(C) CHO-α4/GFP cells were transfected as indicated and then analyzed in scratch wound assays, as described in the legend to Figure 2, except that 24-well dishes were coated with 10 μg/ml CCBD. (B) CHO-α4/GFP cells were transfected as indicated and then analyzed in scratch wound assays, as described in the legend to Figure 2, except that 24-well dishes were coated with 2 μg/ml CS-1. Similarly, expression of ADAM12 in CHO-α4/GFP cells caused a reduction in cell motility in both transwell (Figure 2C) and scratch wound (Figure 2D) assays.
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To produce protein with higher purity, a 6Xhistidine (His) tag was added to the 3′ end of the disintegrin domain coding sequence, using a site-directed mutagenesis kit (Stratagene, La Jolla, CA). Mouse ADAM12 disintegrin domain in the vector pGEX-2T was provided by Anna Zolkiewski (Kansas State University, Manhattan, KS). Mouse anti-α4 integrin functional blocking monoclonal antibody (mAb) HP2/1 and mouse anti-α5 integrin functional blocking antibody CBL497 were obtained from Chemicon International (Temecula, CA). Many recombinant ADAM disintegrin domains have been shown to interact with a variety of β1 integrins, including α4β1 (Tomczuk et al., 2003; White, 2003; Bridges et al., 2004; White et al., 2005).
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The patterns of inhibition are consistent with the integrin binding profiles of the respective ADAM disintegrin domains on cells with activated β1 integrins (Bax et al., 2004) (Figure 1; our unpublished data). However, the mechanisms by which ADAMs function in cell migration remain unclear, both for ADAMs with (e.g., Xenopus ADAM13) and for those without (e.g., Caenorhabditis elegans ADAM14; unc71) proteolytic activity. ADAM (and ADAM-like) proteins have been implicated in several cell migration events (Nishiwaki et al., 2000; Alfandari et al., 2001; Martin et al., 2002; Huang et al., 2003; Seals and Courtneidge, 2003; White, 2003). As seen in Figure 8B, unlike ADAM12, expression of ADAM19 and 33 (individually) also inhibited α5β1-mediated cell migration. As seen in Figure 8A, expression of both ADAM19 and ADAM33 inhibited α4β1-mediated cell migration.
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For scratch wound assays, 24-well dishes were precoated with 10 μg/ml FN, and migration assays were conducted as described in Materials and Methods. Cells were either mock transfected or transfected to express full-length ADAM12, and transwell migration assays (A, C, E, and G) or scratch wound assays (B, D, F, and H) were conducted 24 h after transfection. Cells expressing α4β1 but not α5β1 integrin bind to recombinant ADAM12 disintegrin domain.