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Naik, “Data associated with Keratins coordinate tissue spreading .” Institute of Science and Technology Austria, 2026.","chicago":"Naik, Suyash. “Data Associated with Keratins Coordinate Tissue Spreading .” Institute of Science and Technology Austria, 2026. <a href=\"https://doi.org/10.15479/AT-ISTA-21137\">https://doi.org/10.15479/AT-ISTA-21137</a>."},"oa_version":"Published Version","day":"24","OA_place":"repository","oa":1,"date_created":"2026-02-04T16:38:02Z","contributor":[{"contributor_type":"researcher","first_name":"Yann-Edwin","last_name":"Keta"},{"last_name":"Henkes","first_name":"Silke ","contributor_type":"supervisor"},{"id":"39427864-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0002-0912-4566","first_name":"Carl-Philipp J","contributor_type":"supervisor","last_name":"Heisenberg"},{"contributor_type":"supervisor","first_name":"Edouard B","last_name":"Hannezo","orcid":"0000-0001-6005-1561","id":"3A9DB764-F248-11E8-B48F-1D18A9856A87"}],"year":"2026","department":[{"_id":"GradSch"},{"_id":"CaHe"},{"_id":"EdHa"}],"date_updated":"2026-06-10T09:44:10Z","acknowledgement":"We thank all members of the Heisenberg, Henkes, and Hannezo groups for their support. We are also grateful to the Imaging and Optics, Scientific Computing, Life Science Support, and Cryo-Electron Microscopy facilities at ISTA for their technical assistance and support. Numerical simulations were performed using the computational resources from Lorentz Institute and the Academic Leiden Interdisciplinary Cluster Environment (ALICE) provided by Leiden University, and from PMMH provided by Sorbonne Université. S.N has received funding from European Union’s Horizon 2020 research and innovation programme (grant agreement No. 665385). This work was supported by the Austrian Science Fund (FWF) under projects PAT5044023 and W1250 awarded to C.-P.H.","_id":"21137"},{"file":[{"creator":"nhino","date_created":"2026-07-13T09:16:20Z","content_type":"application/pdf","success":1,"relation":"main_file","checksum":"66444afd243dce7d383d52d44e8d34a4","access_level":"open_access","file_id":"22283","file_name":"Main_text_and_figures.pdf","date_updated":"2026-07-13T09:16:20Z","file_size":12477675},{"creator":"nhino","date_created":"2026-07-13T09:16:25Z","checksum":"90bceb34de64ec792c5de117f0890d05","content_type":"application/pdf","relation":"main_file","success":1,"file_name":"Supplementary_figures.pdf","file_size":4545901,"date_updated":"2026-07-13T09:16:25Z","access_level":"open_access","file_id":"22284"},{"relation":"main_file","success":1,"content_type":"video/mp4","checksum":"9d9ab89c372142f2ffb6c8c625334d7f","file_id":"22285","access_level":"open_access","date_updated":"2026-07-13T09:16:28Z","file_size":10349451,"file_name":"Supplementary_Video1.mp4","creator":"nhino","date_created":"2026-07-13T09:16:28Z"}],"supplementarymaterial":"yes","file_date_updated":"2026-07-13T09:16:28Z","has_accepted_license":"1","month":"07","project":[{"_id":"8f060199-16d5-11f0-9cad-f3253b266c46","grant_number":"PAT 5044023","name":"Keratins in epithelial tissue spreading"},{"name":"Mechanosensitive signaling activation in the crosstalk between mechanical force and tissuefluidity","_id":"34dd7f3b-11ca-11ed-8bc3-856f2c87f5da","grant_number":"LTF 16-2022"}],"user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","corr_author":"1","type":"preprint","author":[{"last_name":"Hino","full_name":"Hino, Naoya","first_name":"Naoya","id":"5299a9ce-7679-11eb-a7bc-d1e62b936307"},{"id":"e3b3eda7-fd4d-11eb-8fd8-c40af7a478b1","first_name":"Tushna","full_name":"Kapoor, Tushna","last_name":"Kapoor"},{"last_name":"Gubbala","full_name":"Gubbala, Uday R","first_name":"Uday R","id":"bb4a0dc4-32c9-11ee-b5ce-a97ceedd5924"},{"orcid":"0000-0001-6005-1561","id":"3A9DB764-F248-11E8-B48F-1D18A9856A87","last_name":"Hannezo","full_name":"Hannezo, Edouard B","first_name":"Edouard B"},{"orcid":"0000-0002-0912-4566","id":"39427864-F248-11E8-B48F-1D18A9856A87","last_name":"Heisenberg","full_name":"Heisenberg, Carl-Philipp J","first_name":"Carl-Philipp J"}],"publisher":"Institute of Science and Technology Austria","date_published":"2026-07-14T00:00:00Z","title":"Apical