{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,13]],"date-time":"2026-11-13T04:30:29Z","timestamp":1794544229162,"version":"4.9.1"},"reference-count":87,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,7,25]],"date-time":"2026-07-25T00:00:00Z","timestamp":1784937600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T00:00:00Z","timestamp":1787788800000},"content-version":"vor","delay-in-days":33,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Commun"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Locomotion - whether walking, running, or crawling - depends on the precise coordination of forces between the body and its surroundings. Two critical factors in this process are the force that resists the relative motion between two bodies, and mechanosensation, the body\u2019s ability to sense and respond to mechanical forces. Together, these processes enable efficient locomotion across diverse environments, yet how body-environment interaction forces shape neural activity in freely moving animals remains poorly understood. Here we show that the \u2018gentle touch\u2019 receptor neurons (TRNs) in the\n                    <jats:italic>Caenorhabditis elegans<\/jats:italic>\n                    body wall are sensitive to dynamic surface mechanics. Using a combination of calcium imaging and traction force microscopy in freely behaving animals informed by micromechanical stimulation, connectome-scale simulations, and optimal control theory, we identify friction force and crawling velocity as key determinants of mechanoreceptor activity. Mutations disrupting MEC-4 DEG\/ENaC ion channel activity in body wall mechanoreceptors produce lethargic animals with impaired proprioceptive regulation, suggesting functional coupling between surface mechanoreceptors and proprioceptors. Together, our findings reveal that gentle \u2018touch\u2019 receptors encode body-environment interaction forces\u00a0during movement to optimize locomotor energetics.\n                  <\/jats:p>","DOI":"10.1038\/s41467-026-75352-7","type":"journal-article","created":{"date-parts":[[2026,7,25]],"date-time":"2026-07-25T15:24:34Z","timestamp":1784993074000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Mechanosensory encoding of\u00a0surface mechanics optimizes locomotion"],"prefix":"10.1038","volume":"17","author":[{"given":"Aleksandra","family":"Pidde","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1514-2883","authenticated-orcid":false,"given":"Montserrat","family":"Porta-de-la-Riva","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Costanza","family":"Agazzi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carmen","family":"Mart\u00ednez-Fern\u00e1ndez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alice","family":"Lorrach","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ashutosh","family":"Bijalwan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Neus","family":"Sanfeliu-Cerd\u00e1n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alba","family":"Calatayud-Sanchez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-7612-2792","authenticated-orcid":false,"given":"Eric","family":"Torralba-Sales","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ravi","family":"Das","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9 J.","family":"Mu\u00f1oz","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0501-5036","authenticated-orcid":false,"given":"Michael","family":"Krieg","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,25]]},"reference":[{"key":"75352_CR1","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1146\/annurev-cellbio-100913-013426","volume":"31","author":"S Katta","year":"2015","unstructured":"Katta, S., Krieg, M. & Goodman, M. B. Feeling force: Physical and physiological principles enabling sensory mechanotransduction. Annu. Rev. Cell Dev. Biol. 31, 347\u2013371 (2015).","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"75352_CR2","doi-asserted-by":"publisher","first-page":"260","DOI":"10.1016\/j.neubiorev.2008.07.001","volume":"34","author":"RI Dunbar","year":"2010","unstructured":"Dunbar, R. I. The social role of touch in humans and primates: behavioural function and neurobiological mechanisms. Neurosci. Biobehav. Rev. 34, 260\u2013268 (2010).","journal-title":"Neurosci. Biobehav. Rev."