junctional localization of its interactors. Curr
Biol 27:3783–3795.e8
32. Then C, Bergler T, Jeblick R, Jung B,
Banas B, Kr€ amer BK (2011) Hypertonic stress
promotes the upregulation and phosphorylation of zonula occludens 1. Nephron Physiol
119:p11–p21
33. Fanning AS, Ma TY, Anderson JM (2002)
Isolation and functional characterization of
the actin binding region in the tight junction
protein ZO-1. FASEB J 16:1835–1837
34. Fanning AS, Little BP, Rahner C,
Utepbergenov D, Walther Z, Anderson JM
(2007) The unique-5 and -6 motifs of ZO-1
regulate tight junction strand localization and
scaffolding properties. Mol Biol Cell
18:721–731
35. Umeda K, Ikenouchi J, Katahira-Tayama S,
Furuse K, Sasaki H, Nakayama M, Matsui T,
Tsukita S, Furuse M, Tsukita S (2006) ZO-1
and ZO-2 independently determine where
claudins are polymerized in tight-junction
strand formation. Cell 126:741–754
36. Ikenouchi J, Umeda K, Tsukita S, Furuse M,
Tsukita S (2007) Requirement of ZO-1 for
the formation of belt-like adherens junctions
during epithelial cell polarization. J Cell Biol
176:779–786
37. Rodgers LS, Beam MT, Anderson JM, Fanning AS (2013) Epithelial barrier assembly
requires coordinated activity of multiple
domains of the tight junction protein ZO-1.
J Cell Sci 126:1565–1575
38. Odenwald MA, Choi W, Buckley A,
Shashikanth N, Joseph NE, Wang Y, Warren
MH, Buschmann MM, Pavlyuk R,
Hildebrand J, Margolis B, Fanning AS,
Turner JR (2017) ZO-1 interactions with
F-actin and occludin direct epithelial polarization and single lumen specification in 3D culture. J Cell Sci 130:243–259
39. Fanning AS, Anderson JM (2009) Zonula
occludens-1 and -2 are cytosolic scaffolds
that regulate the assembly of cellular junctions. Ann N Y Acad Sci 1165:113–120
40. Ullmer C, Schmuck K, Figge A, Lu ¨bbert H
(1998) Cloning and characterization of
MUPP1, a novel PDZ domain protein.
FEBS Lett 424:63–68
41. Asse ´mat E, Bazellie `res E, Pallesi-Pocachard E,
Le Bivic A, Massey-Harroche D (2008) Polarity complex proteins. Biochim Biophys Acta
1778:614–630
42. Lanaspa M, Andres-Hernando A, Rivard C,
Dai Y, Berl T (2008) Hypertonic stress
increases claudin-4 expression and tight junction integrity in association with MUPP1 in
IMCD3 cells. Proc Natl Acad Sci U S A
105:15797–15802
43. Latorre IJ, Roh MH, Frese KK, Weiss RS,
Margolis B, Javier RT (2005) Viral
oncoprotein-induced mislocalization of select
PDZ proteins disrupts tight junctions and
causes polarity defects in epithelial cells. J
Cell Sci 118:4283–4293
44. Shin K, Straight S, Margolis B (2005) PATJ
regulates tight junction formation and polarity in mammalian epithelial cells. J Cell Biol
168:705–711
45. Straight SW, Shin K, Fogg VC, Fan S, Liu C-J,
Roh M, Margolis B (2004) Loss of PALS1
expression leads to tight junction and polarity
defects. Mol Biol Cell 15:1981–1990
46. Sewduth RN, Kovacic H, Jaspard-Vinassa B,
Jecko V, Wavasseur T, Fritsch N, Pernot M,
Jeaningros S, Roux E, Dufourcq P,
Couffinhal T, Dupla `a C (2017) PDZRN3
destabilizes endothelial cell-cell junctions
through a PKCζ-containing polarity complex
to increase vascular permeability. Sci Signal
10:464
47. Be ´camel C, Figge A, Poliak S, Dumuis A,
Peles E, Bockaert J, Lu ¨bbert H, Ullmer C
(2001)
Interaction
of
serotonin
5-hydroxytryptamine type 2C receptors with
PDZ10 of the multi-PDZ domain protein
MUPP1. J Biol Chem 276:12974–12982
48. Fujita E, Tanabe Y, Imhof BA, Momoi MY,
Momoi T (2012) A complex of synaptic adhesion molecule CADM1, a molecule related to
autism spectrum disorder, with MUPP1 in
the cerebellum. J Neurochem 123:886–894
49. Baliova M, Juhasova A, Jursky F (2014) Using
a collection of MUPP1 domains to investigate
the similarities of neurotransmitter transporters C-terminal PDZ motifs. Biochem Biophys Res Commun 454:25–29
50. Wang S-C, Low TYF, Nishimura Y, Gole L,
Yu W, Motegi F (2017) Cortical forces and
CDC-42 control clustering of PAR proteins
for Caenorhabditis elegans embryonic polarization. Nat Cell Biol 19:988–995
51. Munro E, Nance J, Priess JR (2004) Cortical
flows powered by asymmetrical contraction
transport PAR proteins to establish and maintain anterior-posterior polarity in the early
C. elegans embryo. Dev Cell 7:413–424
52. Goehring NW, Hoege C, Grill SW, Hyman
AA (2011) PAR proteins diffuse freely across
the anterior–posterior boundary in polarized
C. elegans embryos. J Cell Biol 193:583–594
53. Goehring NW, Trong PK, Bois JS,
Chowdhury D, Nicola EM, Hyman AA,
Grill SW (2011) Polarization of PAR proteins
Mechanoregulation of PDZ Proteins
271
Biol 27:3783–3795.e8
32. Then C, Bergler T, Jeblick R, Jung B,
Banas B, Kr€ amer BK (2011) Hypertonic stress
promotes the upregulation and phosphorylation of zonula occludens 1. Nephron Physiol
119:p11–p21
33. Fanning AS, Ma TY, Anderson JM (2002)
Isolation and functional characterization of
the actin binding region in the tight junction
protein ZO-1. FASEB J 16:1835–1837
34. Fanning AS, Little BP, Rahner C,
Utepbergenov D, Walther Z, Anderson JM
(2007) The unique-5 and -6 motifs of ZO-1
regulate tight junction strand localization and
scaffolding properties. Mol Biol Cell
18:721–731
35. Umeda K, Ikenouchi J, Katahira-Tayama S,
Furuse K, Sasaki H, Nakayama M, Matsui T,
Tsukita S, Furuse M, Tsukita S (2006) ZO-1
and ZO-2 independently determine where
claudins are polymerized in tight-junction
strand formation. Cell 126:741–754
36. Ikenouchi J, Umeda K, Tsukita S, Furuse M,
Tsukita S (2007) Requirement of ZO-1 for
the formation of belt-like adherens junctions
during epithelial cell polarization. J Cell Biol
176:779–786
37. Rodgers LS, Beam MT, Anderson JM, Fanning AS (2013) Epithelial barrier assembly
requires coordinated activity of multiple
domains of the tight junction protein ZO-1.
