Patient Scientists Elucidate Origin of Eukaryotes

Guest post by John Tillman

In 1962 paper, Roger Stanier and C. B. van Niel established the division of cellular organization into prokaryotes and eukaryotes, defining prokaryotes as those organisms, such as bacteria, which lack a cell nucleus.

Ernst Mayr (1904-2005) called the evolution of eukaryotes–cells with nuclei–“perhaps the most important and dramatic event in the history of life”. Eukaryotic cells also contain various organelles, such as mitochondria, their powerhouses, and in plants and algae, photosynthetic chloroplasts, which use sunlight to make sugar from water and carbon dioxide. This momentous milestone probably occurred between 1.8 and 2.2 billion years ago, but possibly longer.

In 1966, Lynn Margulis (1938-2011), the first Mrs. Carl Sagan, proposed that eukaryotic cells resulted from endosymbiosis, via engulfment of the ancestor of mitochondria by another prokaryote. Experimental evidence for this hypothesis came in 1978, when Robert Schwartz and Margaret Dayhoff demonstrated the descent of mitochondria from bacteria and of chloroplasts from cyanobacteria. During the 1980s, the DNA of mitochondria and chloroplasts was found to differ from their host’s nuclear genetic material, which validated endosymbiosis as a real evolutionary process.

Meanwhile, in 1977, Carl Woese (1928-2012) and George Fox defined Archaea as a third domain of life, with fellow prokaryotic Bacteria and with the much more complex Eukaryota.

In 1999, M. W. Gray, et. al., found strong phylogenetic evidence that an alphaproteobacterium was the ancestor of the mitochondrion. This left open the question of what kind of prokaryote engulfed the first proto-mitochondrion. Similarities in the membranes of archaea and eukaryotes suggested that the endosymbiosis wasn’t between two bacteria, but was a union of two separate domains. If so, then phylogenetically, there were arguably only two domains, not three, since the nuclear DNA of us eukaryotes descends from archaea.

In 2017, various researchers identified the Asgard superphylum as the closest archaean relatives of eukaryotes, but this group was known only from its DNA, recovered from seafloor sediments. Without knowing what Asgard archaeans even look like, let alone their behavior, scientists couldn’t shed light on how endosymbiotic events might have happened.

But now long, hard work by real scientists has helped to unravel this mystery. Thanks to the remarkable persistence and painstaking practice of Japanese microbiologists, science now knows enough about one member of the Asgard superphylum to form an educated hypothesis regarding ancient endosymbiosis and the origin on unicellular eukaryotes and their multicellular descendants, ie animals, fungi and plants.

Please read this Science article for the fascinating details:

https://www.sciencemag.org/news/2019/08/tentacled-microbe-could-be-missing-link-between-simple-cells-and-complex-life

This discovery offers another instance of the importance to life in general of all kinds of symbiosis. We eukaryotes swim in a sea of microbes, most of which aren’t parasites or pathogens. Many make our lives possible. Even the simplest animals–sponges–often form symbiotic relationships with oxygen-producing cyanobacteria. Similarly, lichen are mutualistic partnerships between fungi and algae or cyanobacteria. “Coral symbiosis is a three-player game”:

https://www.nature.com/articles/d41586-019-00949-6

While H. sapiens relies less on gut prokaryotes than do termites, ruminants and other animals needing to break down cellulose, our bodies still contain about 1.4 microbes for each human cell.

Gut prokaryotes: not just for methane any more!

[Edit Addendum sent by John Tillman.

Might add that it appears that eukaryotes developed only once.  Soon after the origin of eukaryotes, sexual reproduction evolved, perhaps in response to reactive oxygen species produced by pre-mitochondria.  Since both photosynthetic and heterotrophic eukaryotes have sex, this development seems to have occurred after proteobacteria (a major gram-negative phylum) were engulfed to become pre-mitochondria but before the inclusion of cyanobacteria as pre-chloroplasts.
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chadb
August 13, 2019 4:46 am

Simple question. I see a lot of comments from people educated in evolutionary biology, so please forgive my ignorance.
What is the point of DNA without DNA polymerase ? Did DNA and DNA polymerase spontaneously co-evolve, and if so where did the code for DNA polymerase come from?
The reason I ask is that DNA polymerase seems to be a fairly complicated protein and it seems essential to any sort of DNA based life. However, in order for an organism to build DNA polymerase it has to have the blueprint in the DNA itself, so did DNA polymerase exist first, or did the code to build DNA polymerase slowly evolve over time, and if it evolved how in the world did any organism actually reproduce without DNA polymerase ?
I have no background in evolutionary biology, so I find this chicken-egg problem to be a bit intractable.