},{"key":"75352_CR3","doi-asserted-by":"publisher","first-page":"737","DOI":"10.1016\/j.neuron.2014.05.001","volume":"82","author":"F McGlone","year":"2014","unstructured":"McGlone, F., Wessberg, J. & Olausson, H. Discriminative and affective touch: sensing and feeling. Neuron 82, 737\u2013755 (2014).","journal-title":"Neuron"},{"key":"75352_CR4","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1007\/s00221-003-1804-y","volume":"156","author":"PF Meyer","year":"2004","unstructured":"Meyer, P. F., Oddsson, L. I. & Luca, C. J. D. The role of plantar cutaneous sensation in unperturbed stance. Exp. Brain Res. 156, 505\u2013512 (2004).","journal-title":"Exp. Brain Res."},{"key":"75352_CR5","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/S0966-6362(03)00095-X","volume":"20","author":"E Eils","year":"2004","unstructured":"Eils, E. et al. Reduced plantar sensation causes a cautious walking pattern. Gait Posture 20, 54\u201360 (2004).","journal-title":"Gait Posture"},{"key":"75352_CR6","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1007\/s00221-004-1868-3","volume":"157","author":"PF Meyer","year":"2004","unstructured":"Meyer, P. F., Oddsson, L. I. E. & Luca, C. J. D. Reduced plantar sensitivity alters postural responses to lateral perturbations of balance. Exp. Brain Res 157, 526\u2013536 (2004).","journal-title":"Exp. Brain Res"},{"key":"75352_CR7","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1534\/genetics.118.300241","volume":"212","author":"MB Goodman","year":"2019","unstructured":"Goodman, M. B. & Sengupta, P. How Caenorhabditis elegans senses mechanical stress, temperature, and other physical stimuli. Genetics 212, 25\u201351 (2019).","journal-title":"Genetics"},{"key":"75352_CR8","doi-asserted-by":"publisher","first-page":"1116","DOI":"10.1039\/C6LC01165A","volume":"17","author":"AL Nekimken","year":"2017","unstructured":"Nekimken, A. L. et al. Pneumatic stimulation of C. elegans mechanoreceptor neurons in a microfluidic trap. Lab Chip 17, 1116\u20131127 (2017).","journal-title":"Lab Chip"},{"key":"75352_CR9","doi-asserted-by":"publisher","first-page":"358","DOI":"10.1016\/0012-1606(81)90459-0","volume":"82","author":"M Chalfie","year":"1981","unstructured":"Chalfie, M. & Sulston, J. Developmental genetics of the mechanosensory neurons of Caenorhabditis elegans. Dev. Biol. 82, 358\u2013370 (1981).","journal-title":"Dev. Biol."},{"key":"75352_CR10","doi-asserted-by":"crossref","unstructured":"Chew, Y. L. et al. An Afferent Neuropeptide System Transmits Mechanosensory Signals Triggering Sensitization and Arousal in C. elegans. Neuron 99, 1233\u20131246 (2018).","DOI":"10.1016\/j.neuron.2018.08.003"},{"key":"75352_CR11","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-020-60994-4","volume":"10","author":"PD McClanahan","year":"2020","unstructured":"McClanahan, P. D. et al. A quiescent state following mild sensory arousal in Caenorhabditis elegans is potentiated by stress. Sci. Rep. 10, 1\u201316 (2020).","journal-title":"Sci. Rep."},{"key":"75352_CR12","doi-asserted-by":"crossref","unstructured":"Staum, M. et al. Behavioral adjustment of C. elegans to mechanosensory loss requires intact mechanosensory neurons. PLoS Biol. 22, e3002729 (2024).","DOI":"10.1371\/journal.pbio.3002729"},{"key":"75352_CR13","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1038\/367470a0","volume":"367","author":"K Hong","year":"1994","unstructured":"Hong, K. & Driscoll, M. A transmembrane domain of the putative channel subunit MEC-4 influences mechanotransduction and neurodegeneration in C. elegans. Nature 367, 470\u2013473 (1994).","journal-title":"Nature"},{"key":"75352_CR14","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1038\/nn1362","volume":"8","author":"R O\u2019Hagan","year":"2005","unstructured":"O\u2019Hagan, R., Chalfie, M. & Goodman, M. B. The MEC-4 deg\/enac channel of Caenorhabditis elegans touch receptor neurons transduces mechanical signals. Nat. Neurosci. 