---
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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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file_date_updated: 2026-03-24T07:21:43Z
has_accepted_license: '1'
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:
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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
_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: 2026-04-07T11:58:57Z
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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  date_created: 2025-10-28T13:10:08Z
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file_date_updated: 2025-10-28T13:10:26Z
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:
  record:
  - 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:
  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)
type: dissertation
user_id: ba8df636-2132-11f1-aed0-ed93e2281fdd
year: '2025'
...
---
OA_place: repository
_id: '20465'
abstract:
- lang: eng
  text: For tissues to spread, they must be deformable while maintaining their structural
    integrity. How these opposing requirements are balanced within spreading tissues
    is not yet well understood. Here, we show that keratin intermediate filaments
    function in epithelial spreading by adapting tissue mechanical resilience to the
    stresses arising in the tissue during the spreading process. By analysing the
    expansion of the enveloping cell layer (EVL) over the large yolk cell in early
    zebrafish embryos in vivo, we found that keratin network maturation in EVL cells
    is promoted by stresses building up within the spreading tissue. Through genetic
    interference and tissue rheology experiments, complemented by a vertex model with
    mechanochemical feedback, we demonstrate that stress-induced keratin network maturation
    in the EVL increases tissue viscosity, which is essential for preventing tissue
    rupture. Interestingly, keratins are also required in the yolk cell for mechanosensitive
    actomyosin network contraction and flow, the force-generating processes pulling
    the EVL. These dual mechanosensitive functions of keratins enable a balance between
    pulling force production in the yolk cell and the mechanical resilience of the
    EVL against stresses generated by these pulling forces, thereby ensuring uniform
    and robust tissue spreading.
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
- first_name: Yann-Edwin
  full_name: Keta, Yann-Edwin
  last_name: Keta
- first_name: Kornelija
  full_name: Pranjic-Ferscha, Kornelija
  id: 4362B3C2-F248-11E8-B48F-1D18A9856A87
  last_name: Pranjic-Ferscha
- 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: Silke
  full_name: Henkes, Silke
  last_name: Henkes
- 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: Naik S, Keta Y-E, Pranjic-Ferscha K, Hannezo EB, Henkes S, Heisenberg C-PJ.
    Keratins coordinate tissue spreading by balancing spreading forces with tissue
    material properties. <i>bioRxiv</i>. doi:<a href="https://doi.org/10.1101/2025.02.14.638262">10.1101/2025.02.14.638262</a>
  apa: Naik, S., Keta, Y.-E., Pranjic-Ferscha, K., Hannezo, E. B., Henkes, S., &#38;
    Heisenberg, C.-P. J. (n.d.). Keratins coordinate tissue spreading by balancing
    spreading forces with tissue material properties. <i>bioRxiv</i>. Cold Spring
    Harbor Laboratory. <a href="https://doi.org/10.1101/2025.02.14.638262">https://doi.org/10.1101/2025.02.14.638262</a>
  chicago: Naik, Suyash, Yann-Edwin Keta, Kornelija Pranjic-Ferscha, Edouard B Hannezo,
    Silke Henkes, and Carl-Philipp J Heisenberg. “Keratins Coordinate Tissue Spreading
    by Balancing Spreading Forces with Tissue Material Properties.” <i>BioRxiv</i>.
    Cold Spring Harbor Laboratory, n.d. <a href="https://doi.org/10.1101/2025.02.14.638262">https://doi.org/10.1101/2025.02.14.638262</a>.
  ieee: S. Naik, Y.-E. Keta, K. Pranjic-Ferscha, E. B. Hannezo, S. Henkes, and C.-P.
    J. Heisenberg, “Keratins coordinate tissue spreading by balancing spreading forces
    with tissue material properties,” <i>bioRxiv</i>. Cold Spring Harbor Laboratory.
  ista: Naik S, Keta Y-E, Pranjic-Ferscha K, Hannezo EB, Henkes S, Heisenberg C-PJ.
