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1

Sunyer, Raimon, and Xavier Trepat. "Durotaxis." Current Biology 30, no. 9 (2020): R383—R387. http://dx.doi.org/10.1016/j.cub.2020.03.051.

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2

Huang, Yuxing, Jing Su, Jiayong Liu, et al. "YAP Activation in Promoting Negative Durotaxis and Acral Melanoma Progression." Cells 11, no. 22 (2022): 3543. http://dx.doi.org/10.3390/cells11223543.

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Directed cell migration towards a softer environment is called negative durotaxis. The mechanism and pathological relevance of negative durotaxis in tumor progression still requires in-depth investigation. Here, we report that YAP promotes the negative durotaxis of melanoma. We uncovered that the RhoA-myosin II pathway may underlie the YAP enhanced negative durotaxis of melanoma cells. Acral melanoma is the most common subtype of melanoma in non-Caucasians and tends to develop in a stress-bearing area. We report that acral melanoma patients exhibit YAP amplification and increased YAP activity.
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3

Puleo, Julieann I., Sara S. Parker, Mackenzie R. Roman, et al. "Mechanosensing during directed cell migration requires dynamic actin polymerization at focal adhesions." Journal of Cell Biology 218, no. 12 (2019): 4215–35. http://dx.doi.org/10.1083/jcb.201902101.

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The mechanical properties of a cell’s microenvironment influence many aspects of cellular behavior, including cell migration. Durotaxis, the migration toward increasing matrix stiffness, has been implicated in processes ranging from development to cancer. During durotaxis, mechanical stimulation by matrix rigidity leads to directed migration. Studies suggest that cells sense mechanical stimuli, or mechanosense, through the acto-myosin cytoskeleton at focal adhesions (FAs); however, FA actin cytoskeletal remodeling and its role in mechanosensing are not fully understood. Here, we show that the
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4

Style, R. W., Y. Che, S. J. Park, et al. "Patterning droplets with durotaxis." Proceedings of the National Academy of Sciences 110, no. 31 (2013): 12541–44. http://dx.doi.org/10.1073/pnas.1307122110.

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5

Hartman, Christopher D., Brett C. Isenberg, Samantha G. Chua, and Joyce Y. Wong. "Vascular smooth muscle cell durotaxis depends on extracellular matrix composition." Proceedings of the National Academy of Sciences 113, no. 40 (2016): 11190–95. http://dx.doi.org/10.1073/pnas.1611324113.

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Mechanical compliance has been demonstrated to be a key determinant of cell behavior, directing processes such as spreading, migration, and differentiation. Durotaxis, directional migration from softer to more stiff regions of a substrate, has been observed for a variety of cell types. Recent stiffness mapping experiments have shown that local changes in tissue stiffness in disease are often accompanied by an altered ECM composition in vivo. However, the importance of ECM composition in durotaxis has not yet been explored. To address this question, we have developed and characterized a polyacr
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6

Yuehua, YANG, and JIANG Hongyuan. "Research Advances in Cell Durotaxis." 应用数学和力学 42, no. 10 (2021): 999–1007. http://dx.doi.org/10.21656/1000-0887.420265.

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7

Bueno, Jesus, Yuri Bazilevs, Ruben Juanes, and Hector Gomez. "Wettability control of droplet durotaxis." Soft Matter 14, no. 8 (2018): 1417–26. http://dx.doi.org/10.1039/c7sm01917c.

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8

Doering, Charles R., Xiaoming Mao, and Leonard M. Sander. "Random walker models for durotaxis." Physical Biology 15, no. 6 (2018): 066009. http://dx.doi.org/10.1088/1478-3975/aadc37.

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9

Stefanoni, Filippo, Maurizio Ventre, Francesco Mollica, and Paolo A. Netti. "A numerical model for durotaxis." Journal of Theoretical Biology 280, no. 1 (2011): 150–58. http://dx.doi.org/10.1016/j.jtbi.2011.04.001.

