Sandra Citi



The tight junction protein cingulin regulates the vascular response to burn injury in a mouse model. Zhuravleva K, Goertz O, Wölkart G, Guillemot L, Petzelbauer P, Lehnhardt M, Schmidt K, Citi S, Schossleitner K. Microvasc Res. 2020 Aug.

Scaffolding proteins of vertebrate apical junctions: structure, functions and biophysicsRouaud F., Sluysmans S., Flinois A., Shah J., Vasileva E., Citi S. BBA. October 2020. REVIEW

Cooperative binding of the tandem WW domains of PLEKHA7 to PDZD11 promotes conformation-dependent interaction with tetraspanin 33. Rouaud F., Tessaro F. , Aimaretti L. , Scapozza L., Citi S. J. Biol Chem. May 2020


The mechanobiology of tight junctions. Citi SBiophys Rev. 2019 Oct 4. REVIEW

R40.76 binds to the α domain of ZO-1: role of ZO-1 (α+) in epithelial differentiation and mechano-sensing. Rouaud F, Vasileva E, Spadaro D, Tsukita S, Citi S. Tissue Barriers. 2019 Aug 22

LncRNA EPR controls epithelial proliferation by coordinating Cdkn1a transcription and mRNA decay response to TGF-β. Rossi M, Bucci G, Rizzotto D, Bordo D, Marzi MJ, Puppo M, Flinois A, Spadaro D, Citi S, Emionite L, Cilli M, Nicassio F, Inga A, Briata P, Gherzi R. Nat Commun. 2019 Apr 29;


A Dock-and-Lock Mechanism Clusters ADAM10 at Cell-Cell Junctions to Promote α-Toxin Cytotoxicity. Shah J, Rouaud F, Guerrera D, Vasileva E, Popov LM, Kelley WL, Rubinstein E, Carette JE, Amieva MR, Citi S. Cell Rep. 2018 Nov 20

The role of microtubules in the regulation of epithelial junctions. Vasileva E, Citi S. Tissue Barriers. 2018 Nov 5. REVIEW

Intestinal barriers protect against disease. Citi S. Science. 2018 Mar 9. REVIEW


Tension-Dependent Stretching Activates ZO-1 to Control the Junctional Localization of Its Interactors. Spadaro D, Le S, Laroche T, Mean I, Jond L, Yan J, Citi S. Curr Biol. 2017 Dec 18

Cell-specific diversity in the expression and organization of cytoplasmic plaque proteins of apical junctions. Vasileva E, Sluysmans S, Bochaton-Piallat ML, Citi S. Ann N Y Acad Sci. 2017 Oct

The role of apical cell-cell junctions and associated cytoskeleton in mechanotransduction. Sluysmans S, Vasileva E, Spadaro D, Shah J, Rouaud F, Citi S. Biol Cell. 2017 Apr. REVIEW


Grete Kellenberger-Gujer: Molecular biology research pioneer. Citi S, Berg DE. Bacteriophage. 2016 Apr 5. REVIEW

Role of Cingulin in Agonist-induced Vascular Endothelial Permeability. Tian Y, Gawlak G, Tian X, Shah AS, Sarich N, Citi S, Birukova AA. J Biol Chem. 2016 Nov 4

Cingulin and actin mediate midbody-dependent apical lumen formation during polarization of epithelial cells. Mangan AJ, Sietsema DV, Li D, Moore JK, Citi S, Prekeris R. Nat Commun. 2016 Aug 3

PLEKHA7: Cytoskeletal adaptor protein at center stage in junctional organization and signaling. Shah J, Guerrera D, Vasileva E, Sluysmans S, Bertels E, Citi S. Int J Biochem Cell Biol. 2016 Jun. REVIEW

PLEKHA7 Recruits PDZD11 to Adherens Junctions to Stabilize Nectins. Guerrera D, Shah J, Vasileva E, Sluysmans S, Méan I, Jond L, Poser I, Mann M, Hyman AA, Citi S. J Biol Chem. 2016 May 20

Evidence That Cingulin Regulates Endothelial Barrier Function In Vitro and In Vivo. Schossleitner K, Rauscher S, Gröger M, Friedl HP, Finsterwalder R, Habertheuer A, Sibilia M, Brostjan C, Födinger D, Citi S, Petzelbauer P. Arterioscler Thromb Vasc Biol. 2016 Apr


