You and I, and all the living things that we can easily see belong to the most recently evolved of the three great domains of life, the Eukarya. The vast bulk of organisms that we can’t see unaided are prokaryotes, divided into the Bacteria and the Archaea. Their genetic material floats around in their cell’s fluid, while ours resides mainly in the eukaryote cell’s nucleus with a bit in various organelles known as mitochondria and the chloroplasts of plant cells. Unlike the chicken and egg question, that concerning which came first, prokaryotes or eukaryotes, is answered by DNA. Eukaryote DNA contains a lot from prokaryotes, but the converse does not hold. That contrast posed the question of how eukaryotes arose from the two earlier, simpler forms of life, the answer to which Lynn Margulis suggested to be a whole series of symbiotic relationships among various prokaryotes that shared a host cell; her hypothesis of endosymbiosis. Now, the vast majority of eukaryotes depend on free oxygen for their metabolism, so when the first of them arose boils down to the period of geological history following the Great Oxidation Event around 2.4 billion years ago.

Molecular-clock estimates based on the range of variation in the genomes of a wide range of eukaryotes suggest it took place sometime between 1000 and 2000 Ma. A better means of homing in on a date for the Last Eukaryote Common Ancestor (LECA – as opposed to that of the first organism LUCA) would be that of the earliest fossil to show eukaryote affinities. Grypania from 1.85 Ga, a sort of whorl-like fossil, is a good candidate and is widely thought to be the earliest of our kind but lacks signs of actual cells. More convincing fossils – known generically as acritarchs – from times between 1.5 and 1.0 Ga look like primitive fungi, red algae and slime moulds. A comprehensive review of the microfossils of the Palaeoproterozoic (2.5 to 1.6 Ga) includes both prokaryotes and probable early eukaryotes (Javaux, E.J. & Lepot, K. 2017. The Paleoproterozoic fossil record: Implications for the evolution of the biosphere during Earth’s middle-age. Earth Science Reviews, v. 176, p. 68-86; doi: 10.1016/j.earscirev.2017.10.0001). Yet, despite rapidly accumulating evidence, especially from rocks in China, the picture remains one of monotony; for instance Grypania spans the best part of half a billion years. Bacteria and Archaea cannot be distinguished easily in the absence of preserved DNA. Despite evidence for oxygen in the oceans and atmosphere, apart from a few shallow-water oxygenated examples the chemistry of Palaeoproterozoic marine sediments is dominated by mineralogical outcomes of reducing chemistry. Many chemical isotopic environmental proxies ‘flat-line’ to the extent that the early Proterozoic is sometimes referred to as the ‘boring billion’, yet our ultimate precursors were part of the marine ecosystem. That is, unless one accepts the possibility that that fossils labelled ‘eukaryote’ are colonial prokaryotes – evidence for cell nuclei is sparse. Endosymbiosis, although an attractive model for eukaryote origins, is not proven. The reason for lingering scepticism is that there are only a tiny number of modern examples of prokaryote cells ending up inside those of other prokaryotes.

Whatever, chemical biomarkers in sediments older than about 720 Ma indicate that prokaryotes were the only notable primary producers in the oceans until the Neoproterozoic. Microscopic fossils that are inescapably eukaryotes in the form of amoeba suddenly emerge around that time. This development from the lingering marginality of early eukaryotes to thriving ecosystems that they dominated thereafter is a puzzle seeking a plausible explanation. It coincides with the onset of the Snowball Earth glaciations of the Cryogenian Period (850 to 635 Ma) and a rise in atmospheric and presumably oceanic oxygen. Then macroscopic eukaryotes ‘bloomed’ into distinctively different forms in the Ediacaran Period (635 to 541 Ma) and thereafter. Before the Cryogenian we can perhaps regard eukaryan life and the endosymbiosis that may have given rise to it as a series of ecological experiments repeatedly knocked-back by chemical conditions and competition with the vastly more abundant prokaryotes.