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10

Parida, Lipika, and Venkat Padmanabhan. "Durotaxis in Nematode Caenorhabditis elegans." Biophysical Journal 111, no. 3 (2016): 666–74. http://dx.doi.org/10.1016/j.bpj.2016.06.030.

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11

DuChez, Brian J., Andrew D. Doyle, Emilios K. Dimitriadis, and Kenneth M. Yamada. "Durotaxis by Human Cancer Cells." Biophysical Journal 116, no. 4 (2019): 670–83. http://dx.doi.org/10.1016/j.bpj.2019.01.009.

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12

Moriyama, Kousuke, and Satoru Kidoaki. "Cellular Durotaxis Revisited: Initial-Position-Dependent Determination of the Threshold Stiffness Gradient to Induce Durotaxis." Langmuir 35, no. 23 (2018): 7478–86. http://dx.doi.org/10.1021/acs.langmuir.8b02529.

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13

Feng, Jingchen, Herbert Levine, Xiaoming Mao, and Leonard M. Sander. "Cell motility, contact guidance, and durotaxis." Soft Matter 15, no. 24 (2019): 4856–64. http://dx.doi.org/10.1039/c8sm02564a.

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14

Novikova, Elizaveta A., Matthew Raab, Dennis E. Discher, and Cornelis Storm. "Cellular Durotaxis from Differentially Persistent Motility." Biophysical Journal 112, no. 3 (2017): 436a. http://dx.doi.org/10.1016/j.bpj.2016.11.2327.

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15

Lazopoulos, Konstantinos A., and Dimitrije Stamenović. "Durotaxis as an elastic stability phenomenon." Journal of Biomechanics 41, no. 6 (2008): 1289–94. http://dx.doi.org/10.1016/j.jbiomech.2008.01.008.

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16

Gomez, Hector, and Mirian Velay-Lizancos. "Thin-film model of droplet durotaxis." European Physical Journal Special Topics 229, no. 2-3 (2020): 265–73. http://dx.doi.org/10.1140/epjst/e2019-900127-x.

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17

Wei, Jie, Xiaofeng Chen, and Bin Chen. "Harnessing structural instability for cell durotaxis." Acta Mechanica Sinica 35, no. 2 (2019): 355–64. http://dx.doi.org/10.1007/s10409-019-00853-2.

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18

Kassianides, Christoforos, Alain Goriely, and Hadrien Oliveri. "The multiscale mechanics of axon durotaxis." Journal of the Mechanics and Physics of Solids 200 (July 2025): 106134. https://doi.org/10.1016/j.jmps.2025.106134.

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19

Raab, Matthew, Joe Swift, P. C. Dave P. Dingal, Palak Shah, Jae-Won Shin, and Dennis E. Discher. "Crawling from soft to stiff matrix polarizes the cytoskeleton and phosphoregulates myosin-II heavy chain." Journal of Cell Biology 199, no. 4 (2012): 669–83. http://dx.doi.org/10.1083/jcb.201205056.

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On rigid surfaces, the cytoskeleton of migrating cells is polarized, but tissue matrix is normally soft. We show that nonmuscle MIIB (myosin-IIB) is unpolarized in cells on soft matrix in 2D and also within soft 3D collagen, with rearward polarization of MIIB emerging only as cells migrate from soft to stiff matrix. Durotaxis is the tendency of cells to crawl from soft to stiff matrix, and durotaxis of primary mesenchymal stem cells (MSCs) proved more sensitive to MIIB than to the more abundant and persistently unpolarized nonmuscle MIIA (myosin-IIA). However, MIIA has a key upstream role: in
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20

McKenzie, Andrew J., Kathryn V. Svec, Tamara F. Williams, and Alan K. Howe. "Protein kinase A activity is regulated by actomyosin contractility during cell migration and is required for durotaxis." Molecular Biology of the Cell 31, no. 1 (2020): 45–58. http://dx.doi.org/10.1091/mbc.e19-03-0131.

