G. E. Davies coauthored a 1970 method for investigating protein quaternary structure through protein cross-linking. With G. R. Stark, G. E. Davies used dimethyl suberimidate, a reagent with two reactive ends, to join nearby protein subunits before analysing the resulting products.[ref]
SDS-PAGE resolved products with masses equal to multiples of the individual protein subunit mass. A series of linked products could therefore reveal an assembly's subunit organization after treatment that would otherwise separate its components.[ref]
For an invented three-subunit assembly with per subunit, partial linking could produce species near , and . Seeing only the middle species would not establish that the original assembly contained two subunits: the completeness and selectivity of the linking reaction matter.

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