John Tillman
Reply to  chadb
August 13, 2019 9:42 am

I was privileged to know Arthur Kornberg, discoverer of DNA polymerase, and his recombinant DNA colleague, Joshua Lederberg, the smartest person I’ve ever met, who got a Nobel for his PhD. thesis, typed by his wife.

Various versions of DNA and RNA polymerase have evolved. It appears that the RNA egg evolved before the DNA polymerase chicken. RNA is both a genetic information storehouse and enzyme capable of catalyzing its own replication. Scientists are investigating how replicated RNA strands might separate without a biological enzyme. The first polymerase might have been a peptide coded for by an early RNA strand.

chadb
Reply to  John Tillman
August 13, 2019 11:17 am

I get that, but somehow RNA morphed into DNA which is only useful if DNA polymerase exists. What I was asking about was not the RNA side, but the DNA side. Were molecules of DNA polymerase floating around before RNA morphed into DNA, or did the DNA polymerase form only after a need arose? This doesn’t make any sense to me at all. Infact, it is confusing enough that I am fully aware I am having trouble forming a coherent question. Even so I will try because I really am interested. (In order to provide a little clarity I will use “DNA polymerase” as a generic term even though there are several forms, and I mean it as distinct from RNA polymerase. I will also use the term “organism” to refer to anything capable of reproducing whether or not it has organelles and a cell wall) It seems like there are a few options
a) An instruction set for forming DNA polymerase evolved on early strands of RNA. This seems ridiculous because instructions for DNA polymerase can serve no function to an RNA based organism.
b) An RNA organism evolved an ability to build DNA based off of the RNA molecule. DNA polymerase then later evolved. This seems even more absurd since the DNA would be non-functioning until the later DNA polymerase evolved
c) An RNA organism evolved a DNA polymerase like molecule that was able to both form DNA and also read it back. That molecule later evolved into DNA polymerase. While this seems feasible it assumes the first evolutionary step was the most complicated – resulting in a molecule that is even more complicated than current DNA polymerase. That would go in the exact opposite direction that evolution is supposed to run (gradual process of small changes accumulating over time).
I’m not seeing an option d, and a, b, and c all appear to fundamentally disagree with the tenets of random mutation and natural selection. What is the valid option d, or must we swallow that the process of RNA to DNA involved forming DNA polymerase in a single evolutionary step?

John Tillman
Reply to  chadb
August 13, 2019 12:34 pm

Lots of papers have been written on the origin of polymerases and their subsequent evolution in viruses and the three domains of cellular life.

Since RNA appears to have preceded double-stranded DNA as an informational macromolecule, RNA polymerase is the place to start. It is hypothesized to have been among the first proteins to evolve. For over 30 years, scientists have suggested that an important vestige of the original enzyme is found in the contemporary bacterial beta’ subunit of DNA-dependent RNA polymerase and its homologues among archaean and eukaryotic enzymes.

https://www.ncbi.nlm.nih.gov/pubmed/3146647

When DNA started replacing RNA as the informational macromolecule, it might have gotten by with a version of RNA polymerase until natural selection improved upon its function, leading to the complex situation today, with RNA-dependent RNA polymerases, RNA-dependent DNA polymerases, DNA-dependent RNA polymerases and DNA-dependent DNA polymerases in bewildering abundance. Variants look to have been transmitted horizontally across domains and from viruses as well as inherited with mutations.

Reverse transcriptases could shed light on the origin of DNA polymerases. While RTs are enzymes of various uses, their main function is as RNA-directed DNA polymerases in first-strand complementary DNA synthesis.

https://www.ncbi.nlm.nih.gov/pubmed/18972389

I hope this helps.

John Tillman
Reply to  chadb
August 13, 2019 5:51 pm

DNA developed in the RNA world, in which RNA polymerases already existed. They would have worked well enough for DNA still to have supplanted RNA in the bioinformation role.

Then natural selection would have worked its wonders to improve RNA-dependent DNA polymerases into the bewildering variety now on offer, to include DNA-dependent RNA and DNA polymerases.

Phil Salmon
August 13, 2019 8:08 am

Excellent and illuminating article, thanks!

Endosymbiosis between different domains!
No wonder it happened only once.

So we’re all archaeans…

John Tillman
Reply to  Phil Salmon
August 13, 2019 11:00 am

So it seems.

You’re welcome, and many thanks to you and all commenters.

I’m glad we managed to keep the discussion civil.

Young
August 13, 2019 9:04 pm

Rather than count galaxies to guess at the chance that life can arise elsewhere, I wonder if it might be easier to look to its development here.

Life appeared almost as soon as the planet cooled enough for life to exist.

That suggests, but does not prove, that the appearance of life on other planets is likely.

John Tillman
Reply to  Young
August 14, 2019 7:50 am

True.

We’ll find out soon whether there is or was life on other worlds in the solar system.