domain mechanosensation regulates tissue tension homeostasis","language":[{"iso":"eng"}],"publication_status":"draft","tmp":{"image":"/images/cc_by.png","legal_code_url":"https://creativecommons.org/licenses/by/4.0/legalcode","short":"CC BY (4.0)","name":"Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)"},"dataavailabilitystatement":"The MATLAB code for image analysis, and the full model code, including all parameter values\r\nand condition-specific settings, are available on GitHub at https://github.com/uday2607/EVL-tension-homeostasis.git.","ddc":["570"],"acknowledged_ssus":[{"_id":"Bio"},{"_id":"LifeSc"},{"_id":"EM-Fac"}],"status":"public","oa_version":"Preprint","day":"14","OA_place":"publisher","das_tickbox":"1","keyword":["Epithelial spreading","tissue tension","mechanosensation","aPKC","Kibra","zebrafish"],"oa":1,"researchdata_availability":"yes","OA_type":"green","article_processing_charge":"No","citation":{"short":"N. Hino, T. Kapoor, U.R. Gubbala, E.B. Hannezo, C.-P.J. Heisenberg, (n.d.).","apa":"Hino, N., Kapoor, T., Gubbala, U. R., Hannezo, E. B., &#38; Heisenberg, C.-P. J. (n.d.). Apical domain mechanosensation regulates tissue tension homeostasis. Institute of Science and Technology Austria.","chicago":"Hino, Naoya, Tushna Kapoor, Uday R Gubbala, Edouard B Hannezo, and Carl-Philipp J Heisenberg. “Apical Domain Mechanosensation Regulates Tissue Tension Homeostasis.” Institute of Science and Technology Austria, n.d.","ieee":"N. Hino, T. Kapoor, U. R. Gubbala, E. B. Hannezo, and C.-P. J. Heisenberg, “Apical domain mechanosensation regulates tissue tension homeostasis.” Institute of Science and Technology Austria.","mla":"Hino, Naoya, et al. <i>Apical Domain Mechanosensation Regulates Tissue Tension Homeostasis</i>. Institute of Science and Technology Austria.","ama":"Hino N, Kapoor T, Gubbala UR, Hannezo EB, Heisenberg C-PJ. Apical domain mechanosensation regulates tissue tension homeostasis.","ista":"Hino N, Kapoor T, Gubbala UR, Hannezo EB, Heisenberg C-PJ. Apical domain mechanosensation regulates tissue tension homeostasis."},"related_material":{"record":[{"relation":"earlier_version","status":"public","id":"21864"}]},"year":"2026","date_updated":"2026-07-14T07:07:41Z","department":[{"_id":"CaHe"},{"_id":"EdHa"},{"_id":"GradSch"}],"acknowledgement":"We thank all members of the Heisenberg group for discussion and feedback on the manuscript, and the Imaging and Optics Facility, the Life Science Support Facility and the Electron Microscopy Facility of the Institute of Science and Technology Austria (ISTA) for their continued support. We are grateful to M. Sonawane (Tata Institute of Fundamental Research, India) for providing the pCS2-HA-aPKC (PKCι)-V260F (DN) and pCS2-HA-aPKC (PKCι)-A122E (CA) plasmids, and to I. Mayer for the discussion. Molecular graphics and analyses were performed with UCSF ChimeraX, developed by the Resource for Biocomputing, Visualization, and Informatics at the University of California, San Francisco, with support from National Institutes of Health R01-GM129325 and the Office of Cyber Infrastructure and Computational Biology, National Institute of Allergy and Infectious Diseases. This research was funded in whole or in part by the Austrian Science Fund (FWF; grant no. PAT5044023) to C.-P.H., and by a JSPS Overseas Research Fellowship and an EMBO Postdoctoral Fellowship (ALTF 16-2022) to N.H.","abstract":[{"lang":"eng","text":"Tissue tension is a key determinant of tissue shape, and its regulation is essential for both morphogenesis and the maintenance of tissue integrity. During zebrafish embryogenesis, the enveloping layer (EVL) – an epithelial monolayer covering the blastoderm – undergoes extensive spreading that is driven by pulling forces exerted at its margin and more than doubles its surface area. Yet whether and how the EVL actively regulates its tissue tension during this process remains unclear. Here, we show that the EVL maintains constant tissue tension while spreading, and that it achieves this by reducing apical cell contractility in response to the same pulling forces that drive its spreading. We identify a mechanosensitive pathway underlying this response, mediated