The adherens junctions control susceptibility to Staphylococcus aureus α-toxin. Popov LM, Marceau CD, Starkl PM, Lumb JH, Shah J, Guerrera D, Cooper RL, Merakou C, Bouley DM, Meng W, Kiyonari H, Takeichi M, Galli SJ, Bagnoli F, Citi S, Carette JE, Amieva MR. Proc Natl Acad Sci U S A. 2015 Nov 17

Distinct E-cadherin-based complexes regulate cell behaviour through miRNA processing or Src and p120 catenin activity. Kourtidis A, Ngok SP, Pulimeno P, Feathers RW, Carpio LR, Baker TR, Carr JM, Yan IK, Borges S, Perez EA, Storz P, Copland JA, Patel T, Thompson EA, Citi S, Anastasiadis PZ. Nat Cell Biol. 2015 Sep;17

The Expression of the Zonula Adhaerens Protein PLEKHA7 Is Strongly Decreased in High Grade Ductal and Lobular Breast Carcinomas. Tille JC, Ho L, Shah J, Seyde O, McKee TA, Citi S.PLoS One. 2015 Aug 13


Epithelial junctions and Rho family GTPases: the zonular signalosome. Citi S, Guerrera D, Spadaro D, Shah J. Small GTPases. 2014. REVIEW

ZO proteins redundantly regulate the transcription factor DbpA/ZONAB. Spadaro D, Tapia R, Jond L, Sudol M, Fanning AS, Citi S. J Biol Chem. 2014 Aug 8

PLEKHA7 modulates epithelial tight junction barrier function. Paschoud S, Jond L, Guerrera D, Citi S. Tissue Barriers. 2014 Jan 1

Cingulin is dispensable for epithelial barrier function and tight junction structure, and plays a role in the control of claudin-2 expression and response to duodenal mucosa injury. Guillemot L, Schneider Y, Brun P, Castagliuolo I, Pizzuti D, Martines D, Jond L, Bongiovanni M, Citi S. J Cell Sci. 2012 Nov 1

The control of gene expression and cell proliferation by the epithelial apical junctional complex. Spadaro D, Tapia R, Pulimeno P, Citi S. Essays Biochem. 2012. REVIEW

Cingulin, paracingulin, and PLEKHA7: signaling and cytoskeletal adaptors at the apical junctional complex. Citi S, Pulimeno P, Paschoud S. Ann N Y Acad Sci. 2012 Jun. REVIEW

Regulation of small GTPases at epithelial cell-cell junctions. Citi S, Spadaro D, Schneider Y, Stutz J, Pulimeno P. Mol Membr Biol. 2011 Oct-Nov. REVIEW

Cingulin and paracingulin show similar dynamic behaviour, but are recruited independently to junctions. Paschoud S, Yu D, Pulimeno P, Jond L, Turner JR, Citi S. Mol Membr Biol. 2011 Feb


PLEKHA7 is an adherens junction protein with a tissue distribution and subcellular localization distinct from ZO-1 and E-cadherin. Pulimeno P, Bauer C, Stutz J, Citi S. PLoS One. 2010 Aug 20


Before 2010

The tight junction protein cingulin regulates gene expression and RhoA signaling.  Citi S., Paschoud S., Pulimeno P., Timolati F., De Robertis F., Jond L., Guillemot L. Ann. NY Acad. Sci. (2009) 1165:88-98

Inducible overexpression of cingulin in stably transfected MDCK cells does not affect tight junction organization and gene expression. Paschoud, S., Citi, S.  Mol. Membr. Biol (2008) 25:1-13

Paracingulin regulates the activity of Rac1 and RhoA GTPases by recruiting Tiam1 and GEF-H1 to epithelial junctions. Guillemot, L., Paschoud, S., Jond, L., Foglia, A., Citi, S.  Mol. Biol. Cell (2008) 19:4442-4453

 The cytoplasmic plaque of tight junctions: A scaffolding and signalling center.  Guillemot, L., Foglia, A., Paschoud, S., Pulimeno, P., Citi, S.   Biochim Biophys Acta (2008) 1778(3):601-13.

 Tight junction formation in early Xenopus laevis embryos: identification and ultrastructural characterization of junctional crests and junctional vesicles.  Cardellini, P., Cirelli A., Citi, S. Cell Tissue Res. (2007) 330:247-56.

Claudin-1 and claudin-5 expression patterns differentiate lung squamous cell carcinomas from adenocarcinomas.  Paschoud, S., Bongiovanni M., Pache J.C., Citi, S. Mod. Pathol. (2007) 20: 947-954.

