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Physics of adherent cells
Rev. Mod. Phys. 85, 1327 – Published 27 August, 2013
DOI: https://doi.org/10.1103/RevModPhys.85.1327
Abstract
One of the most unique physical features of cell adhesion to external surfaces is the active generation of mechanical force at the cell-material interface. This includes pulling forces generated by contractile polymer bundles and networks, and pushing forces generated by the polymerization of polymer networks. These forces are transmitted to the substrate mainly by focal adhesions, which are large, yet highly dynamic adhesion clusters. Tissue cells use these forces to sense the physical properties of their environment and to communicate with each other. The effect of forces is intricately linked to the material properties of cells and their physical environment. Here a review is given of recent progress in our understanding of the role of forces in cell adhesion from the viewpoint of theoretical soft matter physics and in close relation to the relevant experiments.
Article Text
References (404)
- Aegerter-Wilmsen, Tinri, Alister C. Smith, Alix J. Christen, Christof M. Aegerter, Ernst Hafen, and Konrad Basler, 2010, “Exploring the effects of mechanical feedback on epithelial topology,” Development 137, 499.
- Albersdorfer, A., T. Feder, and E Sackmann, 1997, “Adhesion-induced domain formation by interplay of longrange repulsion and short-range attraction force: a model membrane study,” Biophys. J. 73, 245.
- Alberts, Bruce, Alexander Johnson, Peter Walter, Julian Lewis, Martin Raff, and Keith Roberts, 2007, Molecular Biology of the Cell (Taylor & Francis, London), 5th ed.
- Ali, O., H. Guillou, O. Destaing, C. Albigs-Rizo, M. R. Block, and B. Fourcade, 2011, “Cooperativity between integrin activation and mechanical stress leads to integrin clustering,” Biophys. J. 100, 2595.
- Aliee, Maryam, Jens-Christian Röper, Katharina P. Landsberg, Constanze Pentzold, Thomas J. Widmann, Frank Julicher, and Christian Dahmann, 2012, “Physical mechanisms shaping the drosophila dorsoventral compartment boundary,” Curr. Biol. 22, 967.
- Allioux-Guerin, Myriam, Delphine Icard-Arcizet, Christiane Durieux, Sylvie Henon, Francois Gallet, Jean-Claude Mevel, Marie-Jo Masse, Marc Tramier, and Maite Coppey-Moisan, 2009, “Spatiotemporal analysis of cell response to a rigidity gradient: a quantitative study using multiple optical tweezers,” Biophys. J. 96, 238.
- Ambrosi, D., et al., 2011, “Perspectives on biological growth and remodeling,” J. Mech. Phys. Solids 59, 863.
- Arfken, G., and H. Weber, 1995, Mathematical Methods for Physicists (Academic Press, New York).
- Baker, Brendon M., and Christopher S. Chen, 2012, “Deconstructing the third dimension: how 3D culture micro environments alter cellular cues,” J. Cell Sci. 125, 3015.
- Balaban, N. Q., et al., 2001, “Force and focal adhesion assembly: a close relationship studied using elastic micro-patterned substrates,” Nat. Cell Biol. 3, 466.
- Banerjee, S., and L. Giomi, 2013, “Polymorphism and bistability in adherent cells,” Soft Matter 9, 5251.
- Banerjee, S., and M. C. Marchetti, 2011, “Substrate rigidity deforms and polarizes active gels,” Europhys. Lett. 96, 28 003.
- Banerjee, Shiladitya, and M. Cristina Marchetti, 2012, “Contractile stresses in cohesive cell layers on finite-thickness substrates,” Phys. Rev. Lett. 109, 108101.
- Barnhart, Erin L., Kun-Chun Lee, Kinneret Keren, Alex Mogilner, and Julie A. Theriot, 2011, “An adhesion-dependent switch between mechanisms that determine motile cell shape,” PLoS Biol. 9, e1001059.
- Bar-Ziv, R., T. Tlusty, E. Moses, S. A. Safran, and A. Bershadsky, 1999, “Pearling in cells: A clue to understanding cell shape,” Proc. Natl. Acad. Sci. U.S.A. 96, 10 140.
- Basan, M., T. Risler, J. F. Joanny, X. Sastre-Garau, and J. Prost, 2009, “Homeostatic competition drives tumor growth and metastasis nucleation,” HFSP J. 3, 265.
- Bausch, A. R., and K. Kroy, 2006, “A bottom-up approach to cell mechanics,” Nat. Phys. 2, 231.
- Beleggia, M., M. De Graef, and Y. Millev, 2006, “Demagnetization factors of the general ellipsoid: An alternative to the maxwell approach,” Philos. Mag. 86, 2451.
- Bell, G. I., 1978, “Models for specific adhesion of cells to cells,” Science 200, 618.
- Ben-Isaac, E., Y. Park, G. Popescu, F. Brown, N. S. Gov, and Y. Shokef, 2011, “Effective temperature of red-blood-cell membrane fluctuations,” Phys. Rev. Lett. 106, 283103.
- Ben-Zvi, D., B. Shilo, A. Fainsod, and N. Barkai, 2008, “Scaling of the bmp activation gradient in xenopus embryos,” Nature (London) 453, 1205.
- Bershadsky, Alexander, Michael Kozlov, and Benjamin Geiger, 2006, “Adhesion-mediated mechanosensitivity: a time to experiment, and a time to theorize,” Curr. Opin. Cell Biol. 18, 472.
- Bershadsky, Alexander D., Nathalie Q. Balaban, and Benjamin Geiger, 2003, “Adhesion-Dependent cell mechanosensitivity,” Annu. Rev. Cell Dev. Biol. 19, 677.
- Besser, A., and S. A. Safran, 2006, “Force-induced adsorption and anisotropic growth of focal adhesions,” Biophys. J. 90, 3469.
- Besser, A., and U. S. Schwarz, 2007, “Coupling biochemistry and mechanics in cell adhesion: a model for inhomogeneous stress fiber contraction,” New J. Phys. 9, 425.
- Besser, A., and U. S. Schwarz, 2010, “Hysteresis in the cell response to time-dependent substrate stiffness,” Biophys. J. 99, L10.
- Besser, Achim, Julien Colombelli, Ernst H. K. Stelzer, and Ulrich S. Schwarz, 2011, “Viscoelastic response of contractile filament bundles,” Phys. Rev. E 83, 051902.
- Bischofs, I. B., F. Klein, D. Lehnert, M. Bastmeyer, and U. S. Schwarz, 2008, “Filamentous network mechanics and active contractility determine cell and tissue shape,” Biophys. J. 95, 3488.
- Bischofs, I. B., S. A. Safran, and U. S. Schwarz, 2004, “Elastic interactions of active cells with soft materials,” Phys. Rev. E 69, 021911.
- Bischofs, I. B., S. S. Schmidt, and U. S. Schwarz, 2009, “Effect of adhesion geometry and rigidity on cellular force distributions,” Phys. Rev. Lett. 103, 048101.
- Bischofs, I. B., and U. S. Schwarz, 2003, “Cell organization in soft media due to active mechanosensing,” Proc. Natl. Acad. Sci. U.S.A. 100, 9274.
- Bischofs, I. B., and U. S. Schwarz, 2005, “Effect of Poisson ratio on cellular structure formation,” Phys. Rev. Lett. 95, 068102.
- Bischofs, I. B., and U. S. Schwarz, 2006, “Collective effects in cellular structure formation mediated by compliant environments: a Monte Carlo study,” Acta Biomaterialia 2, 253.
- Boal, David, 2012, Mechanics of the Cell (Cambridge University Press, Cambridge, England), 2nd ed.
- Brangwynne, Clifford P., Frederick C. MacKintosh, Sanjay Kumar, Nicholas A. Geisse, Jennifer Talbot, L. Mahadevan, Kevin K. Parker, Donald E. Ingber, and David A. Weitz, 2006, “Microtubules can bear enhanced compressive loads in living cells because of lateral reinforcement,” J. Cell Biol. 173, 733.
- Bray, D, 2001, Cell Movements: from Molecules to Motility (Garland, New York), 2nd ed.
- Brown, R. A., R. Prajapati, D. A. McGrouther, I. V. Yannas, and M. Eastwood, 1998, “Tensional homeostasis in dermal fibroblasts: Mechanical responses to mechanical loading in three-dimensional substrates,” J. Cell. Physiol. 175, 323.
- Buehler, Markus J., and Sinan Keten, 2010, “Colloquium: Failure of molecules, bones, and the earth itself,” Rev. Mod. Phys. 82, 1459.
- Burridge, Keith, and Erika S. Wittchen, 2013, “The tension mounts: stress fibers as force-generating mechanotransducers,” J. Cell Biol. 200, 9.
- Bursac, P., G. Lenormand, B. Fabry, M. Oliver, D. A. Weitz, V. Viasnoff, J. P. Butler, and J. Fredberg, 2005, “Cytoskeletal remodelling and slow dynamics in the living cell,” Nat. Mater. 4, 557.
- Butler, James P., Iva Marija Toli-Nrrelykke, Ben Fabry, and Jeffrey J. Fredberg, 2002, “Traction fields, moments, and strain energy that cells exert on their surroundings,” Am. J. Physiol., Cell Physiol. 282, C595.
- Buxboim, A., I. L. Ivanovska, and D. E. Discher, 2010, “Matrix elasticity, cytoskeletal forces and physics of the nucleus: how deeply do cells ’feel’ outside and in?,” J. Cell Sci. 123, 297.
- Califano, Joseph P., and Cynthia A. Reinhart-King, 2010, “Substrate stiffness and cell area predict cellular traction stresses in single cells and cells in contact,” Cell. Mol. Bioeng. 3, 68.
- Cameron, A., J. Frith, and J. Cooper-White, 2011, “The influence of substrate creep on mesenchymal stem cell behaviour and phenotype,” Biomaterials 32, 5979.
- Camps, Otger, L. Mahadevan, and Jean-Franois Joanny, 2012, “Actin network growth under load,” Biophys. J. 102, 1049.
- Canela-Xandri, Oriol, Francesc Sagus, Jaume Casademunt, and Javier Buceta, 2011, “Dynamics and mechanical stability of the developing dorsoventral organizer of the wing imaginal disc,” PLoS Comput. Biol. 7, e1002153.
- Canham, P. B., 1970, “The minimum energy of bending as a possible explanation of the biconcave shape of the human red blood cell,” J. Theor. Biol. 26, 61.
- Carlsson, A. E., 2003, “Growth velocities of branched actin networks,” Biophys. J. 84, 2907.
