J. Cell Biol., Volume 145, Number 4, May 17, 1999 851-863

Paxillin LD4 Motif Binds PAK and PIX through a Novel 95-kD Ankyrin Repeat, ARF-GAP Protein: A Role in Cytoskeletal Remodeling

Christopher E. Turner,^* Michael C. Brown,^* Joseph A. Perrotta,^* M.C. Riedy,^* Sotiris N. Nikolopoulos,^* A. Rosa McDonald,^* Shubha Bagrodia, Sheila Thomas,^§ and Phillip S. Leventhal^*

^* Department of Anatomy and Cell Biology, State University of New York, Health Science Center, Syracuse, New York 13210;  Department of Molecular Medicine, Cornell University, Ithaca, New York 14853; and ^§ Department of Medicine, Beth Israel Deaconess Medical Center, Harvard Medical School, Boston, Massachusetts 02215

Paxillin is a focal adhesion adaptor protein involved in the integration of growth factor- and adhesion-mediated signal transduction pathways. Repeats of a leucine-rich sequence named paxillin LD motifs (Brown M.C., M.S. Curtis, and C.E. Turner. 1998. Nature Struct. Biol. 5:677-678) have been implicated in paxillin binding to focal adhesion kinase (FAK) and vinculin. Here we demonstrate that the individual paxillin LD motifs function as discrete and selective protein binding interfaces. A novel scaffolding function is described for paxillin LD4 in the binding of a complex of proteins containing active p21 GTPase-activated kinase (PAK), Nck, and the guanine nucleotide exchange factor, PIX. The association of this complex with paxillin is mediated by a new 95-kD protein, p95PKL (paxillin-kinase linker), which binds directly to paxillin LD4 and PIX. This protein complex also binds to Hic-5, suggesting a conservation of LD function across the paxillin superfamily. Cloning of p95PKL revealed a multidomain protein containing an NH₂-terminal ARF-GAP domain, three ankyrin-like repeats, a potential calcium-binding EF hand, calmodulin-binding IQ motifs, a myosin homology domain, and two paxillin-binding subdomains (PBS). Green fluorescent protein- (GFP-) tagged p95PKL localized to focal adhesions/complexes in CHO.K1 cells. Overexpression in neuroblastoma cells of a paxillin LD4 deletion mutant inhibited lamellipodia formation in response to insulin-like growth fac- tor-1. Microinjection of GST-LD4 into NIH3T3 cells significantly decreased cell migration into a wound. These data implicate paxillin as a mediator of p21 GTPase-regulated actin cytoskeletal reorganization through the recruitment to nascent focal adhesion structures of an active PAK/PIX complex potentially via interactions with p95PKL.

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• Huang, C.-F., Chen, C.-C., Tung, L., Buu, L.-M., Lee, F.-J. S. (2002). The yeast ADP-ribosylation factor GAP, Gcs1p, is involved in maintenance of mitochondrial morphology. J. Cell Sci. 115: 275-282 [Abstract] [Full Text]  
• Manabe, R.-i., Kovalenko, M., Webb, D. J., Horwitz, A. R. (2002). GIT1 functions in a motile, multi-molecular signaling complex that regulates protrusive activity and cell migration. J. Cell Sci. 115: 1497-1510 [Abstract] [Full Text]  
• Szafer, E., Rotman, M., Cassel, D. (2001). Regulation of GTP Hydrolysis on ADP-ribosylation Factor-1 at the Golgi Membrane. J. Biol. Chem. 276: 47834-47839 [Abstract] [Full Text]  
• Jia, Y., Ransom, R. F., Shibanuma, M., Liu, C., Welsh, M. J., Smoyer, W. E. (2001). Identification and Characterization of hic-5/ARA55 as an hsp27 Binding Protein. J. Biol. Chem. 276: 39911-39918 [Abstract] [Full Text]  
• Berrou, E., Bryckaert, M. (2001). Platelet-derived Growth Factor Inhibits Smooth Muscle Cell Adhesion to Fibronectin by ERK-dependent and ERK-independent Pathways. J. Biol. Chem. 276: 39303-39309 [Abstract] [Full Text]  
• Vogelezang, M. G., Liu, Z., Relvas, J. B., Raivich, G., Scherer, S. S., ffrench-Constant, C. (2001). {alpha}4 Integrin Is Expressed during Peripheral Nerve Regeneration and Enhances Neurite Outgrowth. J. Neurosci. 21: 6732-6744 [Abstract] [Full Text]  

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