by advective triggering of a pattern-forming
system. Science 334:1137–1141
54. Mayer M, Depken M, Bois JS, Ju ¨licher F, Grill
SW (2010) Anisotropies in cortical tension
reveal the physical basis of polarizing cortical
flows. Nature 467:617–621
55. McFadden WM, McCall PM, Gardel ML,
Munro EM (2017) Filament turnover tunes
both force generation and dissipation to control long-range flows in a model actomyosin
cortex. PLoS Comput Biol 13:e1005811
56. Benton R, Johnston DS (2003) Drosophila
PAR-1 and 14-3-3 inhibit Bazooka/PAR-3
to establish complementary cortical domains
in polarized cells. Cell 115:691–704
57. Joberty G, Petersen C, Gao L, Macara IG
(2000) The cell-polarity protein Par6 links
Par3 and atypical protein kinase C to Cdc42.
Nat Cell Biol 2:531–539
58. Renschler FA, Bruekner SR, Salomon PL,
Mukherjee A, Kullmann L, Schu ¨tz-Stoffregen
MC, Henzler C, Pawson T, Krahn MP, Wiesner S (2018) Structural basis for the interaction between the cell polarity proteins Par3
and Par6. Sci Signal 11:517
59. Uroz M, Wistorf S, Serra-Picamal X, Conte V,
Sales-Pardo M, Roca-Cusachs P, Guimera ` R,
Trepat X (2018) Regulation of cell cycle progression by cell-cell and cell-matrix forces.
Nat Cell Biol 20:646–654
60. Ladoux B, Me `ge R-M, Trepat X (2016)
Front-rear polarization by mechanical cues:
from single cells to tissues. Trends Cell Biol
26:420–433
61. Trepat X, Wasserman MR, Angelini TE,
Millet E, Weitz DA, Butler JP, Fredberg JJ
(2009) Physical forces during collective cell
migration. Nat Phys 5:426–430
62. Etienne-Manneville S, Hall A (2003) Cell
polarity: Par6, aPKC and cytoskeletal crosstalk. Curr Opin Cell Biol 15:67–72
63. Lewis A, Di Ciano C, Rotstein OD, Kapus A
(2002) Osmotic stress activates Rac and
Cdc42 in neutrophils: role in hypertonicityinduced actin polymerization. Am J Phys Cell
Phys 282:C271–C279
64. Li S, Chen BPC, Azuma N, Hu Y-L, Wu SZ,
Sumpio BE, Shyy JY-J, Chien S (1999) Distinct roles for the small GTPases Cdc42 and
Rho in endothelial responses to shear stress. J
Clin Invest 103:1141–1150
65. Tang DD, Gunst SJ (2004) The small GTPase
Cdc42 regulates actin polymerization and
tension development during contractile stimulation of smooth muscle. J Biol Chem
279:51722–51728
66. Plutoni C, Bazellieres E, Le Borgne-RochetM, Comunale F, Brugues A, Se ´veno M,
Planchon D, Thuault S, Morin N, Bodin S,
Trepat X, Gauthier-Rouvie `re C (2016)
P-cadherin promotes collective cell migration
via a Cdc42-mediated increase in mechanical
forces. J Cell Biol 212:199–217
67. Whitney DS, Peterson FC, Kittell AW, Egner
JM, Prehoda KE, Volkman BF (2016) Binding of crumbs to the Par-6 CRIB-PDZ module is regulated by Cdc42. Biochemistry
55:1455–1461
68. Garrard SM, Capaldo CT, Gao L, Rosen MK,
Macara IG, Tomchick DR (2003) Structure
of Cdc42 in a complex with the GTPasebinding domain of the cell polarity protein,
Par6. EMBO J 22:1125–1133
69. Su W-H, Mruk DD, Wong EWP, Lui W-Y,
Cheng CY (2012) Polarity protein complex
Scribble/Lgl/Dlg and epithelial cell barriers.
Adv Exp Med Biol 763:149–170
70. Lee S, Fan S, Makarova O, Straight S, Margolis B (2002) A novel and conserved proteinprotein interaction domain of mammalian
Lin-2/CASK binds and recruits SAP97 to
the lateral surface of epithelia. Mol Cell Biol
22:1778–1791
71. Zhu Z-Q, Wang D, Xiang D, Yuan Y-X, Wang
Y (2014) Calcium/calmodulin-dependent
serine protein kinase is involved in exendin4-induced insulin secretion in INS-1 cells.
Metabolism 63:120–126
72. Jiang M, Qiu J, Zhang L, Lu ¨ D, Long M,
Chen L, Luo X (2016) Changes in tension
regulates proliferation and migration of fibroblasts by remodeling expression of ECM proteins. Exp Ther Med 12:1542–1550
73. Porter AP, White GRM, Mack NA, Malliri A
(2019) The interaction between CASK and
the tumour suppressor Dlg1 regulates mitotic
spindle orientation in mammalian epithelia. J
Cell Sci 132:jcs230086
74. Hannezo E, Prost J, Joanny J-F (2014) Theory of epithelial sheet morphology in three
dimensions. Proc Natl Acad Sci U S A
111:27–32
75. Takeichi M (2014) Dynamic contacts: rearranging adherens junctions to drive epithelial
remodelling. Nat Rev Mol Cell Biol
15:397–410
76. Becker S, Wandel E, Wobus M, Schneider R,
Amasheh S, Sittig D, Kerner C, Naumann R,
Hamann J, Aust G (2010) Overexpression of
CD97 in intestinal epithelial cells of transgenic mice attenuates colitis by strengthening
adherens junctions. PLoS One 5:e8507
272
Elsa Bazellie ` res and Andre ´ Le Bivic
system. Science 334:1137–1141
54. Mayer M, Depken M, Bois JS, Ju ¨licher F, Grill
SW (2010) Anisotropies in cortical tension
reveal the physical basis of polarizing cortical
flows. Nature 467:617–621
55. McFadden WM, McCall PM, Gardel ML,
Munro EM (2017) Filament turnover tunes
both force generation and dissipation to control long-range flows in a model actomyosin
cortex. PLoS Comput Biol 13:e1005811
56. Benton R, Johnston DS (2003) Drosophila
PAR-1 and 14-3-3 inhibit Bazooka/PAR-3
to establish complementary cortical domains
in polarized cells. Cell 115:691–704
57. Joberty G, Petersen C, Gao L, Macara IG
(2000) The cell-polarity protein Par6 links
Par3 and atypical protein kinase C to Cdc42.
