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For many years it was assumed that the ATP synthase and other energy-converting complexes are randomly distributed over the inner membrane. The first hint that this is not the case came from deep-etch freeze-fracture electron microscopy, which revealed double rows of macromolecular complexes in the tubular cristae of the single-cell ciliate Paramecium [37]. The double rows were thought to be linear arrays of mitochondrial ATP synthase. This is indeed what they are, but it could only be shown unambiguously more than 20 years later by cryo-ET [30, 38], which revealed rows of ATP synthase dimers in mitochondria of all species investigated [30] (Fig. 4). Until then, the rows were thought to be peculiar to Paramecium.
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