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Two NMR observables, the N ζ H 3  + peak in the HISQC spectrum and N ζ chemical shift difference between the free and heparin-bound forms, can identify binding-interface lysines in protein-heparin complexes. Unlike backbone chemical shifts, these direct probes are stringent and are less prone to either false positives or false negatives.

Direct detection of lysine side chain NH3+ in protein–heparin complexes using NMR spectroscopy

Direct detection of lysine side chain NH₃⁺ in protein–heparin complexes using NMR spectroscopy