---
OA_place: publisher
_id: '22804'
abstract:
- lang: eng
text: 'Embryo cleavage — a series of rapid, reductive cell divisions — is the first
morphogenetic movement following fertilisation. It lays the foundation for subsequent
developmental events, including gastrulation, germ layer specification, organogenesis,
and the establishment of the overall body plan. Two major modes of cleavage exist
in the animal kingdom: holoblastic (complete) and meroblastic (incomplete) cleavage.
Because holoblastic cleavage resembles canonical cytokinesis, its biochemical
and mechanical basis has been extensively studied — an annular contractile ring
forms at the equator and constricts in a purse-string-like manner, leading to
the complete separation of two daughter cells. In contrast, although meroblastic
cleavage occurs widely across the animal kingdom (e.g. in fish, reptiles, birds,
and cephalopod molluscs), its mechanical basis remains largely unclear. During
meroblastic cleavage, the cytokinetic furrow forms only at one pole and does not
traverse the entire embryo, raising the question of how cytokinesis proceeds in
the absence of a closed contractile ring. Moreover, the resulting daughter cells
are not fully separated from the underlying yolk compartment, and how these blastomeres
are subsequently cellularised remains unknown. This thesis takes the zebrafish
as a model organism to address both questions. The first part characterises the
biochemical and mechanical mechanisms underlying non-canonical meroblastic cleavage,
revealing a two-phase process in which actomyosin cable contraction and cadherin-mediated
membrane adhesion act sequentially to drive furrow ingression and invagination.
The second part sheds light on the spatiotemporal dynamics by which individual
blastomeres become cellularised, and uncovers a previously unrecognised contribution
of central blastomeres to the yolk syncytial layer.'
acknowledged_ssus:
- _id: Bio
- _id: LifeSc
acknowledgement: "I gratefully acknowledge the NOMIS Stiftung (Project ID 5061.844,
Cytoplasmic\r\nself-organization into cell-like compartments as a common guiding
principle in\r\nearly animal development) for funding my doctoral studies."
alternative_title:
- ISTA Thesis
article_processing_charge: No
author:
- first_name: Xin
full_name: Tong, Xin
id: 50F65CDC-AA30-11E9-A72B-8A12E6697425
last_name: Tong
citation:
ama: Tong X. Towards a deeper understanding of meroblastic cleavage. 2026. doi:10.15479/AT-ISTA-22804
apa: Tong, X. (2026). Towards a deeper understanding of meroblastic cleavage.
Institute of Science and Technology Austria. https://doi.org/10.15479/AT-ISTA-22804
chicago: Tong, Xin. “Towards a Deeper Understanding of Meroblastic Cleavage.” Institute
of Science and Technology Austria, 2026. https://doi.org/10.15479/AT-ISTA-22804.
ieee: X. Tong, “Towards a deeper understanding of meroblastic cleavage,” Institute
of Science and Technology Austria, 2026.
ista: Tong X. 2026. Towards a deeper understanding of meroblastic cleavage. Institute
of Science and Technology Austria.
mla: Tong, Xin. Towards a Deeper Understanding of Meroblastic Cleavage. Institute
of Science and Technology Austria, 2026, doi:10.15479/AT-ISTA-22804.
short: X. Tong, Towards a Deeper Understanding of Meroblastic Cleavage, Institute
of Science and Technology Austria, 2026.