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Here, we show that localized PKA activity in migrating cells is regulated by cell–matrix tension, correlates with cellular traction forces, is enhanced by acute mechanical stimulation, and is required for durotaxis. This establishes PKA as an effector of cellular mechanotransduction and as a regulator of mechanically guided cell migration.
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21

Liu, Yang, Jiwen Cheng, Hui Yang, and Guang-Kui Xu. "Rotational constraint contributes to collective cell durotaxis." Applied Physics Letters 117, no. 21 (2020): 213702. http://dx.doi.org/10.1063/5.0031846.

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22

Harland, Ben, Sam Walcott, and Sean X. Sun. "Adhesion dynamics and durotaxis in migrating cells." Physical Biology 8, no. 1 (2011): 015011. http://dx.doi.org/10.1088/1478-3975/8/1/015011.

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23

Harland, Ben, Sam Walcott, and Sean X. Sun. "Adhesion Dynamics and Durotaxis in Migrating Cells." Biophysical Journal 100, no. 3 (2011): 303a. http://dx.doi.org/10.1016/j.bpj.2010.12.1855.

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24

Jain, Gaurav, Andrew J. Ford, and Padmavathy Rajagopalan. "Opposing Rigidity-Protein Gradients Reverse Fibroblast Durotaxis." ACS Biomaterials Science & Engineering 1, no. 8 (2015): 621–31. http://dx.doi.org/10.1021/acsbiomaterials.5b00229.

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25

Riaz, Maryam, Marie Versaevel, and Sylvain Gabriele. "On the Mechanism of Durotaxis in Motile Cells." Biophysical Journal 106, no. 2 (2014): 571a. http://dx.doi.org/10.1016/j.bpj.2013.11.3167.

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26

Escribano, Jorge, Raimon Sunyer, María Teresa Sánchez, Xavier Trepat, Pere Roca-Cusachs, and José Manuel García-Aznar. "A hybrid computational model for collective cell durotaxis." Biomechanics and Modeling in Mechanobiology 17, no. 4 (2018): 1037–52. http://dx.doi.org/10.1007/s10237-018-1010-2.

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27

Vicente-Manzanares, Miguel. "Cell Migration: Cooperation between Myosin II Isoforms in Durotaxis." Current Biology 23, no. 1 (2013): R28—R29. http://dx.doi.org/10.1016/j.cub.2012.11.024.

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28

Vicente-Manzanares, Miguel. "Cell Migration: Cooperation between Myosin II Isoforms in Durotaxis." Current Biology 23, no. 5 (2013): 441. http://dx.doi.org/10.1016/j.cub.2013.02.014.

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29

Wieland, Annalena, Pamela L. Strissel, Hannah Schorle, et al. "Brain and Breast Cancer Cells with PTEN Loss of Function Reveal Enhanced Durotaxis and RHOB Dependent Amoeboid Migration Utilizing 3D Scaffolds and Aligned Microfiber Tracts." Cancers 13, no. 20 (2021): 5144. http://dx.doi.org/10.3390/cancers13205144.

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Background: Glioblastoma multiforme (GBM) and metastatic triple-negative breast cancer (TNBC) with PTEN mutations often lead to brain dissemination with poor patient outcome, thus new therapeutic targets are needed. To understand signaling, controlling the dynamics and mechanics of brain tumor cell migration, we implemented GBM and TNBC cell lines and designed 3D aligned microfibers and scaffolds mimicking brain structures. Methods: 3D microfibers and scaffolds were printed using melt electrowriting. GBM and TNBC cell lines with opposing PTEN genotypes were analyzed with RHO-ROCK-PTEN inhibito
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30

Wieland, Annalena, Pamela L. Strissel, Hannah Schorle, et al. "Brain and Breast Cancer Cells with PTEN Loss of Function Reveal Enhanced Durotaxis and RHOB Dependent Amoeboid Migration Utilizing 3D Scaffolds and Aligned Microfiber Tracts." Cancers 13, no. 20 (2021): 5144. http://dx.doi.org/10.3390/cancers13205144.

