---
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acknowledged_ssus:
- _id: Bio
- _id: EM-Fac
- _id: ScienComp
- _id: LifeSc
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.
article_processing_charge: No
author:
- first_name: Suyash
  full_name: Naik, Suyash
  id: 2C0B105C-F248-11E8-B48F-1D18A9856A87
  last_name: Naik
  orcid: 0000-0001-8421-5508
citation:
  ama: Naik S. Data associated with Keratins coordinate tissue spreading . 2026. doi:<a
    href="https://doi.org/10.15479/AT-ISTA-21137">10.15479/AT-ISTA-21137</a>
  apa: Naik, S. (2026). Data associated with Keratins coordinate tissue spreading
    . Institute of Science and Technology Austria. <a href="https://doi.org/10.15479/AT-ISTA-21137">https://doi.org/10.15479/AT-ISTA-21137</a>
  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>.
  ieee: S. Naik, “Data associated with Keratins coordinate tissue spreading .” Institute
    of Science and Technology Austria, 2026.
  ista: Naik S. 2026. Data associated with Keratins coordinate tissue spreading ,
    Institute of Science and Technology Austria, <a href="https://doi.org/10.15479/AT-ISTA-21137">10.15479/AT-ISTA-21137</a>.
  mla: Naik, Suyash. <i>Data Associated with Keratins Coordinate Tissue Spreading
    </i>. Institute of Science and Technology Austria, 2026, doi:<a href="https://doi.org/10.15479/AT-ISTA-21137">10.15479/AT-ISTA-21137</a>.
  short: S. Naik, (2026).
contributor:
- contributor_type: researcher
  first_name: Yann-Edwin
  last_name: Keta
- contributor_type: supervisor
  first_name: 'Silke '
  last_name: Henkes
- contributor_type: supervisor
  first_name: Carl-Philipp J
  id: 39427864-F248-11E8-B48F-1D18A9856A87
  last_name: Heisenberg
  orcid: 0000-0002-0912-4566
- contributor_type: supervisor
  first_name: Edouard B
  id: 3A9DB764-F248-11E8-B48F-1D18A9856A87
  last_name: Hannezo
  orcid: 0000-0001-6005-1561
corr_author: '1'
date_created: 2026-02-04T16:38:02Z
date_published: 2026-03-24T00:00:00Z
date_updated: 2026-06-10T09:44:10Z
day: '24'
department:
- _id: GradSch
- _id: CaHe
- _id: EdHa
doi: 10.15479/AT-ISTA-21137
ec_funded: 1
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  date_created: 2026-03-16T11:51:10Z
  date_updated: 2026-03-16T11:51:10Z
  description: 'Python3 library written in C++20 to integrate vertex models. Please
    read the readme at https://github.com/yketa/cells/blob/main/README.md for detailed
    instructions for installation and usage of the code in this repository. '
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has_accepted_license: '1'
license: https://creativecommons.org/licenses/by-sa/4.0/
month: '3'
oa: 1
oa_version: Published Version
project:
- _id: 2564DBCA-B435-11E9-9278-68D0E5697425
  call_identifier: H2020
  grant_number: '665385'
  name: International IST Doctoral Program
- _id: 8f060199-16d5-11f0-9cad-f3253b266c46
  grant_number: PAT 5044023
  name: Keratins in epithelial tissue spreading
- _id: 252C3B08-B435-11E9-9278-68D0E5697425
  call_identifier: FWF
  grant_number: W1250-B20
  name: Nano-Analytics of Cellular Systems
publisher: Institute of Science and Technology Austria
status: public
title: 'Data associated with Keratins coordinate tissue spreading '
tmp:
  image: /images/cc_by_sa.png
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    BY-SA 4.0)
  short: CC BY-SA (4.0)
type: research_data
user_id: 68b8ca59-c5b3-11ee-8790-cd641c68093d
year: '2026'
...
---
OA_place: publisher
OA_type: green
_id: '22276'
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.
acknowledged_ssus:
- _id: Bio
- _id: LifeSc
- _id: EM-Fac
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.
article_processing_charge: No
author:
- first_name: Naoya
  full_name: Hino, Naoya
  id: 5299a9ce-7679-11eb-a7bc-d1e62b936307
  last_name: Hino
- first_name: Tushna
  full_name: Kapoor, Tushna
  id: e3b3eda7-fd4d-11eb-8fd8-c40af7a478b1
  last_name: Kapoor
- first_name: Uday R
  full_name: Gubbala, Uday R
  id: bb4a0dc4-32c9-11ee-b5ce-a97ceedd5924
  last_name: Gubbala
- first_name: Edouard B
  full_name: Hannezo, Edouard B
  id: 3A9DB764-F248-11E8-B48F-1D18A9856A87
  last_name: Hannezo
  orcid: 0000-0001-6005-1561
- first_name: Carl-Philipp J
  full_name: Heisenberg, Carl-Philipp J
  id: 39427864-F248-11E8-B48F-1D18A9856A87
  last_name: Heisenberg
  orcid: 0000-0002-0912-4566
citation:
  ama: Hino N, Kapoor T, Gubbala UR, Hannezo EB, Heisenberg C-PJ. Apical domain mechanosensation
    regulates tissue tension homeostasis.