8, 43\u201350 (2005).","journal-title":"Nat. Neurosci."},{"key":"75352_CR15","doi-asserted-by":"publisher","first-page":"2383","DOI":"10.1242\/jeb.004572","volume":"210","author":"J Korta","year":"2007","unstructured":"Korta, J., Clark, D. A., Gabel, C. V., Mahadevan, L. & Samuel, A. D. T. Mechanosensation and mechanical load modulate the locomotory gait of swimming C. elegans. J. Exp. Biol. 210, 2383\u20132389 (2007).","journal-title":"J. Exp. Biol."},{"key":"75352_CR16","doi-asserted-by":"publisher","first-page":"2791","DOI":"10.1016\/j.bpj.2012.04.051","volume":"102","author":"F Lebois","year":"2012","unstructured":"Lebois, F. et al. Locomotion control of Caenorhabditis elegans through confinement. Biophys. J. 102, 2791\u20132798 (2012).","journal-title":"Biophys. J."},{"key":"75352_CR17","doi-asserted-by":"publisher","first-page":"14012","DOI":"10.1074\/jbc.M116.718031","volume":"291","author":"S Shi","year":"2016","unstructured":"Shi, S., Luke, C. J., Miedel, M. T., Silverman, G. A. & Kleyman, T. R. Activation of the Caenorhabditis elegans degenerin channel by shear stress requires the MEC-10 subunit. J. Biol. Chem. 291, 14012\u201314022 (2016).","journal-title":"J. Biol. Chem."},{"key":"75352_CR18","first-page":"E2471","volume":"113","author":"AL Eastwood","year":"2015","unstructured":"Eastwood, A. L. et al. Tissue mechanics govern the rapidly adapting and symmetrical response to touch. Proc. Natl. Acad. Sci. USA 113, E2471\u2013E2471 (2015).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"75352_CR19","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1113\/jphysiol.1966.sp007827","volume":"182","author":"BYWR Loewenstein","year":"1966","unstructured":"Loewenstein, B. Y. W. R. & Skalak, R. Mechanical transmission in a Pacinian corpuscle: an analysis and a theory. J. Physiol. 182, 346\u2013378 (1966).","journal-title":"J. Physiol."},{"key":"75352_CR20","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.cell.2021.06.023","volume":"184","author":"SR Taylor","year":"2021","unstructured":"Taylor, S. R. et al. Molecular topography of an entire nervous system. Cell 184, 1\u201319 (2021).","journal-title":"Cell"},{"key":"75352_CR21","doi-asserted-by":"crossref","unstructured":"Xu, S. & Chisholm, A. D. Highly efficient optogenetic cell ablation in C. elegans using membrane-targeted miniSOG. Sci. Rep. 6, 21271 (2016).","DOI":"10.1038\/srep21271"},{"key":"75352_CR22","doi-asserted-by":"publisher","first-page":"647","DOI":"10.1126\/science.aaa7484","volume":"349","author":"EG Govorunova","year":"2015","unstructured":"Govorunova, E. G., Sineshchekov, O. A., Janz, R., Liu, X. & Spudich, J. L. Natural light-gated anion channels: a family of microbial rhodopsins for advanced optogenetics. Science (New York, NY) 349, 647\u2013650 (2015).","journal-title":"Science (New York, NY)"},{"key":"75352_CR23","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1038\/nmeth.4109","volume":"14","author":"H Wang","year":"2017","unstructured":"Wang, H. et al. Cgal, a temperature-robust gal4-uas system for Caenorhabditis elegans. Nat. Methods 14, 145\u2013148 (2017).","journal-title":"Nat. Methods"},{"key":"75352_CR24","doi-asserted-by":"publisher","first-page":"2279","DOI":"10.1016\/j.cub.2005.11.032","volume":"15","author":"G Nagel","year":"2005","unstructured":"Nagel, G. et al. Light activation of channelrhodopsin-2 in excitable cells of Caenorhabditis elegans triggers rapid behavioral responses. Curr. Biol. 15, 2279\u20132284 (2005).","journal-title":"Curr. Biol."},{"key":"75352_CR25","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1016\/j.cub.2012.02.066","volume":"22","author":"SJ Husson","year":"2012","unstructured":"Husson, S. J. et al. Optogenetic analysis of a nociceptor neuron and network reveals ion channels acting downstream of primary sensors. Curr. Biol. 22, 743\u2013752 (2012).","journal-title":"Curr. Biol."