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WW domain
The WW domain,〔 (also known as the rsp5-domain〔 or WWP repeating motif〔) is a modular protein domain that mediates specific interactions with protein ligands. This domain is found in a number of unrelated signaling and structural proteins and may be repeated up to four times in some proteins.〔〔〔〔 Apart from binding preferentially to proteins that are proline-rich, with particular proline-motifs, ()-P-P-()-Y, some WW domains bind to phosphoserine- phosphothreonine-containing motifs.〔 == Structure and ligands ==
The WW domain is one of the smallest protein modules, composed of only 40 amino acids, which mediates specific protein-protein interactions with short proline-rich or proline-containing motifs.〔 Named after the presence of two conserved tryptophans (W), which are spaced 20-22 amino acids apart within the sequence,〔 the WW domain folds into a meandering triple-stranded beta sheet.〔 The identification of the WW domain was facilitated by the analysis of two splice isoforms of YAP gene product, named YAP1-1 and YAP1-2, which differed by the presence of an extra 38 amino acids. These extra amino acids are encoded by a spliced-in exon and represent the second WW domain in YAP1-2 isoform.〔〔 The first structure of the WW domain was determined in solution by NMR approach.〔 It represented the WW domain of human YAP in complex with peptide ligand containing Proline-Proline-x–Tyrosine (PPxY where x = any amino acid) consensus motif.〔〔 Recently, the YAP WW domain structure in complex with SMAD-derived, PPxY motif-containing peptide was further refined.〔 Apart from the PPxY motif, certain WW domains recognize LPxY motif (where L is Leucine),〔 and several WW domains bind to phospho-Serine-Proline (p-SP) or phospho-Threonine-Proline (p-ST) motifs in a phospho-dependent manner.〔 Structures of these WW domain complexes confirmed molecular details of phosphorylation-regulated interactions.〔〔 There are also WW domains that interact with polyprolines that are flanked by arginine residues or interrupted by leucine residues, but they do not contain aromatic amino acids.〔〔
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