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Through mutagenesis, we have identified a single high affinity binding site for gp120 on the human CD4 protein. This site is localized within the V1 domain within residues 41-55. This same collection of mutants was used to partially define the epitopes for 23 CD4 monoclonal antibodies. The locations of these epitopes are consistent with a V-like structure for the V1 domain, as orginally suggested by shared sequence homology. In the context of this structure, the gp120 binding site encompasses the small CDR2 loop.