Cingulin regulates claudin-2 expression and cell proliferation through the small GTPase RhoA.  Guillemot, L., Citi, S. Mol. Biol. Cell (2006) 17:3569-3577.

Cingulin, a cytoskeleton-associated protein of the tight junction.  Guillemot, L., Citi, S.  In : Tight Junctions, 4th Edition (2006) (L. Gonzalez-Mariscal, ed.), Landes Bioscience Publishing.

Binding of GEF-H1 to the tight junction-associated adaptor cingulin results in inhibition of Rho signaling and G1/S phase transition.  Aijaz, S. D’Atri, F., Citi, S., Balda, M.S., Matter, K. Dev. Cell (2005) 8: 777-786.

 Disruption of the cingulin gene does not prevent tight junction formation but alters gene expression.  Guillemot, L., Hammar, E., Kaister, C., Ritz, J., Caille, D., Jond, L., Bauer, C., Meda, P., Citi, S.  J. Cell Sci. (2004) 117: 5245-5256.

Histone deacetylase inhibitors up-regulate the expression of tight junction proteins. Bordin, M., D’Atri, F., Guillemot, L. , Citi, S. Mol. Cancer Res. (2004) 2; 692-701.

What’s new in tight junctions. Citi, S. Cell Commun. Adhes. (2003)10: 51-54.

Evidence for a functional interaction between cingulin and ZO-1 in cultured cells. D’Atri, F., Nadalutti, F., Citi, S.  J. Biol. Chem. (2002) 277:27757-27764.

Molecular complexity of vertebrate tight junctions. D’Atri, F., Citi, S. Mol. Membr. Biol. (2002) 19: 103-112.

Cingulin interacts with F-actin in vitro. D’Atri, F., Citi, S. FEBS Lett. (2001) 507:21-24.

Tight junction protein expression in early Xenopus development and protein interaction studies.  Citi, S., D’Atri, F., Cordenonsi, M., Cardellini, P. In “Cell-cell interactions- A Practical Approach” (2001) T.P. Fleming, ed. (series Editor BD Hames), Oxford University Press, Oxford, UK, pp 153-176.

The cytoplasmic plaque proteins of the tight junction.  Citi, S. In “Tight junctions” 2nd ed. (2001), M. Cereijido and J.M. Anderson, eds., pp. 231-264.

Interaction of Junctional Adhesion Molecule with the Tight Junction Components ZO-1, Cingulin, and Occludin. Bazzoni, G., Martinez Estrada O.M., Orsenigo, F., Cordenonsi, M., Citi, S., Dejana, E.  J. Biol. Chem. (2000) 275: 20520-20526.

 Tight junction biogenesis in the early Xenopus embryo. Fesenko, I., Kurth, T., Sheth, B., Fleming, T.P., Citi, S., Hausen, P. Mech. Dev. (2000) 96: 51-65.

Differentiation of the epithelial apical junctional complex during mouse preimplantation development: a role for rab13 in the early maturation of the tight junction. Sheth, B., Fontaine, J., Ponza, E., McCallum, A., Page A,. Citi, S., Louvard, D., Zahraoui, A., Fleming, TP.  Mech. Dev. (2000) 97:93-104.

Human and Xenopus cingulin share a modular organization of the coiled-coil rod domain: predictions for intra- and intermolecular assembly. Citi, S., D’Atri, F., Parry, D.A.D. J.Struct. Biol. (2000) 131: 135-145.

Introduction: Opening Up Tight Junctions.  Citi, S. In Seminars in Cell and Developmental Biology, issue “Tight Junctiuns” S. Citi Ed. (2000) 11(4), pp. 277-279.

Tight junctions Citi, S., Editor  Seminars in Cell and Developmental Biology (2000) Vol. 11, Issue 4 (pp. 277-324).

Experimental colitis increases small intestine permeability in the rat.   Fries, W., Mazzon, E., Squarzoni, S., Martin, A., Martines, D., Micali, A., Sturniolo, G. C., Citi, S., Longo, G. Lab .Invest. (1999) 79: 49-57.

Xenopus laevis occludin: identification of in vitro phosphorylation sites by protein kinase CK2 and association with cingulin. Cordenonsi, M., Turco, F., D’Atri, F., Hammar, E., Martinucci, G., Meggio, F., Citi, S. Eur. J. Biochem.  (1999) 264: 374-384.

Cingulin contains globular and coiled-coil domains and interacts with ZO-1, ZO-2, ZO-3 and myosin.  Cordenonsi, M., D’Atri, F., Hammar, E., Parry, D.A.D., Kendrick-Jones, J., Shore, D., Citi, S. J. Cell Biol. (1999) 147: 1569-1581.