- Chaikin, P. M., and T. C. Lubensky, 2000, Principles of Condensed Matter Physics (Cambridge University Press, Cambridge, England).
- Chan, C. E., and D. J. Odde, 2008, “Traction dynamics of filopodia on compliant substrates,” Science 322, 1687.
- Chang, C. W., and S. Kumar, 2013, “Vinculin tension distributions of individual stress fibers within cell–matrix adhesions,” J. Cell Sci. 126, 3021.
- Chen, C., R. Krishnan, E. Zhou, A. Ramachandran, D. Tambe, K. Rajendran, R. M. Adam, L. Deng, and J. J. Fredberg, 2010, “Fluidization and resolidification of the human bladder smooth muscle cell in response to transient stretch,” PLoS One 5, e12035.
- Chen, C. S., 2008, “Mechanotransduction—a field pulling together?,” J. Cell Sci. 121, 3285.
- Chen, Christopher S., Milan Mrksich, Sui Huang, George M. Whitesides, and Donald E. Ingber, 1997, “Geometric control of cell life and death,” Science 276, 1425.
- Chen, Daniel T. N., Qi Wen, Paul A. Janmey, John C. Crocker, and Arjun G. Yodh, 2010, “Rheology of soft materials,” Annu. Rev. Condens. Matter Phys. 1, 301.
- Choquet, D., D. P. Felsenfeld, and M. P. Sheetz, 1997, “Extracellular matrix rigidity causes strengthening of integrin-cytoskeleton linkages,” Cell 88, 39.
- Chrzanowska-Wodnicka, M., and K. Burridge, 1996, “Rho-stimulated contractility drives the formation of stress fibers and focal adhesions,” J. Cell Biol. 133, 1403.
- Civelekoglu-Scholey, G., A. Wayne Orr, I. Novak, J.-J. Meister, M. A. Schwartz, and A. Mogilner, 2005, “Model of coupled transient changes of rac, rho, adhesions and stress fibers alignment in endothelial cells responding to shear stress,” J. Theor. Biol. 232, 569.
- Cohen, Miriam, Derk Joester, Benjamin Geiger, and Lia Addadi, 2004, “Spatial and temporal sequence of events in cell adhesion: From molecular recognition to focal adhesion assembly,” Chembiochem 5, 1393.
- Collinsworth, A. M., C. E. Torgan, S. N. Nagda, R. J. Rajalingam, W. E. Kraus, and G. A. Truskey, 2000, “Orientation and length of mammalian skeletal myocytes in response to a unidirectional stretch,” Cell and Tissue Research 302, 243.
- Colombelli, J., A. Besser, H. Kress, E. G. Reynaud, P. Girard, E. Caussinus, U. Haselmann, J. V. Small, U. S. Schwarz, and E. H. K. Stelzer, 2009, “Mechanosensing in actin stress fibers revealed by a close correlation between force and protein localization,” J. Cell Sci. 122, 1665.
- Coughlin, Mark F., and Dimitrije Stamenovic, 2003, “A prestressed cable network model of the adherent cell cytoskeleton,” Biophys. J. 84, 1328.
- Crow, Ailey, Kevin D. Webster, Evan Hohlfeld, WinPin Ng, Phillip Geissler, and Daniel. A. Fletcher, 2012, “Contractile equilibration of single cells to step changes in extracellular stiffness,” Biophys. J. 102, 443.
- Cukierman, Edna, Roumen Pankov, Daron R. Stevens, and Kenneth M. Yamada, 2001, “Taking cell-matrix adhesions to the third dimension,” Science 294, 1708.
- Cuvelier, Damien, Manuel Thery, Yeh-Shiu Chu, Sylvie Dufour, Jean-Paul Thiery, Michel Bornens, Pierre Nassoy, and L. Mahadevan, 2007, “The universal dynamics of cell spreading,” Curr. Biol. 17, 694.
- Dasanayake, Nilushi L., Paul J. Michalski, and Anders E. Carlsson, 2011, “General mechanism of actomyosin contractility,” Phys. Rev. Lett. 107, 118101.
- Davies, P. F., A. Robotewskyj, and M. L. Griem, 1994, “Quantitative studies of endothelial cell adhesion. directional remodeling of focal adhesion sites in response to flow forces,” J. Clinical Investigation 93, 2031.
- De, R., and S. A. Safran, 2008, “Dynamical theory of active cellular response to external stress,” Phys. Rev. E 78, 031923.
- De, Rumi, Assaf Zemel, and Sam A. Safran, 2010, “Theoretical concepts and models of cellular mechanosensing,” Methods Cell Biol. 98C, 143.
- De, Rumi, Assaf Zemel, and Samuel A. Safran, 2007, “Dynamics of cell orientation,” Nat. Phys. 3, 655.
- De, Rumi, Assaf Zemel, and Samuel A. Safran, 2008, “Do cells sense stress or strain? measurement of cellular orientation can provide a clue,” Biophys. J. 94, L29.
- de Beer, Alex, Günter Majer, Sylvie Roke, and Joachim Spatz, 2010, “Continuous photobleaching to study the growth modes of focal adhesions,” J. Adhes. Sci. Technol. 24, 2323.
- de Gennes, P. G., 1979, Scaling Concepts in Polymer Physics (Cornell University Press, Ithaca).
- de Gennes, P. G., 1985, “Wetting: statics and dynamics,” Rev. Mod. Phys. 57, 827.
- de Gennes, P. G., 1992, “Soft matter (nobel lecture),” Angew. Chem., Int. Ed. Engl. 31, 842.
- de Gennes, P. G., and J. Prost, 1995, Physics of Liquid Crystals (Oxford University Press, New York), 2nd ed.
- Deibler, Martin, Joachim P. Spatz, and Ralf Kemkemer, 2011, “Actin fusion proteins alter the dynamics of mechanically induced cytoskeleton rearrangement,” PLoS One 6, e22941.
- Delano-Ayari, H., R. Al Kurdi, M. Vallade, D. Gulino-Debrac, and D. Riveline, 2004, “Membrane and Acto-Myosin tension promote clustering of adhesion proteins,” Proc. Natl. Acad. Sci. U.S.A. 101, 2229.
- Delano-Ayari, H., J. P. Rieu, and M. Sano, 2010, “4D traction force microscopy reveals asymmetric cortical forces in migrating dictyostelium cells,” Phys. Rev. Lett. 105, 248103.
- Dembo, Micah, and Yu-Li Wang, 1999, “Stresses at the Cell-to-Substrate interface during locomotion of fibroblasts,” Biophys. J. 76, 2307.
- Deng, Linhong, Xavier Trepat, James P. Butler, Emil Millet, Kathleen G. Morgan, David A. Weitz, and Jeffrey J. Fredberg, 2006, “Fast and slow dynamics of the cytoskeleton,” Nat. Mater. 5, 636.
- Deshpande, Vikram S, Robert M. McMeeking, and Anthony G. Evans, 2006, “A bio-chemo-mechanical model for cell contractility,” Proc. Natl. Acad. Sci. U.S.A. 103, 14 015.
- Deshpande, Vikram S., Robert M. McMeeking, and Anthony G. Evans, 2007, “A model for the contractility of the cytoskeleton including the effects of Stress-Fibre formation and dissociation,” Proc. R. Soc. A 463, 787.
- Diamant, H., and D. Andelman, 1996, “Kinetics of surfactant adsorption,” J. Phys. Chem. 100, 13 732.
- Dill, Ken A., and Sarina Bromberg, 2010, Molecular Driving Forces: Statistical Thermodynamics in Chemistry and Biology (Garland Publishing, New York).
- Discher, D. E., D. J. Mooney, and P. W. Zandstra, 2009, “Growth factors, matrices, and forces combine and control stem cells,” Science 324, 1673.
- Discher, Dennis E., Paul Janmey, and Yu-li Wang, 2005, “Tissue cells feel and respond to the stiffness of their substrate,” Science 310, 1139.
- Dogterom, Marileen, and Bernard Yurke, 1997, “Measurement of the force-velocity relation for growing microtubules,” Science 278, 856.
- Doi, M., 1996, Introduction to Polymer Physics (Oxford University, New York).
- DuFort, Christopher C., Matthew J. Paszek, and Valerie M. Weaver, 2011, “Balancing forces: architectural control of mechanotransduction,” Nat. Rev. Mol. Cell Biol. 12, 308.
- Duke, T. A. J., 1999, “Molecular model of muscle contraction,” Proc. Natl. Acad. Sci. U.S.A. 96, 2770.
- Eastwood, M., V. C. Mudera, D. A. McGrouther, and R. A. Brown, 1998, “Effect of precise mechanical loading on fibroblast populated collagen lattices: Morphological changes,” Cell Motil. Cytoskeleton 40, 13.
- Edwards, Carina M., and Ulrich S. Schwarz, 2011, “Force localization in contracting cell layers,” Phys. Rev. Lett. 107, 128101.
- Engler, A., S. Sen, H. Sweeney, and D. Discher, 2006, “Matrix elasticity directs stem cell lineage specification,” Cell 126, 677.
- Engler, A. J., C. Carag-Krieger, C. P. Johnson, M. Raab, H.-Y. Speicher, Tang D. W., J. W. Sanger, J. M. Sanger, and D. E. Discher, 2008, “Embryonic cardiomyocytes beat best on a matrix with heart-like elasticity: scar-like rigidity inhibits beating,” J. Cell Sci. 121, 3794.
- Engler, A. J., M. A. Griffin, S. Sen, C. G. Bonnetnann, H. L. Sweeney, and D. E. Discher, 2004, “Myotubes differentiate optimally on substrates with tissue-like stiffness: pathological implications for soft or stiff microenvironments,” J. Cell Biol. 166, 877.
- Engler, Adam, Lucie Bacakova, Cynthia Newman, Alina Hategan, Maureen Griffin, and Dennis Discher, 2004, “Substrate compliance versus ligand density in cell on gel responses,” Biophys. J. 86, 617.
- Erdmann, T., and U. S. Schwarz, 2004a, “Stability of adhesion clusters under constant force,” Phys. Rev. Lett. 92, 108102.
- Erdmann, T., and U. S. Schwarz, 2004b, “Stochastic dynamics of adhesion clusters under shared constant force and with rebinding,” J. Chem. Phys. 121, 8997.
- Erdmann, Thorsten, and Ulrich S. Schwarz, 2012, “Stochastic force generation by small ensembles of myosin ii motors,” Phys. Rev. Lett. 108, 188101.