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Background: Glioblastoma multiforme (GBM) and metastatic triple-negative breast cancer (TNBC) with PTEN mutations often lead to brain dissemination with poor patient outcome, thus new therapeutic targets are needed. To understand signaling, controlling the dynamics and mechanics of brain tumor cell migration, we implemented GBM and TNBC cell lines and designed 3D aligned microfibers and scaffolds mimicking brain structures. Methods: 3D microfibers and scaffolds were printed using melt electrowriting. GBM and TNBC cell lines with opposing PTEN genotypes were analyzed with RHO-ROCK-PTEN inhibito
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31

Shellard, Adam, and Roberto Mayor. "Collective durotaxis along a self-generated stiffness gradient in vivo." Nature 600, no. 7890 (2021): 690–94. http://dx.doi.org/10.1038/s41586-021-04210-x.

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32

Sunyer, R., V. Conte, J. Escribano, et al. "Collective cell durotaxis emerges from long-range intercellular force transmission." Science 353, no. 6304 (2016): 1157–61. http://dx.doi.org/10.1126/science.aaf7119.

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33

Martinez, Jessica S., Ali M. Lehaf, Joseph B. Schlenoff, and Thomas C. S. Keller. "Cell Durotaxis on Polyelectrolyte Multilayers with Photogenerated Gradients of Modulus." Biomacromolecules 14, no. 5 (2013): 1311–20. http://dx.doi.org/10.1021/bm301863a.

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34

Vincent, Ludovic G., Yu Suk Choi, Baldomero Alonso-Latorre, Juan C. del Álamo, and Adam J. Engler. "Mesenchymal stem cell durotaxis depends on substrate stiffness gradient strength." Biotechnology Journal 8, no. 4 (2013): 472–84. http://dx.doi.org/10.1002/biot.201200205.

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35

Pamonag, Michael, Abigail Hinson, Elisha J. Burton, et al. "Individual cells generate their own self-reinforcing contact guidance cues through local matrix fiber remodeling." PLOS ONE 17, no. 3 (2022): e0265403. http://dx.doi.org/10.1371/journal.pone.0265403.

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Directed cell migration arises from cells following a microenvironmental gradient (e.g. of a chemokine) or polarizing feature (e.g. a linear structure). However cells not only follow, but in many cases, also generate directionality cues by modifying their microenvironment. This bi-directional relationship is seen in the alignment of extracellular matrix (ECM) fibers ahead of invading cell masses. The forces generated by many migrating cells cause fiber alignment, which in turn promotes further migration in the direction of fiber alignment via contact guidance and durotaxis. While this positive
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36

Aubry, D., M. Gupta, B. Ladoux, and R. Allena. "Mechanical link between durotaxis, cell polarity and anisotropy during cell migration." Physical Biology 12, no. 2 (2015): 026008. http://dx.doi.org/10.1088/1478-3975/12/2/026008.

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37

Isenberg, Brett C., Paul A. DiMilla, Matthew Walker, Sooyoung Kim, and Joyce Y. Wong. "Vascular Smooth Muscle Cell Durotaxis Depends on Substrate Stiffness Gradient Strength." Biophysical Journal 97, no. 5 (2009): 1313–22. http://dx.doi.org/10.1016/j.bpj.2009.06.021.

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38

Kuntanawat, P., C. Wilkinson, and M. Riehle. "Observation of durotaxis on a well-defined continuous gradient of stiffness." Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 146, no. 4 (2007): S192. http://dx.doi.org/10.1016/j.cbpa.2007.01.421.

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39

Wormer, Duncan B., Kevin A. Davis, James H. Henderson, and Christopher E. Turner. "The Focal Adhesion-Localized CdGAP Regulates Matrix Rigidity Sensing and Durotaxis." PLoS ONE 9, no. 3 (2014): e91815. http://dx.doi.org/10.1371/journal.pone.0091815.

