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Ageing is a fundamental yet poorly understood biological process that affects all eukaryotic life. Deterioration in mitochondria is clearly seen in ageing, but details of the underlying molecular events are largely unknown. Cryo-ET of mitochondria from the short-lived (18 days) model organism Podospora anserina revealed profound age-dependent changes in the inner membrane architecture [56]. In normal mitochondria of young cells, the cristae protrude deeply into the matrix. Formation of cristae depends both on the rows of ATP synthase dimers along the edges [30] and on the MICOS complex at the crista junctions [32]. With increasing age, the cristae recede into the inner boundary membrane and the inter-membrane space widens. The MICOS complex most likely has to come apart for this to happen. Eventually, the matrix breaks up into spherical vesicles within the outer membrane. The ATP synthase dimer rows disperse and the dimers dissociate into monomers. As the inner membrane vesiculates, the sharp local curvature at the dimer rows inverts, so that the ATP synthase monomers are surrounded by a shallow concave membrane environment, rather than the sharply convex curvature at the crista ridges (Fig. 9). Finally, the outer membrane ruptures, releasing the inner membrane vesicles, along with apoptogenic cytochrome c, into the cytoplasm. Cytochrome c activates a cascade of proteolytic caspases, which degrade cellular proteins [55]. The cell enters into apoptosis and dies.
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