- Eshelby, J. D., 1957, “The determination of the elastic field of an ellipsoidal inclusion, and related problems,” Proc. R. Soc. A 241, 376.
- Eshelby, J. D., 1959, “The field outside of an ellipsoidal inclusion,” Proc. R. Soc. A 252, 561.
- Evans, E., and K. Ritchie, 1997, “Dynamic strength of molecular adhesion bonds,” Biophys. J. 72, 1541.
- Evans, Evan A., and David A. Calderwood, 2007, “Forces and bond dynamics in cell adhesion,” Science 316, 1148.
- Farhadifar, R., J. C. Roper, B. Algouy, S. Eaton, and F. Julicher, 2007, “The influence of cell mechanics, cell-cell interactions, and proliferation on epithelial packing,” Curr. Biol. 17, 2095.
- Faust, Uta, Nico Hampe, Wolfgang Rubner, Norbert Kirchgessner, Sam Safran, Bernd Hoffmann, and Rudolf Merkel, 2011, “Cyclic stress at mHz frequencies aligns fibroblasts in direction of zero strain,” PLoS One 6, e28963.
- Fernandez, P., and A. R. Bausch, 2009, “The compaction of gels by cells: a case of collective mechanical activity,” Integr. Biol. 1, 252.
- Fery, Andreas, and Richard Weinkamer, 2007, “Mechanical properties of micro- and nanocapsules: Single-capsule measurements,” Polymer 48, 7221.
- Filippov, A. E., J. Klafter, and M. Urbakh, 2004, “Friction through dynamical formation and rupture of molecular bonds,” Phys. Rev. Lett. 92, 135503.
- Fink, Jenny, et al., 2011, “External forces control mitotic spindle positioning,” Nat. Cell Biol. 13, 771.
- Fletcher, D. A., and P. L. Geissler, 2009, “Active biological materials,” Annu. Rev. Phys. Chem. 60, 469.
- Fletcher, Daniel A., and R. Dyche Mullins, 2010, “Cell mechanics and the cytoskeleton,” Nature (London) 463, 485.
- Fletcher, Daniel A., and Julie A. Theriot, 2004, “An introduction to cell motility for the physical scientist,” Phys. Biol. 1, T1.
- Friedrich, B. M., E. Fischer-Friedrich, N. S. Gov, and S. A. Safran, 2012, “Sarcomeric pattern formation by actin cluster coalescence,” PLoS Comput. Biol. 8, e1002544.
- Friedrich, B. M., and S. A. Safran, 2011, “Nematic order by elastic interactions and cellular rigidity sensing,” Europhys. Lett. 93, 28007.
- Friedrich, Benjamin M., Amnon Buxboim, Dennis E. Discher, and Samuel A. Safran, 2011, “Striated Acto-Myosin fibers can reorganize and register in response to elastic interactions with the matrix,” Biophys. J. 100, 2706.
- Friedrich, Benjamin M., and Samuel A. Safran, 2012, “How cells feel their substrate: spontaneous symmetry breaking of active surface stresses,” Soft Matter 8, 3223.
- Fu, Jianping, Yang-Kao Wang, Michael T. Yang, Ravi A. Desai, Xiang Yu, Zhijun Liu, and Christopher S. Chen, 2010, “Mechanical regulation of cell function with geometrically modulated elastomeric substrates,” Nat. Methods 7, 733.
- Fung, Y. C., 1993, Biomechanics: Mechanical Properties of Living Tissues (Springer-Verlag, New York).
- Gao, Huajian, Jin Qian, and Bin Chen, 2011, “Probing mechanical principles of focal contacts in cellmatrix adhesion with a coupled stochasticelastic modelling framework,” J. R. Soc. Interface 8, 1217.
- Gardel, M. L., J. H. Shin, F. C. MacKintosh, L. Mahadevan, P. Matsudaira, and D. A. Weitz, 2004, “Elastic behavior of cross-linked and bundled actin networks,” Science 304, 1301.
- Gardel, Margaret L., Karen E. Kasza, Clifford P. Brangwynne, Jiayu Liu, and David A. Weitz, 2008, “Mechanical response of cytoskeletal networks,” Methods Cell Biol. 89, 487.
- Gardel, Margaret L., Benedikt Sabass, Lin Ji, Gaudenz Danuser, Ulrich S. Schwarz, and Clare M. Waterman, 2008, “Traction stress in focal adhesions correlates biphasically with actin retrograde flow speed,” J. Cell Biol. 183, 999.
- Gauthier, Nils C., Marc Antoine Fardin, Pere Roca-Cusachs, and Michael P. Sheetz, 2011, “Temporary increase in plasma membrane tension coordinates the activation of exocytosis and contraction during cell spreading,” Proc. Natl. Acad. Sci. U.S.A. 108, 14 467.
- Gavara, Nuria, Pere Roca-Cusachs, Raimon Sunyer, Ramon Farre, and Daniel Navajas, 2008, “Mapping cell-matrix stresses during stretch reveals inelastic reorganization of the cytoskeleton,” Biophys. J. 95, 464.
- Geiger, Benjamin, Joachim P. Spatz, and Alexander D. Bershadsky, 2009, “Environmental sensing through focal adhesions,” Nat. Rev. Mol. Cell Biol. 10, 21.
- Gonzalez-Rodriguez, David, Karine Guevorkian, Stphane Douezan, and Franoise Brochard-Wyart, 2012, “Soft matter models of developing tissues and tumors,” Science 338, 910.
- Gov, N., 2004, “Membrane undulations driven by force fluctuations of active proteins,” Phys. Rev. Lett. 93, 268104.
- Gov, Nir S., 2006, “Dynamics and morphology of microvilli driven by actin polymerization,” Phys. Rev. Lett. 97, 018101.
- Guo, M., and D. Weitz, 2012 (unpublished).
- Gurin, Thomas, Jacques Prost, Pascal Martin, and Jean-Franois Joanny, 2010, “Coordination and collective properties of molecular motors: theory,” Curr. Opin. Cell Biol. 22, 14.
- Guthardt Torres, P., I. B. Bischofs, and U. S. Schwarz, 2012, “Contractile network models for adherent cells,” Phys. Rev. E 85, 011913.
- Haken, H., 1983, Synergetics, an Introduction: Nonequilibrium Phase Transitions and Self-Organization in Physics, Chemistry, and Biology (Springer-Verlag, Berlin).
- Hanggi, Peter, Peter Talkner, and Michal Borkovec, 1990, “Reaction-rate theory: fifty years after Kramers,” Rev. Mod. Phys. 62, 251.
- Hayakawa, K., N. Sato, and T. Obinata, 2001, “Dynamic reorientation of cultured cells and stress fibers under mechanical stress from periodic stretching,” Exp. Cell Res. 268, 104.
- Head, D. A., A. J. Levine, and F. C. MacKintosh, 2003a, “Distinct regimes of elastic response and deformation modes of cross-linked cytoskeletal and semiflexible polymer networks,” Phys. Rev. E 68, 061907.
- Head, David A., Alex J. Levine, and F. C. MacKintosh, 2003b, “Deformation of cross-linked semiflexible polymer networks,” Phys. Rev. Lett. 91, 108102.
- Heil, P., and J. P. Spatz, 2010, “Lateral shear forces applied to cells with single elastic micropillars to influence focal adhesion dynamics,” J. Phys. Condens. Matter 22, 194108.
- Helfrich, W., 1973, “Elastic properties of lipid bilayers: theory and possible experiments,” Z. Naturforsch. C 28, 693.
- Heussinger, Claus, and Erwin Frey, 2006, “Stiff polymers, foams, and fiber networks,” Phys. Rev. Lett. 96, 017802.
- Heussinger, Claus, Boris Schaefer, and Erwin Frey, 2007, “Nonaffine rubber elasticity for stiff polymer networks,” Phys. Rev. E 76, 031906.
- Hoffman, Brenton D., Carsten Grashoff, and Martin A. Schwartz, 2011, “Dynamic molecular processes mediate cellular mechanotransduction,” Nature (London) 475, 316.
- Hoffmann, Max, and Ulrich S. Schwarz, 2013, “A kinetic model for RNA-interference of focal adhesions,” BMC Systems Biology 7, 2.
- Hotulainen, Pirta, and Pekka Lappalainen, 2006, “Stress fibers are generated by two distinct actin assembly mechanisms in motile cells,” J. Cell Biol. 173, 383.
- Howard, J., 2001, Mechanics of Motor Proteins and the Cytoskeleton (Sinauer Associates, Sunderland, MA).
- Hsu, Hui-Ju, Chin-Fu Lee, and Roland Kaunas, 2009, “A dynamic stochastic model of frequency-dependent stress fiber alignment induced by cyclic stretch,” PLoS One 4, e4853.
- Huang, S., and D. E. Ingber, 1999, “The structural and mechanical complexity of cell-growth control,” Nat. Cell Biol. 1, E131.
- Huber, F., J. Schnauss, S. Ronicke, P. Rauch, K. Muller, C. Futterer, and J. Kas, 2013, “Emergent complexity of the cytoskeleton: from single filaments to tissue,” Adv. Phys. 62, 1.
- Hufnagel, L., A. A. Teleman, H. Rouault, S. M. Cohen, and B. I. Shraiman, 2007, “On the mechanism of wing size determination in fly development,” Proc. Natl. Acad. Sci. U.S.A. 104, 3835.
- Hur, Sung Sik, Yihua Zhao, Yi-Shuan Li, Elliot Botvinick, and Shu Chien, 2009, “Live cells exert 3-Dimensional traction forces on their substrata,” Cell. Mol. Bioeng. 2, 425.
- Huxley, A. F., 1957, “Muscle structure and theories of contraction,” Prog. Biophys. Biophys. Chem. 7, 255 [http://www.ncbi.nlm.nih.gov/pubmed/13485191].
- Ingber, D. E., 1993, “Cellular tensegrity—defining new rules of biological design that govern the cytoskeleton,” J. Cell Sci. 104, 613.
- Ingber, D. E., 2003, “Mechanobiology and diseases of mechanotransduction,” Ann. Med. 35, 564.
- Isenberg, B., P. DiMilla, M. Walker, S. Kim, and J. Wong, 2009, “Vascular smooth muscle cell durotaxis depends on substrate stiffness gradient strength,” Biophys. J. 97, 1313.
- Israelachvili, J. N., 2011, Intermolecular and Surface Forces (Academic Press, Waltham), 3rd ed.
- Iyer, K. Venkatesan, Shovamayee Maharana, Soumya Gupta, Albert Libchaber, Tsvi Tlusty, and G. V. Shivashankar, 2012, “Modeling and experimental methods to probe the link between global transcription and spatial organization of chromosomes,” PLoS One 7, e46628.