},{"key":"75352_CR26","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1038\/ncb2915","volume":"16","author":"M Krieg","year":"2014","unstructured":"Krieg, M., Dunn, A. R. & Goodman, M. B. Mechanical control of the sense of touch by \u03b2-spectrin. Nat. Cell Biol. 16, 224\u2013233 (2014).","journal-title":"Nat. Cell Biol."},{"key":"75352_CR27","doi-asserted-by":"publisher","first-page":"1","DOI":"10.7554\/eLife.29913","volume":"7","author":"AD Fouad","year":"2018","unstructured":"Fouad, A. D. et al. Distributed rhythm generators underlie Caenorhabditis elegans forward locomotion. eLife 7, 1\u201334 (2018).","journal-title":"eLife"},{"key":"75352_CR28","doi-asserted-by":"crossref","unstructured":"Bitzenhofer, S. H. et al. Layer-specific optogenetic activation of pyramidal neurons causes beta-gamma entrainment of neonatal networks. Nat. Commun. 8, 14563 (2017).","DOI":"10.1038\/ncomms14563"},{"key":"75352_CR29","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-019-08412-w","volume":"10","author":"LA Wurmthaler","year":"2019","unstructured":"Wurmthaler, L. A., Sack, M., Gense, K., Hartig, J. S. & Gamerdinger, M. A tetracycline-dependent ribozyme switch allows conditional induction of gene expression in Caenorhabditis elegans. Nat. Commun. 10, 1\u20138 (2019).","journal-title":"Nat. Commun."},{"key":"75352_CR30","doi-asserted-by":"crossref","unstructured":"Fang-Yen, C. et al. Biomechanical analysis of gait adaptation in the nematode Caenorhabditis elegans. Proc. Natl. Acad. Sci. USA 107, 20323\u201320328 (2010).","DOI":"10.1073\/pnas.1003016107"},{"key":"75352_CR31","doi-asserted-by":"publisher","first-page":"257","DOI":"10.1038\/s41586-021-03778-8","volume":"596","author":"D Witvliet","year":"2021","unstructured":"Witvliet, D. et al. Connectomes across development reveal principles of brain maturation. Nature 596, 257\u2013261 (2021).","journal-title":"Nature"},{"key":"75352_CR32","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1103\/PhysRevE.89.052805","volume":"89","author":"J Kunert","year":"2014","unstructured":"Kunert, J., Shlizerman, E. & Kutz, J. N. Low-dimensional functionality of complex network dynamics: Neurosensory integration in the Caenorhabditis elegans connectome. Phys. Rev. E\u2014Stat. Nonlinear Soft Matter Phys. 89, 1\u20136 (2014).","journal-title":"Phys. Rev. E\u2014Stat. Nonlinear Soft Matter Phys."},{"key":"75352_CR33","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pcbi.1005261","volume":"13","author":"JM Kunert","year":"2017","unstructured":"Kunert, J. M., Maia, P. D. & Kutz, J. N. Functionality and robustness of injured connectomic dynamics in C. elegans: linking behavioral deficits to neural circuit damage. PLoS Comput. Biol. 13, 1\u201321 (2017).","journal-title":"PLoS Comput. Biol."},{"key":"75352_CR34","doi-asserted-by":"crossref","unstructured":"Stephens, G. J., Johnson-Kerner, B., Bialek, W. & Ryu, W. S. Dimensionality and dynamics in the behavior of C. elegans. PLoS Comput. Biol. 4, e1000028 (2008).","DOI":"10.1371\/journal.pcbi.1000028"},{"key":"75352_CR35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1126\/sciadv.abg4617","volume":"7","author":"R Das","year":"2021","unstructured":"Das, R. et al. An asymmetric mechanical code ciphers curvature-dependent proprioceptor activity. Sci. Adv. 7, 1\u201320 (2021).","journal-title":"Sci. Adv."},{"key":"75352_CR36","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1038\/nn.2581","volume":"13","author":"M Chatzigeorgiou","year":"2010","unstructured":"Chatzigeorgiou, M. et al. Specific roles for DEG\/ENaC and TRP channels in touch and thermosensation in C. elegans nociceptors. Nat. Neurosci. 13, 861\u20138 (2010).","journal-title":"Nat. Neurosci."},{"key":"75352_CR37","first-page":"272\u2013288.e11","volume":"184","author":"E Yemini","year":"2020","unstructured":"Yemini, E. et al. Neuropal: a multicolor atlas for whole-brain neuronal identification in C. elegans. Cell 184, 272\u2013288.e11 (2020).","journal-title":"Cell"},{"key":"75352_CR38","doi-asserted-by":"publisher","first-page":"956","DOI":"10.1523\/JNEUROSCI.05-04-00956.1985","volume":"5","author":"M Chalfie","year":"1985","unstructured":"Chalfie, M. et al. The neural circuit for touch sensitivity in Caenorhabditis elegans.  