corr_author: '1'
date_created: 2026-09-04T11:54:20Z
date_published: 2026-07-12T00:00:00Z
date_updated: 2026-10-05T08:56:10Z
day: '12'
ddc:
- '596'
- '532'
- '572'
degree_awarded: PhD
department:
- _id: GradSch
- _id: CaHe
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doi_confirm: '1'
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fulldoi: https://doi.org/10.15479/AT-ISTA-22804
has_accepted_license: '1'
keyword:
- meroblastic cleavage
- cytokinesis
- active gel theory
- embryogenesis
- zebrafish
- adhesion
language:
- iso: eng
month: '07'
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oa_version: Published Version
page: '121'
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name: Cytoplasmic self-organization into cell-like compartments as a common guiding
principle in early animal development
publication_identifier:
isbn:
- 978-3-99078-080-0
issn:
- 2663-337X
publication_status: published
publisher: Institute of Science and Technology Austria
related_material:
record:
- id: '22777'
relation: research_data
status: public
- id: '22807'
relation: part_of_dissertation
status: public
researchdata_availability: upon request
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
title: Towards a deeper understanding of meroblastic cleavage
tmp:
image: /images/cc_by_nc.png
legal_code_url: https://creativecommons.org/licenses/by-nc/4.0/legalcode
name: Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
short: CC BY-NC (4.0)
type: dissertation
user_id: 8b945eb4-e2f2-11eb-945a-df72226e66a9
year: '2026'
...
---
OA_place: repository
_id: '22777'
acknowledged_ssus:
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- _id: LifeSc
article_processing_charge: No
author:
- first_name: Xin
full_name: Tong, Xin
id: 50F65CDC-AA30-11E9-A72B-8A12E6697425
last_name: Tong
citation:
ama: Tong X. Towards a deeper understanding of meroblastic cleavage - supplementary
movies. 2026. doi:10.15479/AT-ISTA-22777
apa: Tong, X. (2026). Towards a deeper understanding of meroblastic cleavage - supplementary
movies. Institute of Science and Technology Austria. https://doi.org/10.15479/AT-ISTA-22777
chicago: Tong, Xin. “Towards a Deeper Understanding of Meroblastic Cleavage - Supplementary
Movies.” Institute of Science and Technology Austria, 2026. https://doi.org/10.15479/AT-ISTA-22777.
ieee: X. Tong, “Towards a deeper understanding of meroblastic cleavage - supplementary
movies.” Institute of Science and Technology Austria, 2026.
ista: Tong X. 2026. Towards a deeper understanding of meroblastic cleavage - supplementary
movies, Institute of Science and Technology Austria, 10.15479/AT-ISTA-22777.
mla: Tong, Xin. Towards a Deeper Understanding of Meroblastic Cleavage - Supplementary
Movies. Institute of Science and Technology Austria, 2026, doi:10.15479/AT-ISTA-22777.
short: X. Tong, (2026).
corr_author: '1'
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date_published: 2026-09-04T00:00:00Z
date_updated: 2026-10-05T08:56:09Z
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fulldoi: https://doi.org/10.15479/AT-ISTA-22777
has_accepted_license: '1'
month: '9'
oa: 1
oa_version: None
project:
- _id: 9B861AAC-BA93-11EA-9121-9846C619BF3A
name: NOMIS Fellowship Program
- _id: 05943252-7A3F-11EA-A408-12923DDC885E
call_identifier: H2020
grant_number: '851288'
name: Design Principles of Branching Morphogenesis
publisher: Institute of Science and Technology Austria
related_material:
record:
- id: '22804'
relation: used_in_publication
status: public
status: public
title: Towards a deeper understanding of meroblastic cleavage - supplementary movies
tmp:
image: /images/cc_by_nc.png
legal_code_url: https://creativecommons.org/licenses/by-nc/4.0/legalcode
name: Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
short: CC BY-NC (4.0)
type: research_data
user_id: 68b8ca59-c5b3-11ee-8790-cd641c68093d
year: '2026'
...
---
OA_place: repository
OA_type: green
_id: '22807'
abstract:
- lang: eng
text: 'Cleavage - the series of rapid cell divisions that follow fertilization -
marks the onset of metazoan development and represents a deeply conserved evolutionary
process. Across animals, two principal modes exist: complete (holoblastic) and
incomplete (meroblastic) cleavage. While holoblastic cleavage resembles conventional
cytokinesis both in vitro and in vivo, the mechanisms underlying meroblastic cleavage
have remained poorly understood. Using zebrafish embryos as a model, we show that
meroblastic cleavage proceeds through a distinct two-step mechanism. The process
begins with the assembly and contraction of a large, arc-shaped actomyosin cable.