- Jacot, Jeffrey G., Andrew D. McCulloch, and Jeffrey H. Omens, 2008, “Substrate stiffness affects the functional maturation of neonatal rat ventricular myocytes,” Biophys. J. 95, 3479.
- Janmey, P. A., S. Hvidt, J. Lamb, and T. P. Stossel, 1990, “Resemblance of actin-binding protein actin gels to covalently cross-linked networks,” Nature (London) 345, 89.
- Janmey, P. A., J. P. Winer, M. E. Murray, and Q. Wen, 2009, “The hard life of soft cells,” Cell Motil. Cytoskeleton 66, 597.
- Janmey, Paul A., and R. Tyler Miller, 2011, “Mechanisms of mechanical signaling in development and disease,” J. Cell Sci. 124, 9.
- Johnson, K. L., 1985, Contact Mechanics (Cambridge University Press, Cambridge, England).
- Johnson, K. L., K. Kendall, and A. D. Roberts, 1971, “Surface energy and the contact of elastic solids,” Proc. R. Soc. A 324, 301.
- Jones, Richard A. L., 2002, Soft Condensed Matter (Oxford University Press, New York).
- Julicher, F., K. Kruse, J. Prost, and J. F. Joanny, 2007, “Active behavior of the cytoskeleton,” Phys. Rep. 449, 3.
- Julicher, Frank, and Jacques Prost, 1995, “Cooperative molecular motors,” Phys. Rev. Lett. 75, 2618.
- Jungbauer, S., H. J. Gao, J. P. Spatz, and R. Kemkemer, 2008, “Two characteristic regimes in frequency-dependent dynamic reorientation of fibroblasts on cyclically stretched substrates,” Biophys. J. 95, 3470.
- Kaunas, R., H. J. Hsu, and S. Deguchi, 2011, “Sarcomeric model of stretch-induced stress fiber reorganization,” Cell Health and Cytoskeleton 3, 13.
- Kaunas, R., P. Nguyen, S. Usami, and S. Chien, 2005, “Cooperative effects of rho and mechanical stretch on stress fiber organization,” Proc. Natl. Acad. Sci. U.S.A. 102, 15 895.
- Kawakami, K., H. Tatsumi, and M. Sokabe, 2001, “Dynamics of integrin clustering at focal contacts of endothelial cells studied by multimode imaging microscopy,” J. Cell Sci. 114, 3125.
- Kilian, K. A., B. Bugarija, B. T. Lahn, and M. Mrksich, 2010, “Geometric cues for directing the differentiation of mesenchymal stem cells,” Proc. Natl. Acad. Sci. U.S.A. 107, 4872.
- Klein, Franziska, Benjamin Richter, Thomas Striebel, Clemens M. Franz, Georg von Freymann, Martin Wegener, and Martin Bastmeyer, 2011, “Two-component polymer scaffolds for controlled three-dimensional cell culture,” Adv. Mater. 23, 1341.
- Klein, Franziska, Thomas Striebel, Joachim Fischer, Zhongxiang Jiang, Clemens M. Franz, Georg von Freymann, Martin Wegener, and Martin Bastmeyer, 2010, “Elastic fully three-dimensional microstructure scaffolds for cell force measurements,” Adv. Mater. 22, 868.
- Klotzsch, Enrico, Michael L. Smith, Kristopher E. Kubow, Simon Muntwyler, William C. Little, Felix Beyeler, Delphine Gourdon, Bradley J. Nelson, and Viola Vogel, 2009, “Fibronectin forms the most extensible biological fibers displaying switchable force-exposed cryptic binding sites,” Proc. Natl. Acad. Sci. U.S.A. 106, 18 267.
- Klumpp, Stefan, and Reinhard Lipowsky, 2005, “Cooperative cargo transport by several molecular motors,” Proc. Natl. Acad. Sci. U.S.A. 102, 17 284.
- Knoche, Sebastian, and Jan Kierfeld, 2011, “Buckling of spherical capsules,” Phys. Rev. E 84, 046608.
- Koenderink, Gijsje H., Zvonimir Dogic, Fumihiko Nakamura, Poul M. Bendix, Frederick C. MacKintosh, John H. Hartwig, Thomas P. Stossel, and David A. Weitz, 2009, “An active biopolymer network controlled by molecular motors,” Proc. Natl. Acad. Sci. U.S.A. 106, 15 192.
- Kohler, Simone, Volker Schaller, and Andreas R. Bausch, 2011, “Structure formation in active networks,” Nat. Mater. 10, 462.
- Kollmannsberger, P., C. M. Bidan, J. W. C. Dunlop, and P. Fratzl, 2011, “The physics of tissue patterning and extracellular matrix organisation: how cells join forces,” Soft Matter 7, 9549.
- Kollmannsberger, P., and B. Fabry, 2011, “Linear and nonlinear rheology of living cells,” Annu. Rev. Mater. Res. 41, 75.
- Komura, S., K. Tamura, and T. Kato, 2005, “Buckling of spherical shells adhering onto a rigid substrate,” Eur. Phys. J. E 18, 343.
- Korff, T., and H. G. Augustin, 1999, “Tensional forces in fibrillar extracellular matrices control directional capillary sprouting,” J. Cell Sci. 112, 3249.
- Kovar, D. R., 2006, “Molecular details of formin-mediated actin assembly,” Curr. Opin. Cell Biol. 18, 11.
- Krishnan, R., et al., 2009, “Reinforcement versus fluidization in cytoskeletal mechanoresponsiveness,” PLoS One 4, e5486.
- Kruse, K., J. F. Joanny, F. Julicher, J. Prost, and K. Sekimoto, 2004, “Asters, vortices, and rotating spirals in active gels of polar filaments,” Phys. Rev. Lett. 92, 078101.
- Kruse, K., and F. Julicher, 2000a, “Actively contracting bundles of polar filaments,” Phys. Rev. Lett. 85, 1778.
- Kruse, K., and F. Julicher, 2000b, “Actively contracting bundles of polar filaments,” Phys. Rev. Lett. 85, 1778.
- Kruse, K., and F. Julicher, 2003, “Self-organization and mechanical properties of active filament bundles,” Phys. Rev. E 67, 051913.
- Kruse, K., A. Zumdieck, and F. Julicher, 2003, “Continuum theory of contractile fibres,” Europhys. Lett. 64, 716.
- Kumar, Sanjay, Iva Z. Maxwell, Alexander Heisterkamp, Thomas R. Polte, Tanmay P. Lele, Matthew Salanga, Eric Mazur, and Donald E. Ingber, 2006, “Viscoelastic retraction of single living stress fibers and its impact on cell shape, cytoskeletal organization, and extracellular matrix mechanics,” Biophys. J. 90, 3762.
- Kuo, Jean-Cheng, Xuemei Han, Cheng-Te Hsiao, John R. Yates III, and Clare M. Waterman, 2011, “Analysis of the myosin-II-responsive focal adhesion proteome reveals a role for [beta]-Pix in negative regulation of focal adhesion maturation,” Nat. Cell Biol. 13, 383.
- Kurpinski, Kyle, Julia Chu, Craig Hashi, and Song Li, 2006, “Anisotropic mechanosensing by mesenchymal stem cells,” Proc. Natl. Acad. Sci. U.S.A. 103, 16 095.
- Ladoux, Benoit, and Alice Nicolas, 2012, “Physically based principles of cell adhesion mechanosensitivity in tissues,” Rep. Prog. Phys. 75, 116601.
- Landau, L. D., and E. M. Lifshitz, 1970, Theory of Elasticity, Course of Theoretical Physics (Pergamon Press, Oxford), Vol. 7, 2nd ed.
- Landsberg, Katharina P., Reza Farhadifar, Jonas Ranft, Daiki Umetsu, Thomas J. Widmann, Thomas Bittig, Amani Said, Frank Jlicher, and Christian Dahmann, 2009, “Increased cell bond tension governs cell sorting at the drosophila anteroposterior compartment boundary,” Curr. Biol. 19, 1950.
- Lau, K. H., and W. Kohn, 1977, “Elastic interaction of two atoms adsorbed on a solid-surface,” Surf. Sci. 65, 607.
- Lauffenburger, Douglas A., and Alan F. Horwitz, 1996, “Cell migration: A physically integrated molecular process,” Cell 84, 359.
- Lecuit, Thomas, and Pierre-Francois Lenne, 2007, “Cell surface mechanics and the control of cell shape, tissue patterns and morphogenesis,” Nat. Rev. Mol. Cell Biol. 8, 633.
- Lee, Kun-Chun, and Andrea J. Liu, 2009, “Force-Velocity relation for Actin-Polymerization-Driven motility from brownian dynamics simulations,” Biophys. J. 97, 1295.
- Legant, Wesley R., Colin K. Choi, Jordan S. Miller, Lin Shao, Liang Gao, Eric Betzig, and Christopher S. Chen, 2013, “Multidimensional traction force microscopy reveals out-of-plane rotational moments about focal adhesions,” Proc. Natl. Acad. Sci. U.S.A. 110, 881.
- Lele, T. P., J. Pendse, S. Kumar, M. Salanga, J. Karavitis, and D. E. Ingber, 2006, “Mechanical forces alter zyxin unbinding kinetics within focal adhesions of living cells,” J. Cell. Physiol. 207, 187.
- Lenne, Pierre-Francois, and Alice Nicolas, 2009, “Physics puzzles on membrane domains posed by cell biology,” Soft Matter 5, 2841.
- Lenz, M., T. Thoresen, M. L. Gardel, and A. R. Dinner, 2012, “Contractile units in disordered actomyosin bundles arise from f-actin buckling,” Phys. Rev. Lett. 108, 238107.
- Li, Ying, Prabhakar Bhimalapuram, and Aaron R. Dinner, 2010, “Model for how retrograde actin flow regulates adhesion traction stresses,” J. Phys. Condens. Matter 22, 194113.
- Lieleg, O., M. M. Claessens, Y. Luan, and A. R. Bausch, 2008, “Transient binding and dissipation in cross-linked actin networks,” Phys. Rev. Lett. 101, 108101.
- Lim H. W., Gerald, Michael Wortis, and Ranjan Mukhopadhyay, 2002, “Stomatocyte-discocyte-echinocyte sequence of the human red blood cell: Evidence for the bilayer couple hypothesis from membrane mechanics,” Proc. Natl. Acad. Sci. U.S.A. 99, 16766.