J. Neurosci. 5, 956\u2013964 (1985).","journal-title":"J. Neurosci."},{"key":"75352_CR39","doi-asserted-by":"crossref","unstructured":"Li, T., Kumar, S., Poon, H., Leifer, A. M. & Zheng, C. Functional probing of neuronal subtypes via intersectional expression of optogenetic actuators reveals non-linear components in a linear circuit. Cell Rep. 44, 116327 (2025).","DOI":"10.1016\/j.celrep.2025.116327"},{"key":"75352_CR40","doi-asserted-by":"crossref","unstructured":"Rabets, Y., Backholm, M., Dalnoki-Veress, K. & Ryu, W. S. Direct measurements of drag forces in C. elegans crawling locomotion. Biophys J. 107, 1980\u20131987 (2014).","DOI":"10.1016\/j.bpj.2014.09.006"},{"key":"75352_CR41","doi-asserted-by":"publisher","first-page":"1420","DOI":"10.1111\/pbi.12726","volume":"15","author":"M Rajaee","year":"2017","unstructured":"Rajaee, M. & Ow, D. W. A new location to split cre recombinase for protein fragment complementation. Plant Biotechnol. J. 15, 1420\u20131428 (2017).","journal-title":"Plant Biotechnol. J."},{"key":"75352_CR42","doi-asserted-by":"crossref","unstructured":"van der Vaart, A., Godfrey, M. & Portegijs, V. Dose-dependent functions of SWI\/SNF BAF in permitting and inhibiting cell proliferation in vivo. Sci. Adv. 6, eaay3823 (2020).","DOI":"10.1126\/sciadv.aay3823"},{"key":"75352_CR43","doi-asserted-by":"crossref","unstructured":"Goudeau, J. et al. Split-wrmscarlet and split-sfgfp: tools for faster, easier fluorescent labeling of endogenous proteins in Caenorhabditis elegans. Genetics. 217, iyab014 (2021).","DOI":"10.1093\/genetics\/iyab014"},{"key":"75352_CR44","doi-asserted-by":"crossref","unstructured":"Mu\u00f1oz, J. J., Condamin, L. & Doste, D. On the net displacement of contact surface centroid in contractile bodies. Mech. Res. Commun. 119, 103809 (2022).","DOI":"10.1016\/j.mechrescom.2021.103809"},{"key":"75352_CR45","doi-asserted-by":"crossref","unstructured":"Keaveny, E. E., Brown, A. E. Predicting path from undulations for C. elegans using linear and nonlinear resistive force theory. Phys. Biol. 14, 025001 (2017).","DOI":"10.1088\/1478-3975\/aa5ce6"},{"key":"75352_CR46","doi-asserted-by":"crossref","unstructured":"Wang, B., Zhao, P., Rong, Y. et al. Multibody modeling and resistive coefficient identification for Caenorhabditis elegans swimming in viscous fluids. Multibody Syst. Dyn. 66, 721\u2013758 (2026).","DOI":"10.1007\/s11044-025-10101-0"},{"key":"75352_CR47","doi-asserted-by":"publisher","first-page":"6600","DOI":"10.1016\/S0021-9258(18)53293-2","volume":"268","author":"ML Blaxter","year":"1993","unstructured":"Blaxter, M. L. Cuticle surface proteins of wild type and mutant Caenorhabditis elegans.  J. Biol. Chem. 268, 6600\u20136609 (1993).","journal-title":"J. Biol. Chem."},{"key":"75352_CR48","doi-asserted-by":"publisher","first-page":"52893","DOI":"10.1074\/jbc.M409557200","volume":"279","author":"JF Cipollo","year":"2004","unstructured":"Cipollo, J. F., Awad, A. M., Costello, C. E. & Hirschberg, C. B. SRF-3, a mutant of Caenorhabditis elegans, resistant to bacterial infection and to biofilm binding, is deficient in glycoconjugates. J. Biol. Chem. 279, 52893\u201352903 (2004).","journal-title":"J. Biol. Chem."},{"key":"75352_CR49","doi-asserted-by":"publisher","first-page":"1366","DOI":"10.1091\/mbc.e02-08-0479","volume":"14","author":"L Mcmahon","year":"2003","unstructured":"Mcmahon, L., Muriel, J. M., Roberts, B., Quinn, M. & Johnstone, I. L. Two sets of interacting collagens form functionally distinct substructures within a Caenorhabditis elegans extracellular matrix. Mol. Biol. Cell 14, 1366\u20131378 (2003).","journal-title":"Mol. Biol. Cell"},{"key":"75352_CR50","doi-asserted-by":"publisher","first-page":"2989","DOI":"10.1098\/rstb.2011.0167","volume":"366","author":"TJ Prescott","year":"2011","unstructured":"Prescott, T. J., Diamond, M. E. & Wing, A. M. Active touch sensing. Philos. Trans. R. Soc. B: Biol. Sci. 366, 2989\u20132995 (2011).","journal-title":"Philos. Trans. R. Soc. B: Biol. Sci."