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40

Ebata, Hiroyuki, Kousuke Moriyama, Thasaneeya Kuboki, and Satoru Kidoaki. "General cellular durotaxis induced with cell-scale heterogeneity of matrix-elasticity." Biomaterials 230 (February 2020): 119647. http://dx.doi.org/10.1016/j.biomaterials.2019.119647.

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41

Shellard, Adam, and Roberto Mayor. "Publisher Correction: Collective durotaxis along a self-generated stiffness gradient in vivo." Nature 601, no. 7894 (2022): E33. http://dx.doi.org/10.1038/s41586-021-04367-5.

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42

Budde, Ilka, David Ing, Albrecht Schwab, and Zoltan Denes Petho. "Mechanosensitive ion channels are essential for the durotaxis of pancreatic stellate cells." Biophysical Journal 121, no. 3 (2022): 314a. http://dx.doi.org/10.1016/j.bpj.2021.11.1181.

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43

Alert, Ricard, and Jaume Casademunt. "Role of Substrate Stiffness in Tissue Spreading: Wetting Transition and Tissue Durotaxis." Langmuir 35, no. 23 (2018): 7571–77. http://dx.doi.org/10.1021/acs.langmuir.8b02037.

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44

Allena, R., M. Scianna, and L. Preziosi. "A Cellular Potts Model of single cell migration in presence of durotaxis." Mathematical Biosciences 275 (May 2016): 57–70. http://dx.doi.org/10.1016/j.mbs.2016.02.011.

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45

Malik, Adam A., and Philip Gerlee. "Mathematical modelling of cell migration: stiffness dependent jump rates result in durotaxis." Journal of Mathematical Biology 78, no. 7 (2019): 2289–315. http://dx.doi.org/10.1007/s00285-019-01344-5.

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46

Whang, Minji, and Jungwook Kim. "Synthetic hydrogels with stiffness gradients for durotaxis study and tissue engineering scaffolds." Tissue Engineering and Regenerative Medicine 13, no. 2 (2016): 126–39. http://dx.doi.org/10.1007/s13770-016-0026-x.

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47

Marzban, Bahador, Xin Yi, and Hongyan Yuan. "A minimal mechanics model for mechanosensing of substrate rigidity gradient in durotaxis." Biomechanics and Modeling in Mechanobiology 17, no. 3 (2018): 915–22. http://dx.doi.org/10.1007/s10237-018-1001-3.

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48

Zhang, Zhiwen, Phoebus Rosakis, Thomas Y. Hou, and Guruswami Ravichandran. "A minimal mechanosensing model predicts keratocyte evolution on flexible substrates." Journal of The Royal Society Interface 17, no. 166 (2020): 20200175. http://dx.doi.org/10.1098/rsif.2020.0175.

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A mathematical model is proposed for shape evolution and locomotion of fish epidermal keratocytes on elastic substrates. The model is based on mechanosensing concepts: cells apply contractile forces onto the elastic substrate, while cell shape evolution depends locally on the substrate stress generated by themselves or external mechanical stimuli acting on the substrate. We use the level set method to study the behaviour of the model numerically, and predict a number of distinct phenomena observed in experiments, such as (i) symmetry breaking from the stationary centrosymmetric to the well-kno
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49

Lachowski, Dariusz, Ernesto Cortes, Benjamin Robinson, Alistair Rice, Krista Rombouts, and Armando E. Del Río Hernández. "FAK controls the mechanical activation of YAP, a transcriptional regulator required for durotaxis." FASEB Journal 32, no. 2 (2018): 1099–107. http://dx.doi.org/10.1096/fj.201700721r.

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50

Walker, Matthew L., David House, Margrit Betke, and Joyce Y. Wong. "Using Automated Cell Tracking Software to Quantifying Durokinesis and Durotaxis in Real Time." Biophysical Journal 96, no. 3 (2009): 633a. http://dx.doi.org/10.1016/j.bpj.2008.12.3347.

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