    Keratins coordinate tissue spreading by balancing spreading forces with tissue
    material properties. bioRxiv, <a href="https://doi.org/10.1101/2025.02.14.638262">10.1101/2025.02.14.638262</a>.
  mla: Naik, Suyash, et al. “Keratins Coordinate Tissue Spreading by Balancing Spreading
    Forces with Tissue Material Properties.” <i>BioRxiv</i>, Cold Spring Harbor Laboratory,
    doi:<a href="https://doi.org/10.1101/2025.02.14.638262">10.1101/2025.02.14.638262</a>.
  short: S. Naik, Y.-E. Keta, K. Pranjic-Ferscha, E.B. Hannezo, S. Henkes, C.-P.J.
    Heisenberg, BioRxiv (n.d.).
corr_author: '1'
date_created: 2025-10-14T07:25:27Z
date_published: 2025-02-17T00:00:00Z
date_updated: 2026-04-07T11:58:57Z
day: '17'
department:
- _id: CaHe
- _id: EdHa
doi: 10.1101/2025.02.14.638262
language:
- iso: eng
main_file_link:
- open_access: '1'
  url: https://doi.org/10.1101/2025.02.14.638262
month: '02'
oa: 1
oa_version: Preprint
publication: bioRxiv
publication_status: draft
publisher: Cold Spring Harbor Laboratory
related_material:
  record:
  - id: '20441'
    relation: dissertation_contains
    status: public
status: public
title: Keratins coordinate tissue spreading by balancing spreading forces with tissue
  material properties
tmp:
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  legal_code_url: https://creativecommons.org/licenses/by-nd/4.0/legalcode
  name: Creative Commons Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0)
  short: CC BY-ND (4.0)
type: preprint
user_id: 8b945eb4-e2f2-11eb-945a-df72226e66a9
year: '2025'
...
---
_id: '12830'
abstract:
- lang: eng
  text: Interstitial fluid (IF) accumulation between embryonic cells is thought to
    be important for embryo patterning and morphogenesis. Here, we identify a positive
    mechanical feedback loop between cell migration and IF relocalization and find
    that it promotes embryonic axis formation during zebrafish gastrulation. We show
    that anterior axial mesendoderm (prechordal plate [ppl]) cells, moving in between
    the yolk cell and deep cell tissue to extend the embryonic axis, compress the
    overlying deep cell layer, thereby causing IF to flow from the deep cell layer
    to the boundary between the yolk cell and the deep cell layer, directly ahead
    of the advancing ppl. This IF relocalization, in turn, facilitates ppl cell protrusion
    formation and migration by opening up the space into which the ppl moves and,
    thereby, the ability of the ppl to trigger IF relocalization by pushing against
    the overlying deep cell layer. Thus, embryonic axis formation relies on a hydraulic
    feedback loop between cell migration and IF relocalization.
acknowledged_ssus:
- _id: PreCl
- _id: Bio
acknowledgement: We thank Andrea Pauli (IMP) and Edouard Hannezo (ISTA) for fruitful
  discussions and support with the SPIM experiments; the Heisenberg group, and especially
  Feyza Nur Arslan and Alexandra Schauer, for discussions and feedback; Michaela Jović
  (ISTA) for help with the quantitative real-time PCR protocol; the bioimaging and
  zebrafish facilities of ISTA for continuous support; Stephan Preibisch (Janelia
  Research Campus) for support with the SPIM data analysis; and Nobuhiro Nakamura
  (Tokyo Institute of Technology) for sharing α1-Na+/K+-ATPase antibody. This work
  was supported by funding from the European Union (European Research Council Advanced
  grant 742573 to C.-P.H.), postdoctoral fellowships from EMBO (LTF-850-2017) and
  HFSP (LT000429/2018-L2) to D.P., and a PhD fellowship from the Studienstiftung des
  deutschen Volkes to F.P.