},{"key":"75352_CR51","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1016\/j.conb.2016.08.001","volume":"40","author":"NE Bush","year":"2016","unstructured":"Bush, N. E., Solla, S. A. & Hartmann, M. J. Whisking mechanics and active sensing. Curr. Opin. Neurobiol. 40, 178\u2013188 (2016).","journal-title":"Curr. Opin. Neurobiol."},{"key":"75352_CR52","doi-asserted-by":"publisher","first-page":"1503","DOI":"10.1126\/science.1166467","volume":"323","author":"J Scheibert","year":"2009","unstructured":"Scheibert, J., Leurent, S., Prevost, A. & Debr\u00e9geas, G. The role of fingerprints in the coding of tactile information probed with a biomimetic sensor. Science 323, 1503\u20131507 (2009).","journal-title":"Science"},{"key":"75352_CR53","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1039\/B603209P","volume":"2","author":"JP Gong","year":"2006","unstructured":"Gong, J. P. Friction and lubrication of hydrogels\u2014its richness and complexity. Soft Matter 2, 544\u2013552 (2006).","journal-title":"Soft Matter"},{"key":"75352_CR54","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1083\/jcb.146.3.659","volume":"146","author":"MC Hresko","year":"1999","unstructured":"Hresko, M. C., Schriefer, L. A., Shrimankar, P. & Waterston, R. H. Myotactin, a novel hypodermal protein involved in muscle\u2013cell adhesion in Caenorhabditis elegans.  J. Cell Biol. 146, 659\u2013672 (1999).","journal-title":"J. Cell Biol."},{"key":"75352_CR55","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1007\/s11249-013-0225-8","volume":"52","author":"C Shen","year":"2013","unstructured":"Shen, C. Khonsari, M. M., Effect of dimple\u2019s internal structure on hydrodynamic lubrication. Tribol. Lett. 52, 415\u2013430 (2013).","journal-title":"Tribol. Lett."},{"key":"75352_CR56","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1534\/genetics.116.195511","volume":"206","author":"H Schulenburg","year":"2017","unstructured":"Schulenburg, H. & Felix, M. A. The natural biotic environment of Caenorhabditis elegans. Genetics 206, 55\u201386 (2017).","journal-title":"Genetics"},{"key":"75352_CR57","doi-asserted-by":"publisher","first-page":"2243","DOI":"10.1126\/scirobotics.adi2243","volume":"8","author":"T Wang","year":"2023","unstructured":"Wang, T. et al. Mechanical intelligence simplifies control in terrestrial limbless locomotion. Sci. Robot 8, 2243 (2023).","journal-title":"Sci. Robot"},{"key":"75352_CR58","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-021-26276-x","volume":"12","author":"X Zhang","year":"2021","unstructured":"Zhang, X., Naughton, N., Parthasarathy, T. & Gazzola, M. Friction modulation in limbless, three-dimensional gaits and heterogeneous terrains. Nat. Commun. 12, 1\u20138 (2021).","journal-title":"Nat. Commun."},{"key":"75352_CR59","doi-asserted-by":"crossref","unstructured":"Albantakis, L., Bernard, C., Brenner, N., Marder, E. & Narayanan, R. The brain\u2019s best kept secret is its degenerate structure. J. Neurosci. 44, e1339242024 (2024).","DOI":"10.1523\/JNEUROSCI.1339-24.2024"},{"key":"75352_CR60","doi-asserted-by":"crossref","unstructured":"Stiernagle, T. Maintenance of C. elegans. In WormBook: the Online Review of C. elegans Biology 1\u201311 (2006).","DOI":"10.1895\/wormbook.1.101.1"},{"key":"75352_CR61","doi-asserted-by":"crossref","unstructured":"Porta-de-la Riva, M., Fontrodona, L., Villanueva, A. & Cer\u00f3n, J. Basic Caenorhabditis elegans methods: synchronization and observation. J. Vis. Exp. 64, e4019 (2012).","DOI":"10.3791\/4019"},{"key":"75352_CR62","doi-asserted-by":"publisher","first-page":"3959","DOI":"10.1002\/j.1460-2075.1991.tb04966.x","volume":"10","author":"CC Mello","year":"1991","unstructured":"Mello, C. C., Kramer, J. M., Stinchcomb, D. & Ambros, V. Efficient gene transfer in C. elegans: extrachromosomal maintenance and integration of transforming sequences. EMBO J. 10, 3959\u20133970 (1991).","journal-title":"EMBO J."