However, this contractile event alone is insufficient to complete division. A
second phase, driven by cadherin-mediated membrane adhesion, is required to invaginate
the furrow ridge. Strikingly, this transition depends on mechanical uncoupling
of the contractile cable from the surrounding cortex. We demonstrate that such
uncoupling arises from an active nematic instability, which both enhances contractility
along the cable and generates actin depletion zones that relieve lateral connections.
Together, these findings reveal that meroblastic cleavage is governed not by a
single actomyosin-based event but by a sequential interplay between cytoskeletal
contraction and cadherin-dependent adhesion, highlighting a mechanism fundamentally
distinct from canonical cytokinesis.'
acknowledgement: We are grateful to the members of the Hannezo and Heisenberg groups
for discussions and technical advice. We also thank the Imaging and Optics Facility
and the Lab Support Facility at ISTA for their continuous support. Y.I.L. acknowledges
funding from the European Union’s Horizon 2020 research and innovation programme
under the Marie Skodowska-Curie Grant Agreement No. 101034413. The research was
supported by funding to C.-P.H. from the NOMIS Foundation (Project ID 1.844) and
to E.H. from the European Research Council (ERC) under the European Union’s Horizon
2020 research and innovation programme (grant agreement no. 851288).
article_processing_charge: No
author:
- first_name: Xin
full_name: Tong, Xin
id: 50F65CDC-AA30-11E9-A72B-8A12E6697425
last_name: Tong
- first_name: Yuting I
full_name: Li, Yuting I
id: ee7a5ca8-8b71-11ed-b662-b3341c05b7eb
last_name: Li
- first_name: Joséphine
full_name: Schelle, Joséphine
last_name: Schelle
- 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
biorxivid: 1
citation:
ama: Tong X, Li YI, Schelle J, Hannezo EB, Heisenberg C-PJ. Non-canonical cytokinesis
driven by mechanical uncoupling via nematic flows and adhesion-based invagination.
bioRxiv. doi:10.1101/2025.10.15.682552
apa: Tong, X., Li, Y. I., Schelle, J., Hannezo, E. B., & Heisenberg, C.-P. J.
(n.d.). Non-canonical cytokinesis driven by mechanical uncoupling via nematic
flows and adhesion-based invagination. bioRxiv. https://doi.org/10.1101/2025.10.15.682552
chicago: Tong, Xin, Yuting I Li, Joséphine Schelle, Edouard B Hannezo, and Carl-Philipp
J Heisenberg. “Non-Canonical Cytokinesis Driven by Mechanical Uncoupling via Nematic
Flows and Adhesion-Based Invagination.” BioRxiv, n.d. https://doi.org/10.1101/2025.10.15.682552.
ieee: X. Tong, Y. I. Li, J. Schelle, E. B. Hannezo, and C.-P. J. Heisenberg, “Non-canonical
cytokinesis driven by mechanical uncoupling via nematic flows and adhesion-based
invagination,” bioRxiv. .
ista: Tong X, Li YI, Schelle J, Hannezo EB, Heisenberg C-PJ. Non-canonical cytokinesis
driven by mechanical uncoupling via nematic flows and adhesion-based invagination.
bioRxiv, 10.1101/2025.10.15.682552.
mla: Tong, Xin, et al. “Non-Canonical Cytokinesis Driven by Mechanical Uncoupling
via Nematic Flows and Adhesion-Based Invagination.” BioRxiv, doi:10.1101/2025.10.15.682552.
short: X. Tong, Y.I. Li, J. Schelle, E.B. Hannezo, C.-P.J. Heisenberg, BioRxiv (n.d.).