by the scaffold/adaptor protein Kibra regulating the activity of atypical protein kinase C (aPKC) at the apical domain of EVL cells. Under low mechanical stretch, Kibra forms condensates at the base of actin-based apical projections, where it activates Myosin II to increase apical contractility through aPKC downregulation. As mechanical stretch increases, apical projections disassemble, Kibra condensates dissolve, and aPKC activity rises. Elevated aPKC activity in turn reduces apical contractility by reducing Myosin II activity, thereby maintaining constant tissue tension despite increased mechanical stretch. Together, these findings reveal a mechanosensitive mechanism that enables robust adaptation of tissue tension to changing mechanical stretch, ensuring efficient tissue spreading and morphogenesis."}],"_id":"22276","date_created":"2026-07-13T09:03:26Z"},{"supervisor":[{"last_name":"Heisenberg","full_name":"Heisenberg, Carl-Philipp J","first_name":"Carl-Philipp J","orcid":"0000-0002-0912-4566","id":"39427864-F248-11E8-B48F-1D18A9856A87"},{"last_name":"Hannezo","first_name":"Edouard B","full_name":"Hannezo, Edouard B","id":"3A9DB764-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0001-6005-1561"}],"alternative_title":["ISTA Thesis"],"date_created":"2025-10-10T14:58:30Z","_id":"20441","acknowledgement":"I would also like to thank the LSF and Cryo facility at ISTA, which have been helpful in my\r\nexperiments. I would also like to acknowledge FWF, grant DOI 10.55776/PAT5044023 and JKU Nanocell grant DOI \r\n10.55776/W1250 for providing funding for my PhD research. EMBO and FWF for providing funding for travel grants to attend conferences.","abstract":[{"text":"Epithelial spreading plays a pivotal role in the development of organisms especially those\r\nsuch as zebrafish which require the epithelial enveloping layer (EVL) to spread to cover the\r\nsubstantial yolk surface during gastrulation. Epiboly requires the transition of the epithelium\r\nwith cuboidal cells to form a thin, flat squamous epithelial sheet. During this transition, the\r\ncells show tissue-scale mechanosensation with mechanisms such as direct mechanical control\r\nover the axis of cell division.\r\nCytoskeletal intermediate filaments play a crucial role in vertebrate cells, not only facilitating\r\nmechanical stability but also helping facilitate the mechanosensitive response of the cell.\r\nMechanosenstivity displayed by intermediate filaments is due not just to their interesting\r\nphysical properties but also to their interactions with other cytoskeletal elements such as actin\r\nand microtubules. Keratin is the predominant intermediate filament expressed in the EVL.\r\nIt expresses concomitantly with the gastrulation movements of the developing embryo. Our\r\nwork focuses on understanding the role and dynamics of the keratin cytoskeletal network in\r\nmodulating the physical aspects of EVL spreading. We demonstrated with the combination of\r\nphysical characterisation and manipulations of the EVL, utilising a variety of biophysical tools\r\nand microscopy, the mechanistic role of keratin in tissue spreading.\r\nGenerating novel genetic morphants and mutants, we probe the effect that the loss of the\r\nkeratin network has on the physiology of the epithelium and the developing embryo. We\r\nshow that the changing organisation of the keratin network is important for changing EVL\r\nphysical properties as the stress imposed on the EVL increases during epiboly. By modelling\r\nthe epithelium, we study how the mechanical heterogeneity in an epithelium can feed back into\r\na mechanical loop to the maturation of the keratin network and hence affect the mechanics\r\nof the epithelium. However, unlike what would be predicted by the effect of intermediate\r\nfilaments in acting as a security belt and increasing the resistance of the epithelium, we observe\r\nthat loss of keratin leads to a delay in the EVL movement. Using both local aspirations of the\r\nYSL and EVL ablations, we demonstrate the mechanistic facilitation of actin mechanosensation\r\nin a keratin-dependent manner.