and proliferation on epithelial packing. Curr Biol 17:2095
Fletcher AG, Osterfield M, Baker RE, Shvartsman SY (2014) Vertex models of epithelial morphogenesis. Biophys J 106:2291
Hannezo E, Prost J, Joanny J-F (2014) Theory of epithelial sheet morphology in three dimensions. Proc Natl Acad Sci 111:27
Monier B, Gettings M, Gay G, Mangeat T, Schott S, Guarner A, Suzanne M (2015) Apico-basal forces exerted by apoptotic cells drive epithelium folding. Nature 518:245
Alt S, Ganguly P, Salbreux G (2017) Vertex models: from cell mechanics to tissue morphogenesis. Philos Trans R Soc B 372:20150520
Turlier H, Audoly B, Prost J, Joanny J-F (2014) Furrow constriction in animal cell cytokinesis. Biophys J 106:114
Maitra A, Srivastava P, Rao M, Ramaswamy S (2014) Activating membranes. Phys Rev Lett 112:258101
Berthoumieux H, Maître J-L, Heisenberg C-P, Paluch EK, Jülicher F, Salbreux G (2014) Active elastic thin shell theory for cellular deformations. New J Phys 16:065005
Callan-Jones AC, Ruprecht V, Wieser S, Heisenberg CP, Voituriez R (2016) Cortical flow-driven shapes of nonadherent cells. Phys Rev Lett 116:028102
Salbreux G, Jülicher F (2017) Mechanics of active surfaces. Phys Rev E 96:032404
Rowghanian P, Campàs O (2017) Non-equilibrium membrane homeostasis in expanding cellular domains. Biophys J 113:132
Mietke A, Jülicher F, Sbalzarini IF (2019b) Self-organized shape dynamics of active surfaces. Proc Natl Acad Sci 116:29
Torres-Sánchez A, Millán D, Arroyo M (2019) Modelling fluid deformable surfaces with an emphasis on biological interfaces. J Fluid Mech 872:218
Morris RG, Rao M (2019) Active morphogenesis of epithelial monolayers. Phys Rev E 100:022413
Braun E, Keren K (2018) Hydra regeneration: closing the loop with mechanical processes in morphogenesis. BioEssays 0:1700204
Maroudas-Sacks Y, Garion L, Shani-Zerbib L, Livshits A, Braun E, Keren K (2020) Topological defects in the nematic order of actin fibres as organization centres of Hydra morphogenesis. Nat Phys. https://doi.org/10.1038/s41567-020-01083-1
Grill SW (2011) Growing up is stressful: biophysical laws of morphogenesis. Curr Opin Genet Dev 21:647
Naganathan SR, Fürthauer S, Rodriguez J, Fievet BT, Jülicher F, Ahringer J, Cannistraci CV, Grill SW (2018) Morphogenetic degeneracies in the actomyosin cortex. eLife 7:e37677
Acknowledgements
We acknowledge support from the Department of Atomic Energy, Government of India, under project number 12-R&D-TFR-5.10-1100, the Department of Biotechnology, Government of India for a Ramalingaswami re-entry fellowship, and the Max Planck Society via a Max-Planck-Partner-Group at ICTS-TIFR. We thank Aditya Singh Rajput and Siddharth Jha for discussions and comments.
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Kumar, K.V. The Actomyosin Cortex of Cells: A Thin Film of Active Matter. J Indian Inst Sci 101, 97–112 (2021). https://doi.org/10.1007/s41745-020-00220-2
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DOI: https://doi.org/10.1007/s41745-020-00220-2