corr_author: '1'
date_created: 2026-09-04T12:32:21Z
date_published: 2025-10-15T00:00:00Z
date_updated: 2026-10-05T08:56:09Z
day: '15'
department:
- _id: EdHa
- _id: CaHe
doi: 10.1101/2025.10.15.682552
ec_funded: 1
external_id:
biorxivid:
- 2025.10.15.682552
fulldoi: https://doi.org/10.1101/2025.10.15.682552
language:
- iso: eng
main_file_link:
- open_access: '1'
url: https://doi.org/10.1101/2025.10.15.682552
month: '10'
oa: 1
oa_version: Preprint
project:
- _id: fc2ed2f7-9c52-11eb-aca3-c01059dda49c
call_identifier: H2020
grant_number: '101034413'
name: 'IST-BRIDGE: International postdoctoral program'
- _id: 05943252-7A3F-11EA-A408-12923DDC885E
call_identifier: H2020
grant_number: '851288'
name: Design Principles of Branching Morphogenesis
- _id: 9B861AAC-BA93-11EA-9121-9846C619BF3A
name: NOMIS Fellowship Program
publication: bioRxiv
publication_status: draft
related_material:
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relation: dissertation_contains
status: public
status: public
title: Non-canonical cytokinesis driven by mechanical uncoupling via nematic flows
and adhesion-based invagination
tmp:
image: /images/cc_by_nc.png
legal_code_url: https://creativecommons.org/licenses/by-nc/4.0/legalcode
name: Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0)
short: CC BY-NC (4.0)
type: preprint
user_id: 8b945eb4-e2f2-11eb-945a-df72226e66a9
year: '2025'
...
---
DOAJ_listed: '1'
OA_place: publisher
OA_type: gold
_id: '19404'
abstract:
- lang: eng
text: Cell migration is a fundamental process during embryonic development. Most
studies in vivo have focused on the migration of cells using the extracellular
matrix (ECM) as their substrate for migration. In contrast, much less is known
about how cells migrate on other cells, as found in early embryos when the ECM
has not yet formed. Here, we show that lateral mesendoderm (LME) cells in the
early zebrafish gastrula use the ectoderm as their substrate for migration. We
show that the lateral ectoderm is permissive for the animal-pole-directed migration
of LME cells, while the ectoderm at the animal pole halts it. These differences
in permissiveness depend on the lateral ectoderm being more cohesive than the
animal ectoderm, a property controlled by bone morphogenetic protein (BMP) signaling
within the ectoderm. Collectively, these findings identify ectoderm tissue cohesion
as one critical factor in regulating LME migration during zebrafish gastrulation.
acknowledged_ssus:
- _id: Bio
- _id: LifeSc
- _id: ScienComp
acknowledgement: 'We are grateful to the colleagues who contributed to this work with
discussions, technical advice, and feedback on the manuscript: Irene Steccari, David
Labrousse Arias and the other members of the Heisenberg lab, Nicole Amberg, Florian
Pauler, Nicoletta Petridou, Elena Scarpa, and Edouard Hannezo. We also thank the
Imaging and Optics Facility, the Life Science Facility, and the Scientific Computing
Unit at ISTA for support. The Next Generation Sequencing Facility at Vienna BioCenter
Core Facilities performed the RNA-seq for animal and lateral ectoderm. D.B.B. was
supported by the NOMIS Foundation as a NOMIS Fellow and by an EMBO Postdoctoral
Fellowship (ALTF 343-2022). S. Tavano was supported by an EMBO Postdoctoral Fellowship
(ALTF 1159-2018).'
