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Desmosomes: integrators of mechanical integrity in tissues.

The most prominent cell surface attachment site for intermediate filaments (IF) at epithelial cell-cell interfaces is the desmosome (reviewed in Collins and Garrod, 1994; Schmidt et al., 1994; Garrod et al., 1996; Green and Jones, 1996); see Figure 1. By anchoring IF at sites of strong intercellular adhesion, desmosomes create a transcellular network throughout the tissue, and this network is thought to resist forces of mechanical stress. The importance of the IF-cell surface complex has been highlighted by the discovery of mutations in genes encoding IF structural proteins, as well as cell junction proteins that result in blistering diseases of the skin (Steinert and Bale, 1993; Fuchs, 1994; McLean and Lane, 1995). Furthermore, autoimmune antibodies that specifically attack the desmosoma1 cadherin component of desmosomes result in a family of blistering diseases of the skin and oral cavity known as pemphigus (Stanley, 1995). Thus, these cell-cell adhesive structures and their underlying cytoskeletal attachments are clearly important for the integrity of tissues. Desmosomes are related to another class of adhesive junctions called adherens junctions, which mediate attachment of the microfilament cytoskeleton to the cell surface (Kowalczyk and Green, 1996). In adherens junctions, classic calcium-dependent cell-cell adhesion molecules (e.g., E-cadherin) are anchored to the actin cytoskeleton through a complex that includes the armadillo protein family member p-catenin. ,0-catenin binds directly to both E-cadherin and to the actin-associated protein cu-catenin. In desmosomes, two classes of desmosomal cadherins,