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Heparan sulphate-binding sites in the midkine (MK) dimer. In the head-to-head midkine dimer, two CW motifs (+++) form an extended heparan sulphate (HS)-binding site at the dimer interface; in each monomer, the CW motif slopes towards three additional basic residues that further enhance HS affinity. Cell surface HS stabilizes the midkine dimer and is essential for midkine signalling.