The molecular basis for the structure, function and regulation of tight junctions. Citi, S., Cordenonsi, M.  In: “Advances in Molecular and Cell Biology”, Vol. 28 (1999), JAI Press Inc., Stamford, CT, ed by Bittar, D.R. Garrod,  A.J. North, M.A.S. Chidgey, pp. 203-233.

Cingulin.  Citi, S. In: "Guidebook to the cytoskeletal and motor proteins" (1998), T. Kreis and R. Vale, eds, 2nd edition Oxford Univ. Press.

Tight junction proteins.  Citi, S., Cordenonsi, M. (1998).  Biochim. Biophys. Acta 1448: 1-11.

Hepatocyte tight-junctional permeability is increased in rat experimental colitis.  Lora, L., Mazzon, E., Martines, D., Fries, W., Muraca, M., Martin, A., D'Odorico, A., Naccarato, R., Citi, S. Gastroenterology (1997) 113: 1347-1354.

Occludin dephosphorylation in early development of Xenopus laevis.  Cordenonsi, M., Mazzon, M., De Rigo, L., Baraldo, S., Meggio, F., Citi, S.  J. Cell Sci. (1997) 110: 3131-3139.

Tight junctions in early amphibian development: detection of junctional cingulin from the 2-cell stage and its localization at the boundary of distinct membrane domains in dividing blastomeres in low calcium.  Cardellini, P., Davanzo, G., Citi, S. Dev. Dyn.207 (1996) 104-113.

Vascular smooth muscle cells of H-2Kb-tsA58 transgenic mice. Characterization of cell lines with distinct properties.  Ehler, E., Jat, P.S., Noble, M.D., Citi, S., Draeger, A.  Circulation (1995).92: 3289-3296.  

Phosphorylation of the tight junction protein cingulin and the effect of protein kinase activators and inhibitors in MDCK epithelial cells.  Citi, S., Denisenko, N. J. Cell Sci. (1995) 108: 2917-2926.

Different effects of protein kinase inhibitors on the localization of junctional proteins at cell-cell contact sites. Denisenko, N., Burighel, P., Citi, S. J. Cell Sci. (1994) 107, 969-981.

Cytoskeletal involvement in the modulation of cell-cell junctions by the protein kinase inhibitor H-7. Citi, S.,Volberg, T., Denisenko, N., Bershadsky, A., Geiger, B. J. Cell Sci. (1994) 107, 683-692.

The effect of protein kinase inhibitor H-7 on the contractility, integrity and membrane anchorage of the microfilament system.  Volberg, T., Geiger, B., Citi, S., Bershadsky, A.D.  Cell Motil. Cytosk. (1994) 29, 321-338.

The molecular composition of tight junctions.  Citi, S. In: "Molecular Mechanisms of epithelial cell junctions: from Development to Disease", S. Citi ed. (1994), RG Landes Biomedical Publishers, Austin, TX, pp 83-106.

Molecular Mechanisms of epithelial cell junctions: from Development to Disease Citi, S. (1994) RG Landes Biomedical Publishers, Austin, TX.

Tight junction protein cingulin is expressed by maternal and embryonic genomes during early mouse development. Javed, Q., Fleming, T.P., Hay, M., Citi, S. Development (1993) 117, 1135-1144.

Localization of tight junction protein cingulin is temporally and spatially regulated during early mouse development.  Fleming, T.P., Hay, M., Javed, Q., Citi, S.Development (1993) 117, 1145-1151.

 The molecular organization of tight junctions. Citi, S. J. Cell Biol. (1993) 121, 485-489

 Cingulin. Citi, S. In: "Guidebook to the cytoskeletal and motor proteins", T. Kreis and R. Vale, eds. (1993), Oxford Univ. Press,  pp 225-226.

 Protein kinase inhibitors prevent junction dissociation induced by low extracellular calcium in MDCK epithelial cells. Citi, S. J.Cell Biol. (1992).  117, 169-178.

 Polarized functions and permeability properties of rat epididymal epithelial cells in vitro. Byers, S.W., Citi, S., Anderson, J.M., Hoxter, B. J. Reprod. Fertil. (1992) 95, 385-396.

 Cingulin, a specific protein component of tight junctions, is expressed in normal and neoplastic human epithelial tissues. Citi, S., Amorosi, A., Franconi, F., Giotti, A., Zampi, G.C.  Am. J. Pathol. (1991) 138, 781-789.

