In 1955, biologists Britton Chance and G. R. Williams were the first to propose the idea that respiratory enzymes assemble into larger complexes, although the fluid state model remained the standard. However, as early as 1985, researchers had begun isolating complex III/complex IV supercomplexes from bacteria and yeast. Finally, in 2000 Hermann Schägger and Kathy Pfeiffer used Blue Native PAGE to isolate bovine mitochondrial membrane proteins, showing Complex I, III, and IV arranged in supercomplexes.