},{"key":"75352_CR63","doi-asserted-by":"crossref","unstructured":"Paix, A., Folkmann, A. & Seydoux, G. Precision genome editing using CRISPR-Cas9 and linear repair templates in C. elegans. Methods 121\u2013122, 86\u201393 (2017).","DOI":"10.1016\/j.ymeth.2017.03.023"},{"key":"75352_CR64","first-page":"1","volume":"13","author":"N Malaiwong","year":"2023","unstructured":"Malaiwong, N., Porta-de-la Riva, M. & Krieg, M. Linear repair templates Flint: single shot safe harbor transgene integration via fluorescent landmark interference. G3 (Bethesda, MD) 13, 1\u201312 (2023).","journal-title":"G3 (Bethesda, MD)"},{"key":"75352_CR65","doi-asserted-by":"crossref","unstructured":"Tse, J. R. & Engler, A. J. Linear repair templates: preparation of hydrogel substrates with tunable mechanical properties. Curr. Protoc. Cell Biol.Ch. 10 (2010).","DOI":"10.1002\/0471143030.cb1016s47"},{"key":"75352_CR66","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1038\/s41592-018-0112-1","volume":"15","author":"A Javer","year":"2018","unstructured":"Javer, A. et al. Linear repair templates: an open-source platform for analyzing and sharing worm-behavior data. Nat. Methods 15, 645\u2013646 (2018).","journal-title":"Nat. Methods"},{"key":"75352_CR67","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v021.i07","volume":"21","author":"T Duong","year":"2007","unstructured":"Duong, T. Linear repair templates Ks: Kernel density estimation and kernel discriminant analysis for multivariate data in r. J. Stat. Softw. 21, 1\u201316 (2007).","journal-title":"J. Stat. Softw."},{"key":"75352_CR68","doi-asserted-by":"crossref","unstructured":"Chacon, J., Duong, T. Monographs on Statistics and Applied Probability Vol. 160: Multivariate Kernel Smoothing and its Application (Taylor and Francis, 2018).","DOI":"10.1201\/9780429485572"},{"key":"75352_CR69","unstructured":"Sofroniew, N. et al. Napari: a multi-dimensional image viewer for Python https:\/\/zenodo.org\/records\/13863809 (2024)."},{"key":"75352_CR70","doi-asserted-by":"publisher","first-page":"e10","DOI":"10.14440\/jbm.2014.36","volume":"1","author":"AD Edelstein","year":"2014","unstructured":"Edelstein, A. D. et al. Advanced methods of microscope control using \u03bcmanager software. J. Biol. Methods 1, e10 (2014).","journal-title":"J. Biol. Methods"},{"key":"75352_CR71","doi-asserted-by":"publisher","first-page":"226","DOI":"10.1038\/s41592-021-01087-6","volume":"18","author":"H Pinkard","year":"2021","unstructured":"Pinkard, H. et al. Pycro-manager: open-source software for customized and reproducible microscope control. Nat. Methods 18, 226\u2013228 (2021).","journal-title":"Nat. Methods"},{"key":"75352_CR72","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1016\/j.ymeth.2016.09.016","volume":"115","author":"JY Tinevez","year":"2017","unstructured":"Tinevez, J. Y. et al. Trackmate: an open and extensible platform for single-particle tracking. Methods 115, 80\u201390 (2017).","journal-title":"Methods"},{"key":"75352_CR73","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fonc.2016.00150","volume":"6","author":"T Xu","year":"2016","unstructured":"Xu, T. et al. The expanding toolbox of in vivo bioluminescent imaging. Front. Oncol. 6, 1\u20138 (2016).","journal-title":"Front. Oncol."},{"key":"75352_CR74","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1038\/s41592-023-01836-9","volume":"20","author":"M Porta-de-la Riva","year":"2023","unstructured":"Porta-de-la Riva, M. et al. Neural engineering with photons as synaptic transmitters. Nat. Methods 20, 761\u2013769 (2023).","journal-title":"Nat. Methods"},{"key":"75352_CR75","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-022-34661-3","volume":"13","author":"H Setty","year":"2022","unstructured":"Setty, H. et al. Sexually dimorphic architecture and function of a mechanosensory circuit in C. elegans. Nat. Commun. 13, 1\u201336 (2022).","journal-title":"Nat. Commun."},{"key":"75352_CR76","first-page":"1","volume":"2018","author":"H Fehlauer","year":"2018","unstructured":"Fehlauer, H. et al. Using a microfluidics device for mechanical stimulation and high resolution imaging of C. elegans. J. Vis. Exp. 2018, 1\u201312 (2018).","journal-title":"J. Vis. Exp."