13. Kim HY, Jackson TR, Davidson LA (2017) On
the role of mechanics in driving mesenchymalto-epithelial transitions. Semin Cell Dev Biol
67:113–122.
https://doi.org/10.1016/j.
semcdb.2016.05.011
14. Yang J, Weinberg RA (2008) Epithelialmesenchymal transition: at the crossroads of
development and tumor metastasis. Dev Cell
14(6):818–829
15. Thiery JP, Acloque H, Huang RY, Nieto MA
(2009) Epithelial-mesenchymal transitions in
development
and
disease.
Cell
139
(5):871–890
16. Bidarra S, Oliveira P, Rocha S, Saraiva D,
Oliveira C, Barrias C (2016) A 3D in vitro
model to explore the inter-conversion between
epithelial and mesenchymal states during EMT
and its reversion. Sci Rep 6:27072
17. Oster GF, Murray JD, Harris AK (1983)
Mechanical aspects of mesenchymal morphogenesis. J Embryol Exp Morphol 78:83–125
18. Bhumiratana S, Eton RE, Oungoulian SR, Wan
LQ, Ateshian GA, Vunjak-Novakovic G (2014)
Large, stratified, and mechanically functional
human cartilage grown in vitro by mesenchymal condensation. Proc Natl Acad Sci 111
(19):6940–6945
19. Mammoto T, Mammoto A, Torisawa Y-S,
Tat T, Gibbs A, Derda R, Mannix R, de
Bruijn M, Yung CW, Huh D (2011) Mechanochemical control of mesenchymal condensation and embryonic tooth organ formation.
Dev Cell 21(4):758–769
20. Steinberg MS, Gilbert SF (2004) Townes and
Holtfreter (1955): directed movements and
selective adhesion of embryonic amphibian
cells. J Exp Zool A Comp Exp Biol 301
(9):701–706
21. Maitre JL, Berthoumieux H, Krens SF,
Salbreux G, Julicher F, Paluch E, Heisenberg
CP (2012) Adhesion functions in cell sorting
by mechanically coupling the cortices of adhering cells. Science 338(6104):253–256.
https://doi.org/10.1126/science.1225399
22. Harris AK (1976) Is cell sorting caused by
differences in the work of intercellular adhesion? A critique of the Steinberg hypothesis. J
Theor Biol 61(2):267–285
23. Engler AJ, Sen S, Sweeney HL, Discher DE
(2006) Matrix elasticity directs stem cell lineage specification. Cell 126(4):677–689
24. Campa `s O (2016) A toolbox to explore the
mechanics of living embryonic tissues. Semin
Cell Dev Biol 55:119–130
25. Shawky JH, Davidson LA (2015) Tissue
mechanics and adhesion during embryo development. Dev Biol 401(1):152–164. https://
doi.org/10.1016/j.ydbio.2014.12.005
26. Cost A-L, Ringer P, Chrostek-Grashoff A,
Grashoff C (2015) How to measure molecular
forces in cells: a guide to evaluating geneticallyencoded FRET-based tension sensors. Cell
Mol Bioeng 8(1):96–105. https://doi.org/
10.1007/s12195-014-0368-1
27. Conway DE, Breckenridge MT, Hinde E,
Gratton E, Chen CS, Schwartz MA (2013)
Fluid shear stress on endothelial cells modulates mechanical tension across VE-cadherin
and
PECAM-1.
Curr
Biol
23
(11):1024–1030. https://doi.org/10.1016/j.
cub.2013.04.049
28. Grashoff C, Hoffman BD, Brenner MD,
Zhou R, Parsons M, Yang MT, McLean MA,
Sligar SG, Chen CS, Ha T, Schwartz MA
(2010) Measuring mechanical tension across
vinculin reveals regulation of focal adhesion
dynamics.
Nature
466(7303):263–266.
https://doi.org/10.1038/nature09198
29. Leerberg JM, Gomez GA, Verma S, Moussa
EJ, Wu SK, Priya R, Hoffman BD,
Grashoff C, Schwartz MA, Yap AS (2014)
Tension-sensitive actin assembly supports contractility at the epithelial zonula adherens. Curr
Biol 24(15):1689–1699
30. Austen K, Ringer P, Mehlich A, ChrostekGrashoff A, Kluger C, Klingner C, Sabass B,
Zent R, Rief M, Grashoff C (2015) Extracellular rigidity sensing by talin isoform-specific
mechanical linkages. Nat Cell Biol 17
(12):1597–1606
31. Kumar A, Ouyang M, Van den Dries K,
McGhee EJ, Tanaka K, Anderson MD,
Groisman A, Goult BT, Anderson KI, Schwartz
MA (2016) Talin tension sensor reveals novel
features of focal adhesion force transmission
and mechanosensitivity. J Cell Biol 213
(3):371–383
32. Aragona M, Panciera T, Manfrin A, Giulitti S,
Michielin F, Elvassore N, Dupont S, Piccolo S
(2013) A mechanical checkpoint controls multicellular growth through YAP/TAZ regulation by actin-processing factors. Cell 154
(5):1047–1059
33. Kim HY, Jackson TR, Stuckenholz C, Davidson LA (2020) Tissue mechanics drives regeneration of a mucociliated epidermis on the
surface of Xenopus embryonic aggregates.