article_processing_charge: Yes (via OA deal)
article_type: original
author:
- first_name: Karla
  full_name: Huljev, Karla
  id: 44C6F6A6-F248-11E8-B48F-1D18A9856A87
  last_name: Huljev
- first_name: Shayan
  full_name: Shamipour, Shayan
  id: 40B34FE2-F248-11E8-B48F-1D18A9856A87
  last_name: Shamipour
- first_name: Diana C
  full_name: Nunes Pinheiro, Diana C
  id: 2E839F16-F248-11E8-B48F-1D18A9856A87
  last_name: Nunes Pinheiro
  orcid: 0000-0003-4333-7503
- first_name: Friedrich
  full_name: Preusser, Friedrich
  last_name: Preusser
- first_name: Irene
  full_name: Steccari, Irene
  id: 2705C766-9FE2-11EA-B224-C6773DDC885E
  last_name: Steccari
- first_name: Christoph M
  full_name: Sommer, Christoph M
  id: 4DF26D8C-F248-11E8-B48F-1D18A9856A87
  last_name: Sommer
  orcid: 0000-0003-1216-9105
- first_name: Suyash
  full_name: Naik, Suyash
  id: 2C0B105C-F248-11E8-B48F-1D18A9856A87
  last_name: Naik
  orcid: 0000-0001-8421-5508
- 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: Huljev K, Shamipour S, Nunes Pinheiro DC, et al. A hydraulic feedback loop
    between mesendoderm cell migration and interstitial fluid relocalization promotes
    embryonic axis formation in zebrafish. <i>Developmental Cell</i>. 2023;58(7):582-596.e7.
    doi:<a href="https://doi.org/10.1016/j.devcel.2023.02.016">10.1016/j.devcel.2023.02.016</a>
  apa: Huljev, K., Shamipour, S., Nunes Pinheiro, D. C., Preusser, F., Steccari, I.,
    Sommer, C. M., … Heisenberg, C.-P. J. (2023). A hydraulic feedback loop between
    mesendoderm cell migration and interstitial fluid relocalization promotes embryonic
    axis formation in zebrafish. <i>Developmental Cell</i>. Elsevier. <a href="https://doi.org/10.1016/j.devcel.2023.02.016">https://doi.org/10.1016/j.devcel.2023.02.016</a>
  chicago: Huljev, Karla, Shayan Shamipour, Diana C Nunes Pinheiro, Friedrich Preusser,
    Irene Steccari, Christoph M Sommer, Suyash Naik, and Carl-Philipp J Heisenberg.
    “A Hydraulic Feedback Loop between Mesendoderm Cell Migration and Interstitial
    Fluid Relocalization Promotes Embryonic Axis Formation in Zebrafish.” <i>Developmental
    Cell</i>. Elsevier, 2023. <a href="https://doi.org/10.1016/j.devcel.2023.02.016">https://doi.org/10.1016/j.devcel.2023.02.016</a>.
  ieee: K. Huljev <i>et al.</i>, “A hydraulic feedback loop between mesendoderm cell
    migration and interstitial fluid relocalization promotes embryonic axis formation
    in zebrafish,” <i>Developmental Cell</i>, vol. 58, no. 7. Elsevier, p. 582–596.e7,
    2023.
  ista: Huljev K, Shamipour S, Nunes Pinheiro DC, Preusser F, Steccari I, Sommer CM,
    Naik S, Heisenberg C-PJ. 2023. A hydraulic feedback loop between mesendoderm cell
    migration and interstitial fluid relocalization promotes embryonic axis formation
    in zebrafish. Developmental Cell. 58(7), 582–596.e7.
  mla: Huljev, Karla, et al. “A Hydraulic Feedback Loop between Mesendoderm Cell Migration
    and Interstitial Fluid Relocalization Promotes Embryonic Axis Formation in Zebrafish.”
    <i>Developmental Cell</i>, vol. 58, no. 7, Elsevier, 2023, p. 582–596.e7, doi:<a
    href="https://doi.org/10.1016/j.devcel.2023.02.016">10.1016/j.devcel.2023.02.016</a>.
  short: K. Huljev, S. Shamipour, D.C. Nunes Pinheiro, F. Preusser, I. Steccari, C.M.
    Sommer, S. Naik, C.-P.J. Heisenberg, Developmental Cell 58 (2023) 582–596.e7.
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title: A hydraulic feedback loop between mesendoderm cell migration and interstitial
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