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R Gary and A Bretscher
Section of Biochemistry, Molecular and Cell Biology, Cornell University, Ithaca, New York 14853, USA.
Ezrin is a membrane-cytoskeletal linking protein that is concentrated in actin-rich surface structures. It is closely related to the microvillar proteins radixin and moesin and to the tumor suppressor merlin/schwannomin. Cell extracts contain ezrin dimers and ezrin-moesin heterodimers in addition to monomers. Truncated ezrin fusion proteins were assayed by blot overlay to determine which regions mediate self- association. Here we report that ezrin self-association occurs by head- to-tail joining of distinct N-terminal and C-terminal domains. It is likely that these domains, termed N- and C-ERMADs (ezrin-radixin-moesin association domain), are responsible for homotypic and heterotypic associations among ERM family members. The N-ERMAD of ezrin resided within amino acids 1-296; deletion of 10 additional residues resulted in loss of activity. The C-ERMAD was mapped to the last 107 amino acids of ezrin, residues 479-585. The two residues at the C-terminus were required for activity, and the region from 530-585 was insufficient. The C-ERMAD was masked in the native monomer. Exposure of this domain required unfolding ezrin with sodium dodecyl sulfate or expressing the domain as part of a truncated protein. Intermolecular association could not occur unless the C-ERMAD had been made accessible to its N-terminal partner. It can be inferred that dimerization in vivo requires an activation step that exposes this masked domain. The conformationally inaccessible C-terminal region included the F-actin binding site, suggesting that this activity is likewise regulated by masking.
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Y. Doi, M. Itoh, S. Yonemura, S. Ishihara, H. Takano, T. Noda, S. Tsukita, and S. Tsukita Normal Development of Mice and Unimpaired Cell Adhesion/Cell Motility/Actin-based Cytoskeleton without Compensatory Up-regulation of Ezrin or Radixin in Moesin Gene Knockout J. Biol. Chem., January 22, 1999; 274(4): 2315 - 2321. [Abstract] [Full Text] [PDF]
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K Hayashi, S Yonemura, T Matsui, and S Tsukita Immunofluorescence detection of ezrin/radixin/moesin (ERM) proteins with their carboxyl-terminal threonine phosphorylated in cultured cells and tissues J. Cell Sci., January 4, 1999; 112(8): 1149 - 1158. [Abstract] [PDF]
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M Gronholm, M Sainio, F Zhao, L Heiska, A Vaheri, and O Carpen Homotypic and heterotypic interaction of the neurofibromatosis 2 tumor suppressor protein merlin and the ERM protein ezrin J. Cell Sci., January 3, 1999; 112(6): 895 - 904. [Abstract] [PDF]
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Y. K. Tran, O. Bogler, K. M. Gorse, I. Wieland, M. R. Green, and I. F. Newsham A Novel Member of the NF2/ERM/4.1 Superfamily with Growth Suppressing Properties in Lung Cancer Cancer Res., January 1, 1999; 59(1): 35 - 43. [Abstract] [Full Text] [PDF]
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M. Amieva, P Litman, L Huang, E Ichimaru, and H Furthmayr Disruption of dynamic cell surface architecture of NIH3T3 fibroblasts by the N-terminal domains of moesin and ezrin: in vivo imaging with GFP fusion proteins J. Cell Sci., January 1, 1999; 112(1): 111 - 125. [Abstract] [PDF]
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N. Oshiro, Y. Fukata, and K. Kaibuchi Phosphorylation of Moesin by Rho-associated Kinase (Rho-kinase) Plays a Crucial Role in the Formation of Microvilli-like Structures J. Biol. Chem., December 25, 1998; 273(52): 34663 - 34666. [Abstract] [Full Text] [PDF]
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M. L. Borowsky and R. O. Hynes Layilin, A Novel Talin-binding Transmembrane Protein Homologous with C-type Lectins, is Localized in Membrane Ruffles J. Cell Biol., October 19, 1998; 143(2): 429 - 442. [Abstract] [Full Text] [PDF]
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L. Heiska, K. Alfthan, M. Gronholm, P. Vilja, A. Vaheri, and O. Carpen Association of Ezrin with Intercellular Adhesion Molecule-1 and -2 (ICAM-1 and ICAM-2). REGULATION BY PHOSPHATIDYLINOSITOL 4,5-BISPHOSPHATE J. Biol. Chem., August 21, 1998; 273(34): 21893 - 21900. [Abstract] [Full Text] [PDF]
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D. Reczek and A. Bretscher The Carboxyl-terminal Region of EBP50 Binds to a Site in the Amino-terminal Domain of Ezrin That Is Masked in the Dormant Molecule J. Biol. Chem., July 17, 1998; 273(29): 18452 - 18458. [Abstract] [Full Text] [PDF]
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D. R. LaJeunesse, B. M. McCartney, and R. G. Fehon Structural Analysis of Drosophila Merlin Reveals Functional Domains Important for Growth Control and Subcellular Localization J. Cell Biol., June 29, 1998; 141(7): 1589 - 1599. [Abstract] [Full Text] [PDF]
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D. A. Potter, J. S. Tirnauer, R. Janssen, D. E. Croall, C. N. Hughes, K. A. Fiacco, J. W. Mier, M. Maki, and I. M. Herman Calpain Regulates Actin Remodeling during Cell Spreading J. Cell Biol., May 4, 1998; 141(3): 647 - 662. [Abstract] [Full Text] [PDF]
