Tag: Eukaryote evolution

The rise of the eukaryotes

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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.

Structure of a typical animal cell
Structure of a typical eukaryote (animal) cell (credit: Wikipedia)

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.

 

Salt and Earth’s atmosphere

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It is widely known that glacial ice contains a record of Earth’s changing atmospheric composition in the form of bubbles trapped when the ice formed. That is fine for investigations going back about a million years, in particular those that deal with past climate change. Obviously going back to the composition of air tens or hundreds of million years ago cannot use such a handy, direct source of data, but has relied on a range of indirect proxies. These include the number of pores or stomata on fossil plant leaves for CO2, variations in sulfur isotopes for oxygen content and so on. Variation over time of the atmosphere’s content of oxygen has vexed geoscientists a great deal, partly because it has probably been tied to biological evolution: forming by some kind of oxygenic photosynthesis and being essential for the rise to dominance of eukaryotic animals such as ourselves. Its presence or absence also has had a large bearing on weathering and the associated dissolution or precipitation of a variety of elements, predominantly iron. Despite progressively more clever proxies to indicate the presence of oxygen, and intricate geochemical theory through which its former concentration can be modelled, the lack of an opportunity to calibrate any of the models has been a source of deep frustration and acrimony among researchers.

Yet as is often said, there are more ways of getting rid of cats than drowning them in butter. The search has been on for materials that trap air in much the same way as does ice, and one popular, if elusive target has been the bubbles in crystals of evaporite minerals. The trouble is that most halite deposits formed by precipitation of NaCl from highly concentrated brines in evaporating lakes or restricted marine inlets. As a result the bubbles contain liquids that do a grand job of preserving aqueous geochemistry but leave a lot of doubt as regards the provenance of gases trapped within them. For that to be a sample of air rather than gases once dissolved in trapped liquid, the salt needs to have crystallized above the water surface. That may be possible if salt forms from brines so dense that crystals are able to float, or perhaps where minerals such as gypsum form as soil moisture is drawn upwards by capillary action to form ‘desert roses’. A multinational team, led by Nigel Blamey of Brock University in Canada, has published results from Neoproterozoic halite whose chevron-like crystals suggest subaerial formation (Blamey, N.J.F. and 7 others, 2016. Paradigm shift in determining Neoproterozoic atmospheric oxygen. Geology, v. 44, p. 651-654). Multiple analyses of five halite samples from an ~815 Ma-old horizon in a drill core from the Neoproterozoic Canning Basin of Western Australia contained about 11% by volume of oxygen, compared with 25% from Cretaceous salt from China, 20% of late-Miocene age from Italy, and 19 to 22% from samples modern salt of the same type.

Salar de Atacama salt flat in the Chilean puna
Evaporite salts in the Salar de Atacama Chile (credit: Wikipedia)

Although the Neoproterozoic result is only about half that present in modern air, it contradicts results that stem from proxy approaches, which suggest a significant rise in atmospheric oxygenation from 2 to about 18% during the younger Cryogenian and Ediacaran Periods of the Neoproterozoic, when marine animal life made explosive developments at the time of repeated Snowball Earth events. Whether or not this approach can be extended back to the Great Oxygenation Event at around 2.3 Ga ago and before depends on finding evaporite minerals that fit stringent criteria for having formed at the surface: older deposits are known even from the Archaean.

Flourishing life during a Snowball Earth period

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That glacial conditions were able to spread into tropical latitudes during the late Neoproterozoic, Cryogenian Period is now well established, as are the time spans of two such events. http://earth-pages.co.uk/2015/05/21/snowball-earth-events-pinned-down/ But what were the consequences for life that was evolving at the time? That something dramatic was occurring is signalled by a series of perturbations in the carbon-isotope composition of seawater. Its relative proportion of 13C to 12C (δ13C) fell sharply during the two main Snowball events and at other times between 850 to 550 Ma. Since 12C is taken up preferentially by living organisms, falls in δ13C are sometimes attributed to periods when life was unusually suppressed. It is certain that the ‘excursions’ indicate that some process(es) must have strongly affected the way that carbon was cycled in the natural world.

English: Earth, covered in ice.
Artist’s impression of a Snowball Earth as it would appear with today’s continental configuration adjacent to the East Pacific Ocean. (Photo credit: Wikipedia)

The further sea ice extended beyond landmasses during Snowball events the more it would reduce the amount of sunlight reaching the liquid ocean and so photosynthesis would be severely challenged. Indeed, if ice covered the entire ocean surface – the extreme version of the hypothesis – each event must have come close to extinguishing life. An increasing amount of evidence, from climate- and oceanographic modelling and geological observation, suggests that a completely icebound Earth was unlikely. Nevertheless, such dramatic climate shifts would have distressed living processes to the extent that extinction rates were high and so was adaptive radiation of survivors to occupy whatever ecological niches remained or came into being: evolution was thereby speeded up. The roughly half-billion years of the Neoproterozoic hosted the emergence and development of multicellular organisms (metazoan eukaryotes) whose cells contained a nucleus and other bodies such as mitochondria and the chloroplasts of photosynthesisers. This hugely important stage of evolution burst forth shortly after – in a geological sense – the last Snowball event, during the Ediacaran and the Cambrian Explosion. But recent investigations by palaeontologists in glaciogenic rocks from China unearthed a rich diversity of fossil organisms that thrived during a Snowball event (Ye, Q. et al. 2015. The survival of benthic macroscopic phototrophs on a Neoproterozoic snowball Earth. Geology, v. 43, p. 507-510).

The Nantuo Formation in southern China contains glaciogenic sedimentary rocks ascribed to the later Marinoan glaciation (640 to 635 Ma). Unusually, the pebbly Nantuo glaciogenic rocks contain thin layers of siltstones and black shales. The fact that these layers are free of coarse fragments that floating ice may have dropped supports the idea that open water did exist close to glaciated landmasses in what is now southern China. Palaeomagnetic measurements show that the area was at mid-latitudes during the Marinoan event. The really surprising feature is that they contain abundant, easily visible fossils in the form of carbonaceous ribbons , disks, branching masses and some that dramatically resemble complex multi-limbed animals, though they are more likely to be part of an assemblage of algal remains. Whatever their biological affinities, the fossils clearly signify that life happily flourished beneath open water where photosynthesis provided a potential base to a food chain, though no incontrovertible animals occur among them.

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See also: Corsetti, F.A. 2015. Live during Neoproterozoic Snowball Earth. Geology, v. 43, p. 559-560.