- Lipowsky, Reinhard, 1996, “Adhesion of membranes via anchored stickers,” Phys. Rev. Lett. 77, 1652.
- Liverpool, T. B., and M. C. Marchetti, 2003, “Instabilities of isotropic solutions of active polar filaments,” Phys. Rev. Lett. 90, 138102.
- Lo, C. M., H. B. Wang, M. Dembo, and Y. L. Wang, 2000, “Cell movement is guided by the rigidity of the substrate,” Biophys. J. 79, 144.
- Lobkovsky, Alex, Sharon Gentges, Hao Li, David Morse, and T. A. Witten, 1995, “Scaling properties of stretching ridges in a crumpled elastic sheet,” Science 270, 1482.
- Loosli, Y., R. Luginbuehl, and J. G. Snedeker, 2010, “Cytoskeleton reorganization of spreading cells on Micro-Patterned islands: A functional model,” Phil. Trans. R. Soc. A 368, 2629.
- Lu, L., Y. Feng, W. J. Hucker, S. J. Oswald, G. D. Longmore, and F. C. Yin, 2008, “Actin stress fiber preextension in human aortic endothelial cells,” Cell Motil. Cytoskeleton 65, 281.
- Luo, Y. Z., X. Xu, T. Lele, S. Kumar, and D. E. Ingber, 2008, “A multi-modular tensegrity model of an actin stress fiber,” J. Biomech. 41, 2379.
- Macdonald, Alice, A. Rick Horwitz, and Douglas A. Lauffenburger, 2008, “Kinetic model for lamellipodal actin-integrin ’clutch’ dynamics,” Cell Adhesion & Migration 2, 95.
- Mackintosh, F., 2006, in Soft Condensed Matter Physics in Molecular and Cell Biology (CRC Press, Boca Raton, FL).
- MacKintosh, Frederick C., and Christoph F. Schmidt, 2010, “Active cellular materials,” Curr. Opin. Cell Biol. 22, 29.
- Maier, W., and A. Saupe, 1959, Z. Naturforschg. 14, 882.
- Majkut, Stephanie F., and Dennis E. Discher, 2012, “Cardiomyocytes from late embryos and neonates do optimal work and striate best on substrates with tissue-level elasticity: metrics and mathematics,” Biomech. Model. Mechanobiol. 11, 1219.
- Manneville, J. B., P. Bassereau, S. Ramaswamy, and J. Prost, 2001, “Active membrane fluctuations studied by micropipet aspiration,” Phys. Rev. E 64, 021908.
- Marchetti, M. C., J.-F. Joanny, S. Ramaswamy, T. B. Liverpool, J. Prost, Madan Rao, and R. Aditi Simha, 2013, “Soft active matter,” Rev. Mod. Phys. 85, 1143.
- Marcq, Philippe, Natsuhiko Yoshinaga, and Jacques Prost, 2011, “Rigidity sensing explained by active matter theory,” Biophys. J. 101, L33.
- Marcy, Yann, Jean-Franois Joanny, Jacques Prost, and Ccile Sykes, 2007, “Probing friction in actin-based motility,” New J. Phys. 9, 431.
- Marcy, Yann, Jacques Prost, Marie-France Carlier, and Cecile Sykes, 2004, “Forces generated during actin-based propulsion: A direct measurement by micromanipulation,” Proc. Natl. Acad. Sci. U.S.A. 101, 5992.
- Marko, J., and E. Siggia, 1995, “Stretching DNA,” Macromolecules 28, 8759.
- McGarry, J. P., J. Fu, M. T. Yang, C. S. Chen, R. M. McMeeking, A. G. Evans, and V. S. Deshpande, 2009, “Simulation of the contractile response of cells on an array of Micro-Posts,” Phil. Trans. R. Soc. A 367, 3477.
- McKenna, N. M., C. S. Johnson, and Y. L. Wang, 1986, “Formation and alignment of z lines in living chick myotubes microinjected with rhodamine-labeled alpha-actinin,” J. Cell Biol. 103, 2163.
- McMahon, T. A., 1984, Muscles, reflexes, and locomotion (Princeton University Press, Princeton, NJ).
- Menes, R., and S. A. Safran, 1997, “Nonlinear response of membranes to pinning sites,” Phys. Rev. E 56, 1891.
- Mertz, Aaron F., Shiladitya Banerjee, Yonglu Che, Guy K. German, Ye Xu, Callen Hyland, M. Cristina Marchetti, Valerie Horsley, and Eric R. Dufresne, 2012, “Scaling of traction forces with the size of cohesive cell colonies,” Phys. Rev. Lett. 108, 198101.
- Mertz, Aaron F., Yonglu Che, Shiladitya Banerjee, Jill M. Goldstein, Kathryn A. Rosowski, Stephen F. Revilla, Carien M. Niessen, M. Cristina Marchetti, Eric R. Dufresne, and Valerie Horsley, 2013, “Cadherin-based intercellular adhesions organize epithelial cellmatrix traction forces,” Proc. Natl. Acad. Sci. U.S.A. 110, 842.
- Miao, Ling, Udo Seifert, Michael Wortis, and Hans-Gunther Dobereiner, 1994, “Budding transitions of fluid-bilayer vesicles: The effect of area-difference elasticity,” Phys. Rev. E 49, 5389.
- Mitra, Satyajit K., Daniel A. Hanson, and David D. Schlaepfer, 2005, “Focal adhesion kinase: in command and control of cell motility,” Nat. Rev. Mol. Cell Biol. 6, 56.
- Mitragotri, Samir, and Joerg Lahann, 2009, “Physical approaches to biomaterial design,” Nat. Mater. 8, 15.
- Mitrossilis, Demosthene, Jonathan Fouchard, Axel Guiroy, Nicolas Desprat, Nicolas Rodriguez, Ben Fabry, and Atef Asnacios, 2009, “Single-cell response to stiffness exhibits muscle-like behavior,” Proc. Natl. Acad. Sci. U.S.A. 106, 18 243.
- Mitrossilis, Demosthene, Jonathan Fouchard, David Pereira, Francois Postic, Alain Richert, Michel Saint-Jean, and Atef Asnacios, 2010, “Real-time single-cell response to stiffness,” Proc. Natl. Acad. Sci. U.S.A. 107, 16 518.
- Mizuno, Daisuke, Catherine Tardin, C. F. Schmidt, and F. C. MacKintosh, 2007, “Nonequilibrium mechanics of active cytoskeletal networks,” Science 315, 370.
- Mogilner, A., and B. Rubinstein, 2005, “The physics of filopodial protrusion,” Biophys. J. 89, 782.
- Mogilner, Alex, 2006, “On the edge: modeling protrusion,” Curr. Opin. Cell Biol. 18, 32.
- Mogilner, Alex, and Kinneret Keren, 2009, “The shape of motile cells,” Curr. Biol. 19, R762.
- Muller, C. A., and Tilo Pompe, 2013, “Dissipative interactions in cell-matrix adhesion,” Soft Matter 9, 6207.
- Mura, T., 1991, Micromechanics of defects (Kluwer Academic, Dordrecht).
- Murrell, Michael P., and Margaret L. Gardel, 2012, “F-actin buckling coordinates contractility and severing in a biomimetic actomyosin cortex,” Proc. Natl. Acad. Sci. U.S.A. 109, 20 820.
- Nekouzadeh, Ali, Kenneth M. Pryse, Elliot L. Elson, and Guy M. Genin, 2008, “Stretch-activated force shedding, force recovery, and cytoskeletal remodeling in contractile fibroblasts,” J. Biomech. 41, 2964.
- Nelson, C. M., R. P. Jean, J. L. Tan, W. F. Liu, N. J. Sniadecki, A. A. Spector, and C. S. Chen, 2005, “Emergent patterns of growth controlled by multicellular form and mechanics,” Proc. Natl. Acad. Sci. U.S.A. 102, 11 594.
- Nicolas, A., B. Geiger, and S. A. Safran, 2004, “Cell mechanosensitivity controls the anisotropy of focal adhesions,” Proc. Natl. Acad. Sci. U.S.A. 101, 12 520.
- Nicolas, A., and S. A. Safran, 2004, “Elastic deformations of grafted layers with surface stress,” Phys. Rev. E 69, 051902.
- Nicolas, A., and S. A. Safran, 2006a, “Limitation of cell adhesion by the elasticity of the extracellular matrix,” Biophys. J. 91, 61.
- Nicolas, Alice, Achim Besser, and Samuel A. Safran, 2008, “Dynamics of cellular focal adhesions on deformable substrates: consequences for cell force microscopy,” Biophys. J. 95, 527.
- Noguchi, Hiroshi, and Gerhard Gompper, 2005, “Shape transitions of fluid vesicles and red blood cells in capillary flows,” Proc. Natl. Acad. Sci. U.S.A. 102, 14 159.
- Olberding, J. E., M. D. Thouless, E. M. Arruda, and K. Garikipati, 2010, “The non-equilibrium thermodynamics and kinetics of focal adhesion dynamics,” PLoS One 5, e12043.
- Paluch, Ewa, and Carl-Philipp Heisenberg, 2009, “Biology and physics of cell shape changes in development,” Curr. Biol. 19, R790.
- Parekh, Sapun H., Ovijit Chaudhuri, Julie A. Theriot, and Daniel A. Fletcher, 2005, “Loading history determines the velocity of actin-network growth,” Nat. Cell Biol. 7, 1219.
- Parker, Kevin Kit, Amy Lepre Brock, Cliff Brangwynne, Robert J. Mannix, Ning Wang, Emanuele Ostuni, Nicholas A. Geisse, Josehphine C. Adams, George M. Whitesides, and Donald E. Ingber, 2002, “Directional control of lamellipodia extension by constraining cell shape and orienting cell tractional forces,” FASEB J. 16, 1195.
- Pate, E., G. J. Wilson, M. Bhimani, and R. Cooke, 1994, “Temperature dependence of the inhibitory effects of orthovanadate on shortening velocity in fast skeletal muscle,” Biophys. J. 66, 1554.
- Pathak, A., V. S. Deshpande, R. M. McMeeking, and A. G. Evans, 2008, “The simulation of stress fibre and focal adhesion development in cells on patterned substrates,” J. R. Soc. Interface 5, 507.
- Patla, Israel, Tova Volberg, Nadav Elad, Vera Hirschfeld-Warneken, Carsten Grashoff, Reinhard Fassler, Joachim P. Spatz, Benjamin Geiger, and Ohad Medalia, 2010, “Dissecting the molecular architecture of integrin adhesion sites by cryo-electron tomography,” Nat. Cell Biol. 12, 909.
