Proposed mechanism of cristae formation induced by the ATP synthase dimers. (A) ATP synthases (yellow) fold and assemble as monomers in the membrane. (B) As the ATP synthase monomers dimerize, they cause a long-range deformation in the membrane that extends up to 40 nm away from the complex. (C) As the dimers encounter each other, the energy gained from reduced membrane curvature drives the dimers to self-assemble into rows. (D) The dimer rows form a macroscopic membrane ridge that primes the inner membrane to fold and as its surface area increases during mitochondrial development; the membrane invaginates exactly at the location of the dimer rows, and cristae are generated. Inner membrane, light blue; outer membrane, gray.