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Explore the unique evolutionary event that led to the emergence of eukaryotes in this hour-long colloquium talk. Delve into the singular endosymbiosis between an archaeal host cell and a bacterial endosymbiont that occurred only once in 4 billion years of evolution. Discover how the loss of genes from mitochondria enabled a massive expansion in nuclear genome size and gene expression, granting eukaryotes significantly more energy per gene than prokaryotes. Examine the importance of the core bioenergetic genome retained by mitochondria capable of oxidative phosphorylation in supporting the expanded nuclear genome. Investigate the distinct mechanisms of selection on the mitochondrial genome, including uniparental inheritance. Learn about the evolution of the germline in bilaterians and its relationship to mitochondrial function, including the early sequestration of large oocytes containing thousands of mitochondria. Gain insights from Nick Lane of University College London in this Santa Fe Institute colloquium presented on November 7, 2017.