J Cell Sci 126:1565–1575
38. Odenwald MA, Choi W, Buckley A,
Shashikanth N, Joseph NE, Wang Y, Warren
MH, Buschmann MM, Pavlyuk R,
Hildebrand J, Margolis B, Fanning AS,
Turner JR (2017) ZO-1 interactions with
F-actin and occludin direct epithelial polarization and single lumen specification in 3D culture. J Cell Sci 130:243–259
39. Fanning AS, Anderson JM (2009) Zonula
occludens-1 and -2 are cytosolic scaffolds
that regulate the assembly of cellular junctions. Ann N Y Acad Sci 1165:113–120
40. Ullmer C, Schmuck K, Figge A, Lu ¨bbert H
(1998) Cloning and characterization of
MUPP1, a novel PDZ domain protein.
FEBS Lett 424:63–68
41. Asse ´mat E, Bazellie `res E, Pallesi-Pocachard E,
Le Bivic A, Massey-Harroche D (2008) Polarity complex proteins. Biochim Biophys Acta
1778:614–630
42. Lanaspa M, Andres-Hernando A, Rivard C,
Dai Y, Berl T (2008) Hypertonic stress
increases claudin-4 expression and tight junction integrity in association with MUPP1 in
IMCD3 cells. Proc Natl Acad Sci U S A
105:15797–15802
43. Latorre IJ, Roh MH, Frese KK, Weiss RS,
Margolis B, Javier RT (2005) Viral
oncoprotein-induced mislocalization of select
PDZ proteins disrupts tight junctions and
causes polarity defects in epithelial cells. J
Cell Sci 118:4283–4293
44. Shin K, Straight S, Margolis B (2005) PATJ
regulates tight junction formation and polarity in mammalian epithelial cells. J Cell Biol
168:705–711
45. Straight SW, Shin K, Fogg VC, Fan S, Liu C-J,
Roh M, Margolis B (2004) Loss of PALS1
expression leads to tight junction and polarity
defects. Mol Biol Cell 15:1981–1990
46. Sewduth RN, Kovacic H, Jaspard-Vinassa B,
Jecko V, Wavasseur T, Fritsch N, Pernot M,
Jeaningros S, Roux E, Dufourcq P,
Couffinhal T, Dupla `a C (2017) PDZRN3
destabilizes endothelial cell-cell junctions
through a PKCζ-containing polarity complex
to increase vascular permeability. Sci Signal
10:464
47. Be ´camel C, Figge A, Poliak S, Dumuis A,
Peles E, Bockaert J, Lu ¨bbert H, Ullmer C
(2001)
Interaction
of
serotonin
5-hydroxytryptamine type 2C receptors with
PDZ10 of the multi-PDZ domain protein
MUPP1. J Biol Chem 276:12974–12982
48. Fujita E, Tanabe Y, Imhof BA, Momoi MY,
Momoi T (2012) A complex of synaptic adhesion molecule CADM1, a molecule related to
autism spectrum disorder, with MUPP1 in
the cerebellum. J Neurochem 123:886–894
49. Baliova M, Juhasova A, Jursky F (2014) Using
a collection of MUPP1 domains to investigate
the similarities of neurotransmitter transporters C-terminal PDZ motifs. Biochem Biophys Res Commun 454:25–29
50. Wang S-C, Low TYF, Nishimura Y, Gole L,
Yu W, Motegi F (2017) Cortical forces and
CDC-42 control clustering of PAR proteins
for Caenorhabditis elegans embryonic polarization. Nat Cell Biol 19:988–995
51. Munro E, Nance J, Priess JR (2004) Cortical
flows powered by asymmetrical contraction
transport PAR proteins to establish and maintain anterior-posterior polarity in the early
C. elegans embryo. Dev Cell 7:413–424
52. Goehring NW, Hoege C, Grill SW, Hyman
AA (2011) PAR proteins diffuse freely across
the anterior–posterior boundary in polarized
C. elegans embryos. J Cell Biol 193:583–594
53. Goehring NW, Trong PK, Bois JS,
Chowdhury D, Nicola EM, Hyman AA,
Grill SW (2011) Polarization of PAR proteins
Mechanoregulation of PDZ Proteins
271