article_number: '115387'
article_processing_charge: Yes
article_type: original
author:
- first_name: Ste
full_name: Tavano, Ste
id: 2F162F0C-F248-11E8-B48F-1D18A9856A87
last_name: Tavano
orcid: 0000-0001-9970-7804
- first_name: David
full_name: Brückner, David
id: e1e86031-6537-11eb-953a-f7ab92be508d
last_name: Brückner
orcid: 0000-0001-7205-2975
- first_name: Saren
full_name: Tasciyan, Saren
id: 4323B49C-F248-11E8-B48F-1D18A9856A87
last_name: Tasciyan
orcid: 0000-0003-1671-393X
- first_name: Xin
full_name: Tong, Xin
id: 50F65CDC-AA30-11E9-A72B-8A12E6697425
last_name: Tong
- first_name: Roland
full_name: Kardos, Roland
id: 4039350E-F248-11E8-B48F-1D18A9856A87
last_name: Kardos
- first_name: Alexandra
full_name: Schauer, Alexandra
id: 30A536BA-F248-11E8-B48F-1D18A9856A87
last_name: Schauer
orcid: 0000-0001-7659-9142
- first_name: Robert
full_name: Hauschild, Robert
id: 4E01D6B4-F248-11E8-B48F-1D18A9856A87
last_name: Hauschild
orcid: 0000-0001-9843-3522
- 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: Tavano S, Brückner D, Tasciyan S, et al. BMP-dependent patterning of ectoderm
tissue material properties modulates lateral mesendoderm cell migration during
early zebrafish gastrulation. Cell Reports. 2025;44(3). doi:10.1016/j.celrep.2025.115387
apa: Tavano, S., Brückner, D., Tasciyan, S., Tong, X., Kardos, R., Schauer, A.,
… Heisenberg, C.-P. J. (2025). BMP-dependent patterning of ectoderm tissue material
properties modulates lateral mesendoderm cell migration during early zebrafish
gastrulation. Cell Reports. Cell Press. https://doi.org/10.1016/j.celrep.2025.115387
chicago: Tavano, Ste, David Brückner, Saren Tasciyan, Xin Tong, Roland Kardos, Alexandra
Schauer, Robert Hauschild, and Carl-Philipp J Heisenberg. “BMP-Dependent Patterning
of Ectoderm Tissue Material Properties Modulates Lateral Mesendoderm Cell Migration
during Early Zebrafish Gastrulation.” Cell Reports. Cell Press, 2025. https://doi.org/10.1016/j.celrep.2025.115387.
ieee: S. Tavano et al., “BMP-dependent patterning of ectoderm tissue material
properties modulates lateral mesendoderm cell migration during early zebrafish
gastrulation,” Cell Reports, vol. 44, no. 3. Cell Press, 2025.
ista: Tavano S, Brückner D, Tasciyan S, Tong X, Kardos R, Schauer A, Hauschild R,
Heisenberg C-PJ. 2025. BMP-dependent patterning of ectoderm tissue material properties
modulates lateral mesendoderm cell migration during early zebrafish gastrulation.
Cell Reports. 44(3), 115387.
mla: Tavano, Ste, et al. “BMP-Dependent Patterning of Ectoderm Tissue Material Properties
Modulates Lateral Mesendoderm Cell Migration during Early Zebrafish Gastrulation.”
Cell Reports, vol. 44, no. 3, 115387, Cell Press, 2025, doi:10.1016/j.celrep.2025.115387.
short: S. Tavano, D. Brückner, S. Tasciyan, X. Tong, R. Kardos, A. Schauer, R. Hauschild,
C.-P.J. Heisenberg, Cell Reports 44 (2025).
corr_author: '1'
date_created: 2025-03-16T23:01:24Z
date_published: 2025-03-25T00:00:00Z
date_updated: 2026-10-20T08:20:45Z
day: '25'
ddc:
- '570'
department:
- _id: CaHe
- _id: EdHa
- _id: MiSi
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doi: 10.1016/j.celrep.2025.115387
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title: BMP-dependent patterning of ectoderm tissue material properties modulates lateral
mesendoderm cell migration during early zebrafish gastrulation
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...