\r\nFurthermore, using chemical inhibitors of microtubule polymerisation, we provide insight into\r\nthe mechanisms underlying the organisation and distribution of keratin. Interestingly, the\r\nphenotype observed upon this loss of microtubules shows that keratins interact with the nucleus\r\nthrough microtubular interactions. Together with these diverse observations, we describe\r\nthe mechanosensory feedback between resilience and that is critical for uniform and robust\r\nspreading of the epithelium.","lang":"eng"}],"date_updated":"2026-07-06T12:51:41Z","department":[{"_id":"GradSch"},{"_id":"CaHe"},{"_id":"EdHa"}],"year":"2025","related_material":{"record":[{"status":"public","id":"20465","relation":"part_of_dissertation"}]},"citation":{"mla":"Naik, Suyash. <i>Keratins Act as Global Coordinators of Tissue Spreading through Mechanosensitive Feedback</i>. Institute of Science and Technology Austria, 2025, doi:<a href=\"https://doi.org/10.15479/AT-ISTA-20441\">10.15479/AT-ISTA-20441</a>.","ama":"Naik S. Keratins act as global coordinators of tissue spreading through mechanosensitive feedback. 2025. doi:<a href=\"https://doi.org/10.15479/AT-ISTA-20441\">10.15479/AT-ISTA-20441</a>","ista":"Naik S. 2025. Keratins act as global coordinators of tissue spreading through mechanosensitive feedback. Institute of Science and Technology Austria.","short":"S. Naik, Keratins Act as Global Coordinators of Tissue Spreading through Mechanosensitive Feedback, Institute of Science and Technology Austria, 2025.","apa":"Naik, S. (2025). <i>Keratins act as global coordinators of tissue spreading through mechanosensitive feedback</i>. Institute of Science and Technology Austria. <a href=\"https://doi.org/10.15479/AT-ISTA-20441\">https://doi.org/10.15479/AT-ISTA-20441</a>","chicago":"Naik, Suyash. “Keratins Act as Global Coordinators of Tissue Spreading through Mechanosensitive Feedback.” Institute of Science and Technology Austria, 2025. <a href=\"https://doi.org/10.15479/AT-ISTA-20441\">https://doi.org/10.15479/AT-ISTA-20441</a>.","ieee":"S. Naik, “Keratins act as global coordinators of tissue spreading through mechanosensitive feedback,” Institute of Science and Technology Austria, 2025."},"article_processing_charge":"No","publication_identifier":{"isbn":["978-3-99078-069-5"],"issn":["2663-337X"]},"oa":1,"OA_place":"publisher","day":"12","oa_version":"Published Version","acknowledged_ssus":[{"_id":"Bio"},{"_id":"LifeSc"}],"status":"public","page":"105","ddc":["596","597","532"],"tmp":{"image":"/images/cc_by_sa.png","legal_code_url":"https://creativecommons.org/licenses/by-sa/4.0/legalcode","name":"Creative Commons Attribution-ShareAlike 4.0 International Public License (CC BY-SA 4.0)","short":"CC BY-SA (4.0)"},"publication_status":"published","language":[{"iso":"eng"}],"title":"Keratins act as global coordinators of tissue spreading through mechanosensitive feedback","publisher":"Institute of Science and Technology Austria","date_published":"2025-10-12T00:00:00Z","author":[{"id":"2C0B105C-F248-11E8-B48F-1D18A9856A87","orcid":"0000-0001-8421-5508","last_name":"Naik","first_name":"Suyash","full_name":"Naik, Suyash"}],"type":"dissertation","degree_awarded":"PhD","user_id":"ba8df636-2132-11f1-aed0-ed93e2281fdd","corr_author":"1","project":[{"grant_number":"PAT 5044023","name":"Keratins in epithelial tissue spreading","_id":"8f060199-16d5-11f0-9cad-f3253b266c46"},{"grant_number":"W 1250-B20","name":"Nano-Analytics of Cellular Systems","_id":"25AA5F24-B435-11E9-9278-68D0E5697425","call_identifier":"FWF"}],"month":"10","has_accepted_license":"1","file_date_updated":"2025-10-28T13:10:26Z","doi":"10.15479/AT-ISTA-20441","file":[{"creator":"snaik","date_created":"2025-10-28T13:10:08Z","relation":"main_file","success":1,"content_type":"application/pdf","checksum":"2892f04d4a5c18677871c3e06ac1244a","file_id":"20567","access_level":"open_access","date_updated":"2025-10-28T13:10:08Z","file_size":6846189,"file_name":"Thesis_PDFA_.pdf"},{"creator":"snaik","date_created":"2025-10-28T13:10:26Z","checksum":"15934d4465cd0e9b7c32678da9a33a2f","content_type":"application/zip","relation":"source_file","file_name":"Thesis.zip","date_updated":"2025-10-28T13:10:26Z","file_size":8839300,"access_level":"open_access","file_id":"20568"}]}]