Nat Commun 11(1):1–10. https://doi.org/
10.1038/s41467-020-14385-y
34. Davidson LA, Dzamba BD, Keller R, Desimone DW (2008) Live imaging of cell protrusive activity, and extracellular matrix assembly
and remodeling during morphogenesis in the
frog, Xenopus laevis. Dev Dyn 237
(10):2684–2692. https://doi.org/10.1002/
dvdy.21600
286
Hye Young Kim and Lance A. Davidson
the role of mechanics in driving mesenchymalto-epithelial transitions. Semin Cell Dev Biol
67:113–122.
https://doi.org/10.1016/j.
semcdb.2016.05.011
14. Yang J, Weinberg RA (2008) Epithelialmesenchymal transition: at the crossroads of
development and tumor metastasis. Dev Cell
14(6):818–829
15. Thiery JP, Acloque H, Huang RY, Nieto MA
(2009) Epithelial-mesenchymal transitions in
development
and
disease.
Cell
139
(5):871–890
16. Bidarra S, Oliveira P, Rocha S, Saraiva D,
Oliveira C, Barrias C (2016) A 3D in vitro
model to explore the inter-conversion between
epithelial and mesenchymal states during EMT
and its reversion. Sci Rep 6:27072
17. Oster GF, Murray JD, Harris AK (1983)
Mechanical aspects of mesenchymal morphogenesis. J Embryol Exp Morphol 78:83–125
18. Bhumiratana S, Eton RE, Oungoulian SR, Wan
LQ, Ateshian GA, Vunjak-Novakovic G (2014)
Large, stratified, and mechanically functional
human cartilage grown in vitro by mesenchymal condensation. Proc Natl Acad Sci 111
(19):6940–6945
19. Mammoto T, Mammoto A, Torisawa Y-S,
Tat T, Gibbs A, Derda R, Mannix R, de
Bruijn M, Yung CW, Huh D (2011) Mechanochemical control of mesenchymal condensation and embryonic tooth organ formation.
Dev Cell 21(4):758–769
20. Steinberg MS, Gilbert SF (2004) Townes and
Holtfreter (1955): directed movements and
selective adhesion of embryonic amphibian
cells. J Exp Zool A Comp Exp Biol 301
(9):701–706
21. Maitre JL, Berthoumieux H, Krens SF,
Salbreux G, Julicher F, Paluch E, Heisenberg
CP (2012) Adhesion functions in cell sorting
by mechanically coupling the cortices of adhering cells. Science 338(6104):253–256.
https://doi.org/10.1126/science.1225399
22. Harris AK (1976) Is cell sorting caused by
differences in the work of intercellular adhesion? A critique of the Steinberg hypothesis. J
Theor Biol 61(2):267–285
23. Engler AJ, Sen S, Sweeney HL, Discher DE
(2006) Matrix elasticity directs stem cell lineage specification. Cell 126(4):677–689
24. Campa `s O (2016) A toolbox to explore the
mechanics of living embryonic tissues. Semin
Cell Dev Biol 55:119–130
25. Shawky JH, Davidson LA (2015) Tissue
mechanics and adhesion during embryo development. Dev Biol 401(1):152–164. https://
doi.org/10.1016/j.ydbio.2014.12.005
26. Cost A-L, Ringer P, Chrostek-Grashoff A,
Grashoff C (2015) How to measure molecular
forces in cells: a guide to evaluating geneticallyencoded FRET-based tension sensors. Cell
Mol Bioeng 8(1):96–105. https://doi.org/
10.1007/s12195-014-0368-1
27. Conway DE, Breckenridge MT, Hinde E,
Gratton E, Chen CS, Schwartz MA (2013)
Fluid shear stress on endothelial cells modulates mechanical tension across VE-cadherin
and
PECAM-1.
Curr
Biol
23
(11):1024–1030. https://doi.org/10.1016/j.
cub.2013.04.049
28. Grashoff C, Hoffman BD, Brenner MD,
Zhou R, Parsons M, Yang MT, McLean MA,
Sligar SG, Chen CS, Ha T, Schwartz MA
(2010) Measuring mechanical tension across
vinculin reveals regulation of focal adhesion
dynamics.
Nature
466(7303):263–266.
https://doi.org/10.1038/nature09198
29. Leerberg JM, Gomez GA, Verma S, Moussa
EJ, Wu SK, Priya R, Hoffman BD,
Grashoff C, Schwartz MA, Yap AS (2014)
Tension-sensitive actin assembly supports contractility at the epithelial zonula adherens. Curr
Biol 24(15):1689–1699
30. Austen K, Ringer P, Mehlich A, ChrostekGrashoff A, Kluger C, Klingner C, Sabass B,
Zent R, Rief M, Grashoff C (2015) Extracellular rigidity sensing by talin isoform-specific
mechanical linkages. Nat Cell Biol 17
(12):1597–1606
31. Kumar A, Ouyang M, Van den Dries K,
McGhee EJ, Tanaka K, Anderson MD,
Groisman A, Goult BT, Anderson KI, Schwartz
MA (2016) Talin tension sensor reveals novel
features of focal adhesion force transmission
and mechanosensitivity. J Cell Biol 213
(3):371–383
32. Aragona M, Panciera T, Manfrin A, Giulitti S,
Michielin F, Elvassore N, Dupont S, Piccolo S
(2013) A mechanical checkpoint controls multicellular growth through YAP/TAZ regulation by actin-processing factors. Cell 154
(5):1047–1059
33. Kim HY, Jackson TR, Stuckenholz C, Davidson LA (2020) Tissue mechanics drives regeneration of a mucociliated epidermis on the
surface of Xenopus embryonic aggregates.
Nat Commun 11(1):1–10. https://doi.org/
10.1038/s41467-020-14385-y
34. Davidson LA, Dzamba BD, Keller R, Desimone DW (2008) Live imaging of cell protrusive activity, and extracellular matrix assembly
and remodeling during morphogenesis in the
frog, Xenopus laevis. Dev Dyn 237
(10):2684–2692. https://doi.org/10.1002/
dvdy.21600
286
Hye Young Kim and Lance A. Davidson