},{"key":"75352_CR77","first-page":"51873","volume":"16","author":"SG Knoll","year":"2014","unstructured":"Knoll, S. G., Ali, M. Y. & Saif, M. T. A novel method for localizing reporter fluorescent beads near the cell culture surface for traction force microscopy. J. Vis. Exp. 16, 51873 (2014).","journal-title":"J. Vis. Exp."},{"key":"75352_CR78","doi-asserted-by":"crossref","unstructured":"Archit, A. et al. Segment anything for microscopy. Nat. Methods 22, 579\u2013591 (2025).","DOI":"10.1038\/s41592-024-02580-4"},{"key":"75352_CR79","doi-asserted-by":"publisher","unstructured":"Liberzon, A., K\u00e4ufer, T., Bauer, A., Vennemann, P. & Zimmer, E. Openpiv\/openpiv-python: Openpiv-python v0.23.6. Zenodo, https:\/\/doi.org\/10.5281\/zenodo.593157 (2021).","DOI":"10.5281\/zenodo.593157"},{"key":"75352_CR80","doi-asserted-by":"publisher","first-page":"e1008364","DOI":"10.1371\/journal.pcbi.1008364","volume":"17","author":"A Bauer","year":"2021","unstructured":"Bauer, A. et al. pytfm: a tool for traction force and monolayer stress microscopy. PLoS Comput. Biol. 17, e1008364 (2021).","journal-title":"PLoS Comput. Biol."},{"key":"75352_CR81","doi-asserted-by":"publisher","first-page":"C595","DOI":"10.1152\/ajpcell.00270.2001","volume":"282","author":"JP Butler","year":"2002","unstructured":"Butler, J. P., Tolic-N\u00f8rrelykke, I. M., Fabry, B. & Fredberg, J. J. Traction fields, moments, and strain energy that cells exert on their surroundings. Am. J. Physiol. Cell Physiol. 282, C595\u2013C605 (2002).","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"75352_CR82","doi-asserted-by":"crossref","unstructured":"Hox, J., Moerbeek, M. & Van de Schoot, R. Multilevel Analysis Techniques and Applications 3rd edn (Taylor and Francis, 2017).","DOI":"10.4324\/9781315650982"},{"key":"75352_CR83","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.physrep.2018.06.001","volume":"748","author":"G Lancaster","year":"2018","unstructured":"Lancaster, G., Iatsenko, D., Pidde, A., Ticcinelli, V. & Stefanovska, A. Surrogate data for hypothesis testing of physical systems. Phys. Rep. 748, 1\u201360 (2018).","journal-title":"Phys. Rep."},{"key":"75352_CR84","doi-asserted-by":"crossref","unstructured":"Schreiber, T. & Schmitz, A. Surrogate time series. Physica D. 142, 346\u2013382 (2000).","DOI":"10.1016\/S0167-2789(00)00043-9"},{"key":"75352_CR85","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1038\/s41592-019-0470-3","volume":"16","author":"J Ho","year":"2019","unstructured":"Ho, J., Tumkaya, T., Aryal, S., Choi, H. & Claridge-Chang, A. Moving beyond p values: data analysis with estimation graphics. Nat. Methods 16, 565\u2013566 (2019).","journal-title":"Nat. Methods"},{"key":"75352_CR86","doi-asserted-by":"publisher","first-page":"58859","DOI":"10.1038\/349588a0","volume":"349","author":"M Driscol","year":"1991","unstructured":"Driscol, M. & Chalfie, M. The mec-4 gene is a member of a family of Caenorhabditis elegans genes that can mutate to induce neuronal degeneration. Nature 349, 58859\u2013593 (1991).","journal-title":"Nature"},{"key":"75352_CR87","doi-asserted-by":"crossref","unstructured":"Yokobayashi, Y. Aptamer-based and aptazyme-based riboswitches in mammalian cells. Curr. Opin. Chem. Biol. 52, 72\u201378 (2019).","DOI":"10.1016\/j.cbpa.2019.05.018"}],"container-title":["Nature Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-75352-7","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-75352-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-75352-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T11:03:24Z","timestamp":1787828604000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41467-026-75352-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,25]]},"references-count":87,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["75352"],"URL":"https:\/\/doi.org\/10.1038\/s41467-026-75352-7","relation":{},"ISSN":["2041-1723"],"issn-type":[{"value":"2041-1723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,25]]},"assertion":[{"value":"11 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 June 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}],"article-number":"9083"}}