Indirect readout refers to I. a protein's ability to form hydrogen bonds through a network of water molecules II. ability of a protein to bind both 5' to 3' sequences on the opposite strand of a DNA helix.. III. a protein's ability to recognize a specific DNA sequence by its backbone conformation and flexibility. IV. the protein's ability to make hydrogen bonds with phosphate oxygens O I, IV O I, IV O1, IV 1, II, IV III