- Paul, R., P. Heil, J. P. Spatz, and U. S. Schwarz, 2008, “Propagation of mechanical stress through the actin cytoskeleton towards focal adhesions: model and experiment,” Biophys. J. 94, 1470.
- Pearson, H., 2008, “The cellular hullabaloo,” Nature (London) 453, 150.
- Pelham, R. J., and Y. L. Wang, 1997, “Cell locomotion and focal adhesions are regulated by substrate flexibility,” Proc. Natl. Acad. Sci. U.S.A. 94, 13 661.
- Pellegrin, Stphanie, and Harry Mellor, 2007, “Actin stress fibres,” J. Cell Sci. 120, 3491.
- Peterson, Lynda J., Zenon Rajfur, Amy S. Maddox, Christopher D. Freel, Yun Chen, Magnus Edlund, Carol Otey, and Keith Burridge, 2004, “Simultaneous stretching and contraction of stress fibers in vivo,” Mol. Biol. Cell 15, 3497.
- Phillips, Rob, Jane Kondev, and Julie Theriot, 2008, Physical Biology of the Cell (Taylor & Francis, London), 1st ed.
- Pieranski, P., 1983, “Colloidal crystals,” Contemp. Phys. 24, 25.
- Pinto, Ines Mendes, Boris Rubinstein, Andrei Kucharavy, Jay R. Unruh, and Rong Li, 2012, “Actin depolymerization drives actomyosin ring contraction during budding yeast cytokinesis,” Dev. Cell 22, 1247.
- Pirentis, A., and P. Lazopoulos, 2009, “On stress fiber reorientation under plane substrate stretching,” Arch. Appl. Mech. 79, 263.
- Pirentis, A., E. Peruski, A. Jordan, and D. Stamenovic, 2011, “A model for stress fiber realignment caused by cytoskeletal fluidization during cyclic stretching,” Cell. Mol. Bioeng. 4, 67.
- Placais, P.-Y., M. Balland, T. Gurin, J.-F. Joanny, and P. Martin, 2009, “Spontaneous oscillations of a minimal actomyosin system under elastic loading,” Phys. Rev. Lett. 103, 158102.
- Plotnikov, Sergey V., Ana M. Pasapera, Benedikt Sabass, and Clare M. Waterman, 2012, “Force fluctuations within focal adhesions mediate ECM-Rigidity sensing to guide directed cell migration,” Cell 151, 1513.
- Pollard, Thomas D., and Julien Berro, 2008, “Mathematical models and simulations of cellular processes based on actin filaments,” J. Biol. Chem. 284, 5433.
- Pollard, Thomas D., and Gary G. Borisy, 2003, “Cellular motility driven by assembly and disassembly of actin filaments,” Cell 112, 453.
- Pollard, Thomas D., and John A. Cooper, 2009, “Actin, a central player in cell shape and movement,” Science 326, 1208.
- Pompe, T., S. Glorius, T. Bischoff, I. Uhlmann, M. Kaufmann, S. Brenner, and C. Werner, 2009, “Dissecting the impact of matrix anchorage and elasticity in cell adhesion,” Biophys. J. 97, 2154.
- Ponti, A., M. Machacek, S. L. Gupton, C. M. Waterman-Storer, and G. Danuser, 2004, “Two distinct actin networks drive the protrusion of migrating cells,” Science 305, 1782.
- Powers, Thomas R., 2010, “Dynamics of filaments and membranes in a viscous fluid,” Rev. Mod. Phys. 82, 1607.
- Pozrikidis, C., and Costas Pozrikidis, 2003, Modeling and Simulation of Capsules and Biological Cells (CRC Press, Boca Raton, FL).
- Prager-Khoutorsky, Masha, Alexandra Lichtenstein, Ramaswamy Krishnan, Kavitha Rajendran, Avi Mayo, Zvi Kam, Benjamin Geiger, and Alexander D. Bershadsky, 2011, “Fibroblast polarization is a matrix-rigidity-dependent process controlled by focal adhesion mechanosensing,” Nat. Cell Biol. 13, 1457.
- Prass, Marcus, Ken Jacobson, Alex Mogilner, and Manfred Radmacher, 2006, “Direct measurement of the lamellipodial protrusive force in a migrating cell,” J. Cell Biol. 174, 767.
- Pullarkat, Pramod A., Paul Dommersnes, Pablo Fernndez, Jean-Franois Joanny, and Albrecht Ott, 2006, “Osmotically driven shape transformations in axons,” Phys. Rev. Lett. 96, 048104.
- Qian, J., et al., 2013, “A mechanochemical model of cell reorientation on substrates under cyclic stretch,” PLoS One 8, e65864.
- Qian, Jin, Jizeng Wang, and Huajian Gao, 2008, “Lifetime and strength of adhesive molecular bond clusters between elastic media,” Langmuir 24, 1262.
- Ramaswamy, Sriram, 2010, “The mechanics and statistics of active matter,” Annu. Rev. Condens. Matter Phys. 1, 323.
- Ranft, Jonas, Markus Basan, Jens Elgeti, Jean-Francois Joanny, Jacques Prost, and Frank Julicher, 2010, “Fluidization of tissues by cell division and apoptosis,” Proc. Natl. Acad. Sci. U.S.A. 107, 20 863.
- Rauzi, Matteo, Pascale Verant, Thomas Lecuit, and Pierre-Francois Lenne, 2008, “Nature and anisotropy of cortical forces orienting drosophila tissue morphogenesis,” Nat. Cell Biol. 10, 1401.
- Rehfeldt, F., A. Brown, M. Raab, S. Cai, A. L. Zajac, A. Zemel, and D. E. Discher, 2012, “Hyaluronic acid matrices show matrix stiffness in 2d and 3d dictates cytoskeletal order and myosin-ii phosphorylation within stem cells,” Integr. Biol. 4, 422.
- Reinhart-King, Cynthia A., Micah Dembo, and Daniel A. Hammer, 2008, “Cell-cell mechanical communication through compliant substrates,” Biophys. J. 95, 6044.
- Rhee, D., J. M. Sanger, and J. W. Sanger, 1994, “The premyofibril: evidence for its role in myofibrillogenesis,” Cell Motil. Cytoskeleton 28, 1.
- Ridley, Anne J., 2011, “Life at the leading edge,” Cell 145, 1012.
- Riehl, B. D., J-H. Park, I. K. Kwon, and J. Y. Lim, 2012, “Mechanical stretching for tissue engineering: two-dimensional and three-dimensional constructs,” Tissue Engineering Part B 18, 288.
- Riveline, Daniel, Eli Zamir, Nathalie Q. Balaban, Ulrich S. Schwarz, Toshimasa Ishizaki, Shuh Narumiya, Zvi Kam, Benjamin Geiger, and Alexander D. Bershadsky, 2001, “Focal contacts as mechanosensors: Externally applied local mechanical force induces growth of focal contacts by an Mdia1-Dependent and Rock-Independent mechanism,” J. Cell Biol. 153, 1175.
- Roopa, T., A. Mazumder, A. Basu, L. Mahadevan, and G. V. Shivashankar, 2008, “Dynamics of chromatin decondensation reveals the structural integrity of a mechanically prestressed cell nucleus,” Biophys. J. 95, 3028.
- Rubinstein, M., and R. Colby, 2003, Polymer Physics (Oxford University, New York).
- Rudnicki, M. S., et al., 2013, “Nonlinear strain stiffening is not suffcient to explain how far cells can feel on fibrous protein gels,” Biophys. J.. 105, 11.
- Russell, Robert J., Shen-Ling Xia, Richard B. Dickinson, and Tanmay P. Lele, 2009, “Sarcomere mechanics in capillary endothelial cells,” Biophys. J. 97, 1578.
- Sabass, B., M. L. Gardel, C. M. Waterman, and U. S. Schwarz, 2008, “High resolution traction force microscopy based on experimental and computational advances,” Biophys. J. 94, 207.
- Sabass, Benedikt, and Ulrich S. Schwarz, 2010, “Modeling cytoskeletal flow over adhesion sites: competition between stochastic bond dynamics and intracellular relaxation,” J. Phys. Condens. Matter 22, 194112.
- Saez, Alexandre, Axel Buguin, Pascal Silberzan, and Benoit Ladoux, 2005, “Is the mechanical activity of epithelial cells controlled by deformations or forces?,” Biophys. J. 89, L52.
- Saez, Alexandre, Marion Ghibaudo, Axel Buguin, Pascal Silberzan, and Benoit Ladoux, 2007, “Rigidity-driven growth and migration of epithelial cells on microstructured anisotropic substrates,” Proc. Natl. Acad. Sci. U.S.A. 104, 8281.
- Safran, S. A., 1999, “Curvature elasticity of thin films,” Adv. Phys. 48, 395.
- Safran, S. A., and R. De, 2009, “Nonlinear dynamics of cell orientation,” Phys. Rev. E 80, 060901.
- Safran, S. A., and D. R. Hamann, 1979, “Long-range elastic interactions and staging in graphite-intercalation compounds,” Phys. Rev. Lett. 42, 1410.
- Safran, Samuel, 2003, Statistical Thermodynamics of Surfaces, Interfaces, and Membranes (Westview Press, Boulder, CO).
- Saitô, Nobuhiko, Kunihiko Takahashi, and Yasuo Yunoki, 1967, “Statistical mechanical theory of stiff chains,” J. Phys. Soc. Jpn. 22, 219.
- Samuel, J. L., and H. H. Vandenburgh, 1990, “Mechanically induced orientation of adult-rat cardiac myocytes invitro,” In Vitro Cellular & Developmental Biology 26, 905.
- Savir, Yonatan, Elad Noor, Ron Milo, and Tsvi Tlusty, 2010, “Cross-species analysis traces adaptation of rubisco toward optimality in a low-dimensional landscape,” Proc. Natl. Acad. Sci. U.S.A. 107, 3475.
- Schallamach, A., 1963, “A theory of dynamic rubber friction,” Wear 6, 375.
- Schreiber, Christian H., Murray Stewart, and Thomas Duke, 2010, “Simulation of cell motility that reproduces the force-velocity relationship,” Proc. Natl. Acad. Sci. U.S.A. 107, 9141.
- Schwarz, U. S., 2007, “Soft matters in cell adhesion: rigidity sensing on soft elastic substrates,” Soft Matter 3, 263.
- Schwarz, U. S., N. Q. Balaban, D. Riveline, A. Bershadsky, B. Geiger, and S. A. Safran, 2002, “Calculation of forces at focal adhesions from elastic substrate data: the effect of localized force and the need for regularization,” Biophys. J. 83, 1380.
