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Respiratory electron transport complexes are organized as individual entities or combined as large supercomplexes (SC). Gram-negative bacteria deploy a mitochondrial-like cytochrome (cyt)  bc  1  (Complex III, CIII 2 ), and may have specific  cbb  3 -type cyt  c  oxidases (Complex IV, CIV) instead of the canonical  aa  3 -type CIV. Electron transfer between these complexes is mediated by soluble ( c  2 ) and membrane-anchored ( c  y ) cyts. Here, we report the structure of an engineered  bc  1 - cbb  3  type SC (CIII 2 CIV, 5.2 Å resolution) and three conformers of native CIII 2  (3.3 Å resolution). The SC is active in vivo and in vitro, contains all catalytic subunits and cofactors, and two extra transmembrane helices attributed to cyt  c  y  and the assembly factor CcoH. The cyt  c  y  is integral to SC, its cyt domain is mobile and it conveys electrons to CIV differently than cyt  c  2 . The successful production of a native-like functional SC and determination of its structure illustrate the characteristics of membrane-confined and membrane-external respiratory electron transport pathways in Gram-negative bacteria.

Cryo-EM structures of engineered active bc1-cbb3 type CIII2CIV super-complexes and electronic communication between the complexes