  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.
  ista: Hino N, Kapoor T, Gubbala UR, Hannezo EB, Heisenberg C-PJ. Apical domain mechanosensation
    regulates tissue tension homeostasis.
  mla: Hino, Naoya, et al. <i>Apical Domain Mechanosensation Regulates Tissue Tension
    Homeostasis</i>. Institute of Science and Technology Austria.
  short: N. Hino, T. Kapoor, U.R. Gubbala, E.B. Hannezo, C.-P.J. Heisenberg, (n.d.).
corr_author: '1'
das_tickbox: '1'
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."
date_created: 2026-07-13T09:03:26Z
date_published: 2026-07-14T00:00:00Z
date_updated: 2026-07-14T07:07:41Z
day: '14'
ddc:
- '570'
department:
- _id: CaHe
- _id: EdHa
- _id: GradSch
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  relation: main_file
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file_date_updated: 2026-07-13T09:16:28Z
has_accepted_license: '1'
keyword:
- Epithelial spreading
- tissue tension
- mechanosensation
- aPKC
- Kibra
- zebrafish
language:
- iso: eng
month: '07'
oa: 1
oa_version: Preprint
project:
- _id: 8f060199-16d5-11f0-9cad-f3253b266c46
  grant_number: PAT 5044023
  name: Keratins in epithelial tissue spreading
- _id: 34dd7f3b-11ca-11ed-8bc3-856f2c87f5da
  grant_number: LTF 16-2022
  name: Mechanosensitive signaling activation in the crosstalk between mechanical
    force and tissuefluidity
publication_status: draft
publisher: Institute of Science and Technology Austria
related_material:
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    status: public
researchdata_availability: yes
status: public
supplementarymaterial: yes
title: Apical domain mechanosensation regulates tissue tension homeostasis
tmp:
  image: /images/cc_by.png
  legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
  name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
  short: CC BY (4.0)
type: preprint
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
year: '2026'
...
---
OA_place: publisher
_id: '20441'
abstract:
- lang: eng
  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."
acknowledged_ssus:
- _id: Bio
- _id: LifeSc
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."
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Suyash
  full_name: Naik, Suyash
  id: 2C0B105C-F248-11E8-B48F-1D18A9856A87
  last_name: Naik
  orcid: 0000-0001-8421-5508
citation:
  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>
  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.
  ista: Naik S. 2025. Keratins act as global coordinators of tissue spreading through
    mechanosensitive feedback. Institute of Science and Technology Austria.
  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>.
  short: S. Naik, Keratins Act as Global Coordinators of Tissue Spreading through
    Mechanosensitive Feedback, Institute of Science and Technology Austria, 2025.
corr_author: '1'
date_created: 2025-10-10T14:58:30Z
date_published: 2025-10-12T00:00:00Z
date_updated: 2026-07-06T12:51:41Z
day: '12'
ddc:
- '596'
- '597'
- '532'
degree_awarded: PhD
department:
- _id: GradSch
- _id: CaHe
- _id: EdHa
doi: 10.15479/AT-ISTA-20441
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has_accepted_license: '1'
language:
- iso: eng
month: '10'
oa: 1
oa_version: Published Version
page: '105'
project:
- _id: 8f060199-16d5-11f0-9cad-f3253b266c46
  grant_number: PAT 5044023
  name: Keratins in epithelial tissue spreading
- _id: 25AA5F24-B435-11E9-9278-68D0E5697425
  call_identifier: FWF
  grant_number: W 1250-B20
  name: Nano-Analytics of Cellular Systems
publication_identifier:
  isbn:
  - 978-3-99078-069-5
  issn:
  - 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
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  - id: '20465'
    relation: part_of_dissertation
    status: public
status: public
supervisor:
- first_name: Carl-Philipp J
  full_name: Heisenberg, Carl-Philipp J
  id: 39427864-F248-11E8-B48F-1D18A9856A87
  last_name: Heisenberg
  orcid: 0000-0002-0912-4566
- first_name: Edouard B
  full_name: Hannezo, Edouard B
  id: 3A9DB764-F248-11E8-B48F-1D18A9856A87
  last_name: Hannezo
  orcid: 0000-0001-6005-1561
title: Keratins act as global coordinators of tissue spreading through mechanosensitive
  feedback
tmp:
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  name: Creative Commons Attribution-ShareAlike 4.0 International Public License (CC
    BY-SA 4.0)
  short: CC BY-SA (4.0)
type: dissertation
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
year: '2025'
...