Nat Cell Biol 2:531–539
58. Renschler FA, Bruekner SR, Salomon PL,
Mukherjee A, Kullmann L, Schu ¨tz-Stoffregen
MC, Henzler C, Pawson T, Krahn MP, Wiesner S (2018) Structural basis for the interaction between the cell polarity proteins Par3
and Par6. Sci Signal 11:517
59. Uroz M, Wistorf S, Serra-Picamal X, Conte V,
Sales-Pardo M, Roca-Cusachs P, Guimera ` R,
Trepat X (2018) Regulation of cell cycle progression by cell-cell and cell-matrix forces.
Nat Cell Biol 20:646–654
60. Ladoux B, Me `ge R-M, Trepat X (2016)
Front-rear polarization by mechanical cues:
from single cells to tissues. Trends Cell Biol
26:420–433
61. Trepat X, Wasserman MR, Angelini TE,
Millet E, Weitz DA, Butler JP, Fredberg JJ
(2009) Physical forces during collective cell
migration. Nat Phys 5:426–430
62. Etienne-Manneville S, Hall A (2003) Cell
polarity: Par6, aPKC and cytoskeletal crosstalk. Curr Opin Cell Biol 15:67–72
63. Lewis A, Di Ciano C, Rotstein OD, Kapus A
(2002) Osmotic stress activates Rac and
Cdc42 in neutrophils: role in hypertonicityinduced actin polymerization. Am J Phys Cell
Phys 282:C271–C279
64. Li S, Chen BPC, Azuma N, Hu Y-L, Wu SZ,
Sumpio BE, Shyy JY-J, Chien S (1999) Distinct roles for the small GTPases Cdc42 and
Rho in endothelial responses to shear stress. J
Clin Invest 103:1141–1150
65. Tang DD, Gunst SJ (2004) The small GTPase
Cdc42 regulates actin polymerization and
tension development during contractile stimulation of smooth muscle. J Biol Chem
279:51722–51728
66. Plutoni C, Bazellieres E, Le Borgne-RochetM, Comunale F, Brugues A, Se ´veno M,
Planchon D, Thuault S, Morin N, Bodin S,
Trepat X, Gauthier-Rouvie `re C (2016)
P-cadherin promotes collective cell migration
via a Cdc42-mediated increase in mechanical
forces. J Cell Biol 212:199–217
67. Whitney DS, Peterson FC, Kittell AW, Egner
JM, Prehoda KE, Volkman BF (2016) Binding of crumbs to the Par-6 CRIB-PDZ module is regulated by Cdc42. Biochemistry
55:1455–1461
68. Garrard SM, Capaldo CT, Gao L, Rosen MK,
Macara IG, Tomchick DR (2003) Structure
of Cdc42 in a complex with the GTPasebinding domain of the cell polarity protein,
Par6. EMBO J 22:1125–1133
69. Su W-H, Mruk DD, Wong EWP, Lui W-Y,
Cheng CY (2012) Polarity protein complex
Scribble/Lgl/Dlg and epithelial cell barriers.
Adv Exp Med Biol 763:149–170
70. Lee S, Fan S, Makarova O, Straight S, Margolis B (2002) A novel and conserved proteinprotein interaction domain of mammalian
Lin-2/CASK binds and recruits SAP97 to
the lateral surface of epithelia. Mol Cell Biol
22:1778–1791
71. Zhu Z-Q, Wang D, Xiang D, Yuan Y-X, Wang
Y (2014) Calcium/calmodulin-dependent
serine protein kinase is involved in exendin4-induced insulin secretion in INS-1 cells.
Metabolism 63:120–126
72. Jiang M, Qiu J, Zhang L, Lu ¨ D, Long M,
Chen L, Luo X (2016) Changes in tension
regulates proliferation and migration of fibroblasts by remodeling expression of ECM proteins. Exp Ther Med 12:1542–1550
73. Porter AP, White GRM, Mack NA, Malliri A
(2019) The interaction between CASK and
the tumour suppressor Dlg1 regulates mitotic
spindle orientation in mammalian epithelia. J
Cell Sci 132:jcs230086
74. Hannezo E, Prost J, Joanny J-F (2014) Theory of epithelial sheet morphology in three
dimensions. Proc Natl Acad Sci U S A
111:27–32
75. Takeichi M (2014) Dynamic contacts: rearranging adherens junctions to drive epithelial
remodelling. Nat Rev Mol Cell Biol
15:397–410
76. Becker S, Wandel E, Wobus M, Schneider R,
Amasheh S, Sittig D, Kerner C, Naumann R,
Hamann J, Aust G (2010) Overexpression of
CD97 in intestinal epithelial cells of transgenic mice attenuates colitis by strengthening
adherens junctions. PLoS One 5:e8507
272
Elsa Bazellie ` res and Andre ´ Le Bivic