- Schwarz, U. S., and I. B. Bischofs, 2005, “Physical determinants of cell organization in soft media,” Medical Engineering and Physics 27, 763.
- Schwarz, U. S., T. Erdmann, and I. B. Bischofs, 2006, “Focal adhesions as mechanosensors: the two-spring model,” BioSystems 83, 225.
- Schwarz, U. S., and G. Gompper, 2002, in Morphology of Condensed Matter: Physics and Geometry of Spatially Complex Systems, Lecture Notes in Physics, edited by K. R. Mecke and D. Stoyan (Springer, Heidelberg), Vol. 600, pp. 107–151.
- Schwarz, U. S., S. Komura, and S. A. Safran, 2000, “Deformation and tribology of multi-walled hollow nanoparticles,” Europhys. Lett. 50, 762.
- Schwarz, U. S., and S. A. Safran, 2002, “Elastic interactions of cells,” Phys. Rev. Lett. 88, 048102.
- Schwarz, Ulrich S., and Margaret L. Gardel, 2012, “United we stand integrating the actin cytoskeleton and cellmatrix adhesions in cellular mechanotransduction,” J. Cell Sci. 125, 3051.
- Seifert, Udo, 1997, “Configurations of fluid membranes and vesicles,” Adv. Phys. 46, 13.
- Seifert, Udo, Karin Berndl, and Reinhard Lipowsky, 1991, “Shape transformations of vesicles: Phase diagram for spontaneous- curvature and bilayer-coupling models,” Phys. Rev. A 44, 1182.
- Seifert, Udo, and Reinhard Lipowsky, 1990, “Adhesion of vesicles,” Phys. Rev. A 42, 4768.
- Serena, E., S. Zatti, E. Reghelin, A. Pasut, E. Cimetta, and N. Elvasorre, 2010, “Soft substrates drive optimal differentiation of human healthy and dystrophic myotubes,” Integr. Biol. 2, 193.
- Shao, Danying, Wouter-Jan Rappel, and Herbert Levine, 2010, “Computational model for cell morphodynamics,” Phys. Rev. Lett. 105, 108104.
- Shemesh, T., B. Geiger, A. D. Bershadsky, and M. M. Kozlov, 2005, “Focal adhesions as mechanosensors: a physical mechanism,” Proc. Natl. Acad. Sci. U.S.A. 102, 12 383.
- Shemesh, Tom, D. Alexander Bershadsky, and Michael M. Kozlov, 2012, “Physical model for Self-Organization of actin cytoskeleton and adhesion complexes at the cell front,” Biophys. J. 102, 1746.
- Shemesh, Tom, Alexander B. Verkhovsky, Tatyana M. Svitkina, Alexander D. Bershadsky, and Michael M. Kozlov, 2009, “Role of focal adhesions and mechanical stresses in the formation and progression of the lamellum interface,” Biophys. J. 97, 1254.
- Shirinsky, V. P., A. S. Antonov, K. G. Birukov, A. A. Sobolevsky, Y. A. Romanov, N. V. Kabaeva, G. N. Antonova, and V. N. Smirnov, 1989, “Mechano-chemical control of human endothelium orientation and size,” J. Cell Biol. 109, 331.
- Shokef, Y., and S. A. Safran, 2012a, “Erratum: Scaling laws for the response of nonlinear elastic media with implications for cell mechanics,” Phys. Rev. Lett. 109, 169901.
- Shokef, Y., and S. A. Safran, 2012b, “Scaling laws for the response of nonlinear elastic media with implications for cell mechanics,” Phys. Rev. Lett. 108, 178103.
- Shraiman, B. I., 2005, “Mechanical feedback as a possible regulator of tissue growth,” Proc. Natl. Acad. Sci. U.S.A. 102, 3318.
- Shtengel, Gleb, et al., 2009, “Interferometric fluorescent super-resolution microscopy resolves 3D cellular ultrastructure,” Proc. Natl. Acad. Sci. U.S.A. 106, 3125.
- Siems, R., 1968, “Mechanical interactions of point defects,” Phys. Status Solidi (b) 30, 645.
- Smith, Ana, and Udo Seifert, 2005, “Effective adhesion strength of specifically bound vesicles,” Phys. Rev. E 71, 061902.
- Smith, Ana, Kheya Sengupta, Stefanie Goennenwein, Udo Seifert, and Erich Sackmann, 2008, “Force-Induced growth of adhesion domains is controlled by receptor mobility,” Proc. Natl. Acad. Sci. U.S.A. 105, 6906.
- Soares e Silva, Marina, Martin Depken, Bjrn Stuhrmann, Marijn Korsten, Fred C. MacKintosh, and Gijsje H. Koenderink, 2011, “Active multistage coarsening of actin networks driven by myosin motors,” Proc. Natl. Acad. Sci. U.S.A. 108, 9408.
- Solon, Jerome, Ilya Levental, Kheya Sengupta, Penelope C. Georges, and Paul A. Janmey, 2007, “Fibroblast adaptation and stiffness matching to soft elastic substrates,” Biophys. J. 93, 4453.
- Srinivasan, M., and S. Walcott, 2009, “Binding site models of friction due to the formation and rupture of bonds: State-function formalism, force-velocity relations, response to slip velocity transients, and slip stability,” Phys. Rev. E 80, 046124.
- Stachowiak, Matthew, R., Patrick M. McCall, Todd Thoresen, Hayri E. Balcioglu, Lisa Kasiewicz, Margaret L. Gardel, and Ben O’Shaughnessy, 2012, “Self-organization of myosin II in reconstituted actomyosin bundles,” Biophys. J. 103, 1265.
- Stachowiak, Matthew, R., and Ben O’Shaughnessy, 2008, “Kinetics of stress fibers,” New J. Phys. 10, 025002.
- Stachowiak, Matthew, R., and Ben O’Shaughnessy, 2009, “Recoil after severing reveals stress fiber contraction mechanisms,” Biophys. J. 97, 462.
- Stamenovic, D., S. M. Mijailovich, I. M. Tolic-Norrelykke, J. X. Chen, and N. Wang, 2002, “Cell prestress. II. contribution of microtubules,” Am. J. Physiol. Cell Physiol. 282, C617.
- Stamenovic, Dimitrije, and Donald E. Ingber, 2009, “Tensegrity-guided self assembly: from molecules to living cells,” Soft Matter 5, 1137.
- Stamenovic, Dimitrije, Noah Rosenblatt, Martin Montoya-Zavala, Benjamin D. Matthews, Shaohua Hu, Bela Suki, Ning Wang, and Donald E. Ingber, 2007, “Rheological behavior of living cells is timescale-dependent,” Biophys. J. 93, L39.
- Storm, C., J. J. Pastore, F. C. MacKintosh, T. C. Lubensky, and P. A. Janmey, 2005, “Nonlinear elasticity in biological gels,” Nature (London) 435, 191.
- Stricker, Jonathan, Yvonne Aratyn-Schaus, Patrick W. Oakes, and Margaret L. Gardel, 2011, “Spatiotemporal constraints on the Force-Dependent growth of focal adhesions,” Biophys. J. 100, 2883.
- Stricker, Jonathan, Tobias Falzone, and Margaret L. Gardel, 2010, “Mechanics of the f-actin cytoskeleton,” J. Biomech. 43, 9.
- Svitkina, Tatyana M., and Gary G. Borisy, 1999, “ complex and actin depolymerizing Factor/Cofilin in dendritic organization and treadmilling of actin filament array in lamellipodia,” J. Cell Biol. 145, 1009.
- Takakuda, K., and H. Miyairi, 1996, “Tensile behaviour of fibroblasts cultured in collagen gel,” Biomaterials 17, 1393.
- Tan, John L., Joe Tien, Dana M. Pirone, Darren S. Gray, Kiran Bhadriraju, and Christopher S. Chen, 2003, “Cells lying on a bed of microneedles: An approach to isolate mechanical force,” Proc. Natl. Acad. Sci. U.S.A. 100, 1484.
- Tang, Xin, Piyush Bajaj, Rashid Bashir, and Taher A. Saif, 2011, “How far cardiac cells can see each other mechanically,” Soft Matter 7, 6151.
- Tawada, Katsuhisa, and Ken Sekimoto, 1991, “Protein friction exerted by motor enzymes through a weak-binding interaction,” J. Theor. Biol. 150, 193.
- Tempel, M., G. Isenberg, and E. Sackmann, 1996, “Temperature-induced sol-gel transition and microgel formation in alpha-actinin cross-linked actin networks: a rheological study,” Phys. Rev. E 54, 1802.
- Thery, Manuel, Andrea Jimenez-Dalmaroni, Victor Racine, Michel Bornens, and Frank Julicher, 2007, “Experimental and theoretical study of mitotic spindle orientation,” Nature (London) 447, 493.
- Thery, Manuel, Anne Pepin, Emilie Dressaire, Yong Chen, and Michel Bornens, 2006, “Cell distribution of stress fibres in response to the geometry of the adhesive environment,” Cell Motil. Cytoskeleton 63, 341.
- Thompson, D’Arcy Wentworth, 1992, On Growth and Form (Cambridge University Press, Cambridge, England).
- Thoresen, Todd, Martin Lenz, and Margaret L. Gardel, 2011, “Reconstitution of contractile actomyosin bundles,” Biophys. J. 100, 2698.
- Thoresen, Todd, Martin Lenz, and Margaret L. Gardel, 2013, “Thick filament length and isoform composition determine self-organized contractile units in actomyosin bundles,” Biophys. J. 104, 655.
- Tondon, Abhishek, Hui-Ju Hsu, and Roland Kaunas, 2012, “Dependence of cyclic stretch-induced stress fiber reorientation on stretch waveform,” J. Biomech. 45, 728.
- Trappmann, Britta, et al., 2012, “Extracellular-matrix tethering regulates stem-cell fate,” Nat. Mater. 11, 642.
- Trepat, X., L. Deng, S. An, D. Navajas, D. Tschumperlin, W. Gerthoffer, J. Butler, and J. Fredberg, 2007, “Universal physical responses to stretch in the living cell,” Nature (London) 447, 592.
- Trepat, X., M. Grabulosa, F. Puig, G. N. Maksym, D. Navajas, and R. Farre, 2004, “Viscoelasticity of human alveolar epithelial cells subjected to stretch,” Am. J. Physiol. 287, L1025.
