Intramolecular covalent linkages in Gram-positive pili and MSCRAMMs. (A) Lys-Asn intramolecular isopeptide bond in Spy0128, (B) Thr-Gln ester bond in Cpe0147, and (C) Internal thioester in Spy0125. The side chains of residues are colored: yellow, residues forming the bonds; cyan, catalytic residues in Cpe0147; green, residues contributing to the proton shuttles. In orange are parts of the β-strands that are covalently linked through the bonds.