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Y. Fukata, K. Kimura, N. Oshiro, H. Saya, Y. Matsuura, and K. Kaibuchi Association of the Myosin-binding Subunit of Myosin Phosphatase and Moesin: Dual Regulation of Moesin Phosphorylation by Rho-associated Kinase and Myosin Phosphatase J. Cell Biol., April 20, 1998; 141(2): 409 - 418. [Abstract] [Full Text] [PDF]
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S. F. Pietromonaco, P. C. Simons, A. Altman, and L. Elias Protein Kinase C-theta Phosphorylation of Moesin in the Actin-binding Sequence J. Biol. Chem., March 27, 1998; 273(13): 7594 - 7603. [Abstract] [Full Text] [PDF]
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S. Yonemura, M. Hirao, Y. Doi, N. Takahashi, T. Kondo, S. Tsukita, and S. Tsukita Ezrin/Radixin/Moesin (ERM) Proteins Bind to a Positively Charged Amino Acid Cluster in the Juxta-Membrane Cytoplasmic Domain of CD44, CD43, and ICAM-2 J. Cell Biol., February 23, 1998; 140(4): 885 - 895. [Abstract] [Full Text] [PDF]
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T. Matsui, M. Maeda, Y. Doi, S. Yonemura, M. Amano, K. Kaibuchi, S. Tsukita, and S. Tsukita Rho-Kinase Phosphorylates COOH-terminal Threonines of Ezrin/Radixin/Moesin (ERM) Proteins and Regulates Their Head-to-Tail Association J. Cell Biol., February 9, 1998; 140(3): 647 - 657. [Abstract] [Full Text] [PDF]
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R. J. Shaw, M. Henry, F. Solomon, and T. Jacks RhoA-Dependent Phosphorylation and Relocalization of ERM Proteins into Apical Membrane/Actin Protrusions in Fibroblasts Mol. Biol. Cell, February 1, 1998; 9(2): 403 - 419. [Abstract] [Full Text]
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D. Reczek, M. Berryman, and A. Bretscher Identification of EBP50: A PDZ-containing Phosphoprotein that Associates with Members of the Ezrin-Radixin-Moesin Family J. Cell Biol., October 6, 1997; 139(1): 169 - 179. [Abstract] [Full Text] [PDF]
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K. Takahashi, T. Sasaki, A. Mammoto, K. Takaishi, T. Kameyama, S. Tsukita, S. Tsukita, and Y. Takai Direct Interaction of the Rho GDP Dissociation Inhibitor with Ezrin/Radixin/Moesin Initiates the Activation of the Rho Small G Protein J. Biol. Chem., September 12, 1997; 272(37): 23371 - 23375. [Abstract] [Full Text] [PDF]
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C. Roy, M. Martin, and P. Mangeat A Dual Involvement of the Amino-terminal Domain of Ezrin in F- and G-actin Binding J. Biol. Chem., August 8, 1997; 272(32): 20088 - 20095. [Abstract] [Full Text] [PDF]
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T. Crepaldi, A. Gautreau, P. M. Comoglio, D. Louvard, and M. Arpin Ezrin Is an Effector of Hepatocyte Growth Factor-mediated Migration and Morphogenesis in Epithelial Cells J. Cell Biol., July 28, 1997; 138(2): 423 - 434. [Abstract] [Full Text] [PDF]
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A Bretscher, D Reczek, and M Berryman Ezrin: a protein requiring conformational activation to link microfilaments to the plasma membrane in the assembly of cell surface structures J. Cell Sci., January 12, 1997; 110(24): 3011 - 3018. [Abstract] [PDF]
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M Sainio, F Zhao, L Heiska, O Turunen, M den Bakker, E Zwarthoff, M Lutchman, G. Rouleau, J Jaaskelainen, A Vaheri, et al. Neurofibromatosis 2 tumor suppressor protein colocalizes with ezrin and CD44 and associates with actin-containing cytoskeleton J. Cell Sci., January 9, 1997; 110(18): 2249 - 2260. [Abstract] [PDF]
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X. Yao, L. Cheng, and J. G. Forte Biochemical Characterization of Ezrin-Actin Interaction J. Biol. Chem., March 22, 1996; 271(12): 7224 - 7229. [Abstract] [Full Text] [PDF]
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R. Nguyen, D. Reczek, and A. Bretscher Hierarchy of Merlin and Ezrin N- and C-terminal Domain Interactions in Homo- and Heterotypic Associations and Their Relationship to Binding of Scaffolding Proteins EBP50 and E3KARP J. Biol. Chem., March 2, 2001; 276(10): 7621 - 7629. [Abstract] [Full Text] [PDF]
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N. R. Filipenko and D. M. Waisman The C Terminus of Annexin II Mediates Binding to F-actin J. Biol. Chem., February 9, 2001; 276(7): 5310 - 5315. [Abstract] [Full Text] [PDF]
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P. Poullet, A. Gautreau, G. Kadare, J.-A. Girault, D. Louvard, and M. Arpin Ezrin Interacts with Focal Adhesion Kinase and Induces Its Activation Independently of Cell-matrix Adhesion J. Biol. Chem., September 28, 2001; 276(40): 37686 - 37691. [Abstract] [Full Text] [PDF]
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A. Ivetic, J. Deka, A. Ridley, and A. Ager The Cytoplasmic Tail of L-selectin Interacts with Members of the Ezrin-Radixin-Moesin (ERM) Family of Proteins. CELL ACTIVATION-DEPENDENT BINDING OF MOESIN BUT NOT EZRIN J. Biol. Chem., January 11, 2002; 277(3): 2321 - 2329. [Abstract] [Full Text] [PDF]
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