- Trepat, X., F. Puig, N. Gavara, J. J. Fredberg, R. Farre, and D. Navajas, 2006, “Effect of stretch on structural integrity and micromechanics of human alveolar epithelial cell monolayers exposed to thrombin,” Am. J. Physiol. 290, L1104.
- Trichet, Lea, Jimmy Le Digabel, Rhoda J. Hawkins, Sri Ram Krishna Vedula, Mukund Gupta, Claire Ribrault, Pascal Hersen, Raphael Voituriez, and Benoit Ladoux, 2012, “Evidence of a Large-Scale mechanosensing mechanism for cellular adaptation to substrate stiffness,” Proc. Natl. Acad. Sci. U.S.A. 109, 6933.
- Tseng, Qingzong, Eve Duchemin-Pelletier, Alexandre Deshiere, Martial Balland, Herv Guillou, Odile Filhol, and Manuel Thry, 2012, “Spatial organization of the extracellular matrix regulates cellcell junction positioning,” Proc. Natl. Acad. Sci. U.S.A. 109, 1506.
- Urban, Edit, Sonja Jacob, Maria Nemethova, Guenter P. Resch, and J. Victor Small, 2010, “Electron tomography reveals unbranched networks of actin filaments in lamellipodia,” Nat. Cell Biol. 12, 429.
- van Kampen, N. G., 1992, Stochastic Processes in Physics and Chemistry (Elsevier, Amsterdam).
- Veigel, Claudia, Justin E. Molloy, Stephan Schmitz, and John Kendrick-Jones, 2003, “Load-dependent kinetics of force production by smooth muscle myosin measured with optical tweezers,” Nat. Cell Biol. 5, 980.
- Verkhovsky, A. B., and G. G. Borisy, 1993, “Non-Sarcomeric mode of myosin II organization in the fibroblast lamellum,” J. Cell Biol. 123, 637.
- Vianay, B., J. Kafer, E. Planus, M. Block, F. Graner, and H. Guillou, 2010, “Single cells spreading on a protein lattice adopt an energy minimizing shape,” Phys. Rev. Lett. 105, 128101.
- Vilfan, Andrej, and Thomas Duke, 2003, “Instabilities in the transient response of muscle,” Biophys. J. 85, 818.
- Vliegenthart, G. A., and G. Gompper, 2006a, “Forced crumpling of self-avoiding elastic sheets,” Nat. Mater. 5, 216.
- Vliegenthart, Gerard Adriaan, and Gerhard Gompper, 2006b, “Mechanical deformation of spherical viruses with icosahedral symmetry,” Biophys. J. 91, 834.
- Vogel, V., and M. P. Sheetz, 2006, “Local force and geometry sensing regulate cell functions,” Nat. Rev. Mol. Cell Biol. 7, 265.
- Vogel, V., and M. P. Sheetz, 2009, “Cell fate regulation by coupling mechanical cycles to biochemical signaling pathways,” Curr. Opin. Cell Biol. 21, 38.
- von Wichert, G., B. Haimovich, G. S. Feng, and M. P. Sheetz, 2003, “Force-dependent integrin-cytoskeleton linkage formation requires downregulation of focal complex dynamics by shp2,” EMBO J. 22, 5023.
- Wagner, H., and H. Horner, 1974, “Elastic interaction and phase-transition in coherent metal-hydrogen systems,” Adv. Phys. 23, 587.
- Walcott, S., and S. X. Sun, 2010, “A mechanical model of actin stress fiber formation and substrate elasticity sensing in adherent cells,” Proc. Natl. Acad. Sci. U.S.A. 107, 7757.
- Walcott, Sam, Dong-Hwee Kim, Denis Wirz, and Sean X. Sun, 2011, “Nucleation and decay initiation are the stiffness-sensitive phases of focal adhesion maturation,” Biophys. J. 101, 2919.
- Wang, J. H. C., P. Goldschmidt-Clermont, J. Wille, and F. C. P. Yin, 2001, “Specificity of endothelial cell reorientation in response to cyclic mechanical stretching,” J. Biomech. 34, 1563.
- Wang, J. H. C., and E. S. Grood, 2000, “The strain magnitude and contact guidance determine orientation response of fibroblasts to cyclic substrate strains,” Connect. Tissue Res. 41, 29.
- Wang, N., J. D. Tytell, and D. E. Ingber, 2009, “Mechanotransduction at a distance: mechanically coupling the extracellular matrix with the nucleus,” Nat. Rev. Mol. Cell Biol. 10, 75.
- Wei, Z., V. S. Deshpande, R. M. McMeeking, and A. G. Evans, 2008, “Analysis and interpretation of stress-fiber organization in cells subject to cyclic stretch,” J. Biomed. Eng. 130, 031009.
- Weichsel, J., and U. S. Schwarz, 2010, “Two competing orientation patterns explain experimentally observed anomalies in growing actin networks,” Proc. Natl. Acad. Sci. U.S.A. 107, 6304.
- Weichsel, Julian, Edit Urban, J. Victor Small, and Ulrich S. Schwarz, 2012, “Reconstructing the orientation distribution of actin filaments in the lamellipodium of migrating keratocytes from electron microscopy tomography data,” Cytometry Part A 81A, 496.
- Weikl, Thomas R., and Reinhard Lipowsky, 2001, “Adhesion-induced phase behavior of multicomponent membranes,” Phys. Rev. E 64, 011903.
- Wen, Q., and P. Janmey, 2013, “Effects of non-linearity on cell-ECM interactions,” Experimental Cell Research, in press [http://dx.doi.org/10.1016/j.yexcr.2013.05.017].
- Wilhelm, Jan, and Erwin Frey, 2003, “Elasticity of stiff polymer networks,” Phys. Rev. Lett. 91, 108103.
- Winer, J,, S. Oake, and P. A. Janmey, 2009, “Non-linear elasticity of extracellular matrices enables contractile cells to communicate local position and orientation,” PLoS One 4, e6382.
- Wolfenson, Haguy, Alexander Bershadsky, Yoav I. Henis, and Benjamin Geiger, 2011, “Actomyosin-generated tension controls the molecular kinetics of focal adhesions,” J. Cell Sci. 124, 1425.
- Wottawah, Falk, Stefan Schinkinger, Bryan Lincoln, Revathi Ananthakrishnan, Maren Romeyke, Jochen Guck, and Josef Kas, 2005, “Optical rheology of biological cells,” Phys. Rev. Lett. 94, 098103.
- Xu, J. Y., D. Wirtz, and T. D. Pollard, 1998, “Dynamic cross-linking by alpha-actinin determines the mechanical properties of actin filament networks,” J. Biol. Chem. 273, 9570.
- Yeung, T., P. C. Georges, L. A. Flanagan, B. Marg, M. Ortiz, M. Funaki, N. Zahir, W. Y. Ming, V. Weaver, and P. A. Janmey, 2005, “Effects of substrate stiffness on cell morphology, cytoskeletal structure, and adhesion,” Cell Motil. Cytoskeleton 60, 24.
- Yoshinaga, N., J.-F. Joanny, J. Prost, and P. Marcq, 2010, “Polarity patterns of stress fibers,” Phys. Rev. Lett. 105, 238103.
- Zahn, J., I. Louban, S. Jungbauer, M. Bissinger, D. Kaufmann, R. Kemkemer, and J. Spatz, 2011, “Age-dependent changes in microscale stiffness and mechanoresponses of cells,” Small 7, 1480.
- Zaidel-Bar, R., C. Ballestrem, Z. Kam, and B. Geiger, 2003, “Early molecular events in the assembly of matrix adhesions at the leading edge of migrating cells,” J. Cell Sci. 116, 4605.
- Zaidel-Bar, Ronen, Shalev Itzkovitz, Avi Ma’ayan, Ravi Iyengar, and Benjamin Geiger, 2007, “Functional atlas of the integrin adhesome,” Nat. Cell Biol. 9, 858.
- Zand, Martin S., and Guenter Albrecht-Buehler, 1989, “What structures, besides adhesions, prevent spread cells from rounding up?,” Cell Motil. Cytoskeleton 13, 195.
- Zemel, A., I. B. Bischofs, and S. A. Safran, 2006, “Active elasticity of gels with contractile cells,” Phys. Rev. Lett. 97, 128103.
- Zemel, A., F. Rehfeldt, A. E. X. Brown, D. E. Discher, and S. A. Safran, 2010a, “Cell shape, spreading symmetry, and the polarization of stress-fibers in cells,” J. Phys. Condens. Matter 22, 194110.
- Zemel, A., F. Rehfeldt, A. E. X. Brown, D. E. Discher, and S. A. Safran, 2010b, “Optimal matrix rigidity for stress-fibre polarization in stem cells,” Nat. Phys. 6, 468.
- Zemel, A., and S. A. Safran, 2007, “Active self-polarization of contractile cells in asymmetrically shaped domains,” Phys. Rev. E 76, 021905.
- Zeng, Y., A. K. Yip, S-K Teo, and K.-H. Chiam, 2012, “A three-dimensional random network model of the cytoskeleton and its role in mechanotransduction and nucleus deformation,” Biomechan. Model Mechanobiol. 11, 49.
- Zhao, X. H., C. Laschinger, P. Arora, K. Szàszi, A. Kapus, and C. A. McCulloch, 2007, “Force activates smooth muscle alpha-actin promoter activity through the rho signaling pathway,” J. Cell Sci. 120, 1801.
- Ziebert, F., and I. S. Aranson, 2013, “Effects of Adhesion Dynamics and Substrate Compliance on the Shape and Motility of Crawling Cells,” PLoS One 8, e64511.
- Ziebert, Falko, Sumanth Swaminathan, and Igor S. Aranson, 2012, “Model for Self-Polarization and motility of keratocyte fragments,” J. R. Soc. Interface 9, 1084.
- Ziebert, Falko, and Walter Zimmermann, 2004, “Pattern formation driven by nematic ordering of assembling biopolymers,” Phys. Rev. E 70, 022902.
- Zilman, A. G., and S. A. Safran, 2003, “Role of cross-links in bundle formation, phase separation and gelation of long filaments,” Europhys. Lett. 63, 139.
- Zimmermann, Juliane, Claudia Brunner, Mihaela Enculescu, Michael Goegler, Allen Ehrlicher, Josef Kas, and Martin Falcke, 2012, “Actin filament elasticity and retrograde flow shape the Force-Velocity relation of motile cells,” Biophys. J. 102, 287.
- Zuckerman, D., and R. Bruinsma, 1995, “Statistical mechanics of membrane adhesion by reversible molecular bonds,” Phys. Rev. Lett. 74, 3900.