{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T21:20:18Z","timestamp":1792790418292,"version":"4.5.0"},"posted":{"date-parts":[[2026,10,3]]},"group-title":"Biophysics","reference-count":86,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,10,3]],"date-time":"2026-10-03T00:00:00Z","timestamp":1790985600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Science Foundation grant NSF Directorate for Engineering","award":["CMMI-2449148"],"award-info":[{"award-number":["CMMI-2449148"]}],"id":[{"id":"https:\/\/ror.org\/00b6sbb32","id-type":"ROR","asserted-by":"crossref"}]},{"name":"NIH National Institute of Arthritis and Musculoskeletal and Skin Disease","award":["5R21AR083064-02"],"award-info":[{"award-number":["5R21AR083064-02"]}],"id":[{"id":"https:\/\/ror.org\/006zn3t30","id-type":"ROR","asserted-by":"crossref"}]},{"name":"NSF Graduate Research Fellowship","award":["DGE-2139899"],"award-info":[{"award-number":["DGE-2139899"]}]},{"name":"National Science Foundation","award":["DMR-2327094"],"award-info":[{"award-number":["DMR-2327094"]}]},{"name":"National Science Foundation","award":["DMR-1719875"],"award-info":[{"award-number":["DMR-1719875"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,10,8]]},"abstract":"<jats:p>Large shear deformation in cartilage can lead to tissue failure and injury, which are often precursors to diseases such as post-traumatic osteoarthritis. Under shear, cartilage strain-stiffens prior to failure, which makes stiffening an important indicator of changes to the loading of the extracellular matrix that gives rise to this tissue\u2019s properties. While a variety of stiffening mechanisms ranging from uncrimping to fiber-alignment have been explored, whether such mechanisms apply to the depth dependent shear strain stiffening in articular cartilage remains poorly understood. Here, we present experiments and theory that support the idea that the strain-stiffening behavior is caused by a mechanical phase transition from a random isotropic network whose mechanics arise primarily from bending-dominated modes to one where the mechanics arise from stretching-dominated modes. Indeed, consistent with network model predictions, we find that stress-strain data for all depths can be collapsed onto a single universal curve that diverges at a depth-dependent critical strain. These results show that increasing the local concentration of collagen, which constitutes the primary load-bearing network in cartilage, decreases the critical strain at which the strain-stiffening transition occurs, while increasing the low-strain elastic modulus. Furthermore, we demonstrate that bulk compression leads to a higher critical strain and decreased low-strain elastic modulus. Network simulations capture this rich behavior, and indicate that under compression the network collapses, which allows for larger strains before the transition from bending-dominated to stretching-dominated mechanics occurs. Collectively, these findings shed light on the roles that tissue composition and axial strain play in the mechanisms that organize how articular cartilage stiffens during the lead-up to injury.<\/jats:p>\n                <jats:sec>\n                  <jats:title>Significance Statement<\/jats:title>\n                  <jats:p>We show that strain stiffening in cartilage, which is a precursor to tissue damage, is governed by a mechanical phase transition from a network whose properties arise from bending-dominated modes to one where the mechanics arise from stretching-dominated modes. This framework provides insights into the possible failure mechanisms that arise under extreme loading and points to new approaches for prevention and repair.<\/jats:p>\n                <\/jats:sec>","DOI":"10.64898\/2026.09.28.751808","type":"posted-content","created":{"date-parts":[[2026,10,4]],"date-time":"2026-10-04T04:10:12Z","timestamp":1791087012000},"source":"Crossref","is-referenced-by-count":0,"title":["Shear Strain Stiffening in Cartilage is Governed by a Mechanical Phase Transition"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9918-2614","authenticated-orcid":false,"given":"Japheth","family":"Omonira","sequence":"first","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5294-0251","authenticated-orcid":false,"given":"Joshua D.","family":"Clugston","sequence":"additional","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0361-007X","authenticated-orcid":false,"given":"Anna R.","family":"Barth","sequence":"additional","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9325-2586","authenticated-orcid":false,"given":"Stephen J.","family":"Thornton","sequence":"additional","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9126-0892","authenticated-orcid":false,"given":"James P.","family":"Sethna","sequence":"additional","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0397-8886","authenticated-orcid":false,"given":"Moumita","family":"Das","sequence":"additional","affiliation":[{"name":"School of Physics and Astronomy, Rochester Institute of Technology, Rochester, New York 14623, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1094-6433","authenticated-orcid":false,"given":"Lawrence J.","family":"Bonassar","sequence":"additional","affiliation":[{"name":"Meinig School of Biomedical Engineering, Cornell University, Ithaca, New York 14853, USA"},{"name":"Sibley School of Mechanical and Aerospace Engineering, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9218-043X","authenticated-orcid":false,"given":"Itai","family":"Cohen","sequence":"additional","affiliation":[{"name":"Department of Physics, Cornell University, Ithaca, New York 14853, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026101210550538000_2026.09.28.751808v2.1","doi-asserted-by":"publisher","DOI":"10.1016\/0043-1648(62)90176-X"},{"key":"2026101210550538000_2026.09.28.751808v2.2","doi-asserted-by":"crossref","first-page":"2005513","DOI":"10.1002\/adma.202005513","article-title":"Recent Progress in Cartilage Lubrication","volume":"33","year":"2021","journal-title":"Adv. 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