 Localization of myosin in the cytoskeleton of brush border cells using monoclonal antibodies and confocal laser-beam scanning microscopy. Citi, S., Kendrick-Jones, J.  Tissue & Cell (1991) 23, 789-799.

 The role of phosphorylation in development of tight junctions in cultured renal epithelial (MDCK) cells.  Nigam, S., Denisenko, N., Rodriguez-Boulan, E., Citi, S.  Biochem. Biophys. Res. Commun. (1991) 181, 548-553. 

Molecular approaches to study tight junctions: characterization of ZO-1 and cingulin.  Citi, S., Kendrick-Jones, J., & Shore, D. In "Form and function in zoology", G. Lanzavecchia and R. Valvassori eds. (1991), Selected Symposia Monographs UZI 5, Mucchi, Modena, pp. 103-121.

Cingulin: characterization and localization. Citi, S., Sabanay, H., Kendrick-Jones, J. & Geiger, B. J. Cell Sci. (1989) 93, 107-122.

Parallel modulation of brush border myosin conformation and enzyme activity induced by monoclonal antibodies. Citi, S., Cross, R.A., Bagshaw, C.R., & Kendrick-Jones, J.  J. Cell Biol. (1989) 109, 549-556.

 ZO-1 and cingulin: tight junction proteins with distinct identities and localizations. Stevenson, B.R., Heintzelman, M.B., Anderson, J.M., Citi, S., Mooseker, M.S.  Am. J.Physiol. (1989) 257, Cell Physiol. 26, C621-C628.

 How phosphorylation controls the self-assembly of vertebrate smooth and nonmuscle myosins. Cross, R.A., Citi, S. & Kendrick-Jones, J.  Biochem. Soc. Trans. (1988) 16, 501-503.

 Cingulin, a new peripheral component of tight junctions. Citi, S., Sabanay, H., Jakes, R., Geiger, B. & Kendrick-Jones, J.  Nature (1988) 333, 272-276.

Brush border myosin filament assembly and interaction with actin investigated with monoclonal antibodies. Citi, S. & Kendrick-Jones, J.  J. Muscle Res. Cell Motil. (1988)  9, 306-319.

 Active site trapping of nucleotide by smooth and nonmuscle myosins.  Cross, R.A., Jackson, A.P., Citi, S., Kendrick-Jones, J., & Bagshaw, C.R. J.Mol. Biol. (1988) 203, 173-181.

 Modulation of cellular morphology and locomotory activity by antibodies against myosin. Hoener, B., Citi, S., Kendrick-Jones, J., & Jockusch, B.M. J. Cell Biol. (1988) 107, 2181-2189.

Monoclonal antibodies as probes to modulate the conformation, filament assembly and enzymatic activities of myosin. Citi, S. & Kendrick-Jones, J. In : Sarcomeric and non-sarcomeric muscles: basic and applied research prospects for the 90's (U. Carraro ed.) (1988), Unipress Padova, Italy, pp. 739-743.

 Studies on the structure and conformation of brush border myosin using monoclonal antibodies. Citi, S. & Kendrick-Jones, J.  Eur. J. Biochem. (1987) 165, 315-325. 

 The polymerisation of vertebrate smooth muscle and nonmuscle myosins. Kendrick-Jones, J., Smith, R., Craig, R., & Citi, S.. J. Mol. Biol. (1987)  198, 241-252.

The effects of light chain phosphorylation and vertebrate skeletal myosin on the stability of nonmuscle myosin filaments.  Citi, S., Smith, R., & Kendrick-Jones, J. J. Mol. Biol. (1987) 198, 253-262.

Regulation of nonmuscle myosin structure and function. Citi, S. & Kendrick-Jones, J. Bioessays  (1987) 7, 155-159.

Regulation of "contraction" in nonmuscle cells: the role of light chain phosphorylation.  Citi, S. & Kendrick-Jones, J. In: Advances in Myochemistry (ed. G. Benzi), J. Libbey Eurotext, Paris (1987), pp 347-348.

Regulation and organization of nonmuscle myosins. Kendrick-Jones, J., Smith, R., Citi, S. In: "Nature and function of cytoskeletal proteins in motility and transport" (Progress in Zoology, Vol. 34, K.E. Wohlfarth-Botterman, ed. (1987)) Fischer-Verlag, Stuttgart, pp. 87-96.

Regulation in vitro of brush border myosin by light chain phosphorylation. Citi, S., Kendrick.Jones, J.. J.Mol. Biol. (1986) 188, 369-382.