Trying to understand how the complex eukaryotic cell evolved is not fully understood. How did a simple prokaryote cell with no membrane-enclosed organelles give rise to cells that contained mitochondria, chloroplasts and a nucleus? One possible answer for this is the viral eukaryogenesis (VE) hypothesis.
According to the VC hypothesis the mitochondria was believed to have originally been a free-living marine alpha-proteobacterium. Possibly 2 billion years ago, this alpha-proteobacterium found its way into a larger archaeon. Whether this was an infection of the archaeon or the archaeon cell consuming the alpha-proteobacterium via phagocytosis remains unclear. In either case, the cells then developed an evolutionary symbiosis. The same hypothesis explains the rise of chloroplasts having come from an engulfed cyanobacterium and eventually gave rise to algae and then plants. What remains less understood is the sudden development of a nucleus within a cell. The VC hypothesis tries to explain this as being the result of a virus that infected the cell but instead of killing the cell became an essential part of it.
In 2003, a new virus was described that infects the genus of amoebae called Acanthamoeba. What is special about this virus is its giant size, both physically (750nm) and the size of its double-stranded DNA genome (1.2Mb) (Raoult et al, 2004). This is actually bigger than some marine bacteria. These viruses are now called mimiviruses (Mimiviridae) and have been given the nickname ‘giruses’.
In this study, an ancestor of the mimivirus is believed to be the origin of the eukaryotic nucleus for several reasons. Just like a eukaryotic nucleus the mimivirus genome is double-stranded, there are repetitive base sequences at their telomeres and have DNA polymerases that are homologous to eukaryotes but not prokaryotes. Mimiviruses also contain enzymes that are used in the capping of mRNA to protect it from degradation inside the host’s cytoplasm. This is not seen in prokaryotic cells but is in eukaryotic nuclei. This also means that once the viral capsid is inside the host cell it is able to carry out DNA replication and transcription independently. Although it still relies on the cell for translation and energy production.
Bell (2009) suggests that like some other viruses, the mimivirus causes the cell membrane to pinch off internal vesicles that is then packaged with DNA to produce new virions. Possibly, as mimivirus are so big, the host genetic material started to get packaged into these vesicles too. The virus remained in the cell and took on the role of DNA storage instead.
The VC hypothesis was originally suggested back in 2001 but with the discovery of the mimivirus a couple of years later this hypothesis had even more evidence to back it up. It is also an expansion on the endosymbiotic theory which is an older and better known theory. It is still just a theory of coarse but I think one that has a lot of strong evidence to back it up. It will be very interesting to see if there are more breakthroughs in this field in the near future.
A Review of:
Bell, P. J. L. (2009) The Viral Eukaryogenesis Hypothesis: A Key Role for Viruses in the Emergence of Eukaryotes from a Prokaryotic World Environment. Ann. NY Acad. Sci, 1178, 91-105.
Additional References:
Raoult, D. Audic, S. Robert, C. Abergel, C. Renesto, P. Ogata, H. La Scola, B. Suzan, M and Claverie, J (2004) The 1.2-Megabase Genome Sequence of Mimivirus. Science, 306, 1344-1350.
9 comments:
There are very exciting devlopmenst int this field with huge controversy raging. It's the study of marine systems which is providing some of th biggest surprises. Here's a new paper on megaviruses from marine samples http://www.pnas.org/content/108/42/17486
This paper came out earlier in the year - a huge virus infecting one of the commonest marine protista. http://www.ncbi.nlm.nih.gov/m/pubmed/21559486/
And there are other ideas coming from probing GOS data. It will be god to follow up in one of the seminars.
Thanks Colin, I will check them out. Its a fascinating subject. I can see why its so controversial.
Hi Lee,
Interesting review. There seems to be several separate endosymbiotic events that took place in early microbial evolution, evident when comparing structures of the mitochondria and the chloroplasts. Mitochondria is believed to have evolved from alpha-proteobacteria while chloroplast containing organisms have a variety of organelle structures, from the typical inner and outer membrane to some having 3-4 membranes. This opens up the idea of several endosymbiotic events (including eukaryotic cell to eukaryotic cell endosymbiosis and the evolution of hydrogenosomes) which don't seem to be all that uncommon and could still be happening as evolution progresses.
Certain species living today, such as the Pelomyxa amoeba, do not have mitochondria but are in symbiosis with aerobic bacteria that play the same role as the mitochondria. An interesting model for supporters of endosymbiosis.
The points on your review of the mimivirus being the ancestor of the nucleus is very plausible indeed. I wonder whether there is some homology in the DNA sequence of the virus and the DNA that encodes the nucleus of the eukaryotic cell.
Hi Mario
yes there are several genes found in this 'girus' that are homologous to eukaryotes, like the mRNA capping enzyme that adds a 5' cap. The paper by Raoult et al (2004) goes into alot of detail. Accorrding to Bell (2009) this puts the mimivirus right at the base of the eukaryotic phylogenetic tree. Very contraversal!
I wanted to right more about it but my word count was getting abit high
Hey Lee,
This is a really interesting subject and is one of the reasons why we are here today. I was just wondering if you have ever read the book Power, Sex and Suicide by Nick Lane. It's a really interesting book talking about the evolution of mitochondria. It's a good read if you're interested, and it talks about the amoeba but as it was published in 2005 probably not as up to date as the paper you were reading!
Hi Arainna
I have heard of that book, not read it sadly. I think I will give it a try. I would like to learn more about the mitochondria side of things.
Hi Lee!
This paper presents really interesting theories. I like how it explores the basics and beginnings of all life really.
I too have read that book - Power, Sex and Suicide. I believe i still have a copy if you'd like a read. It has a lot of useful aspects, but as a general read, it wasn't a book that i got along with very well.
In addition to your review, I wonder if you know why the virus remained within the cell and became an essential part of it? As this is clearly an unusual path for a virus to take.
Hi Sami
Thanks for the comment. If I could borrow that book please that would be great thanks Sami.
To be honest I am not really sure why the virus suddenly decided to stay in the cell. Like you said unusual why it would do that. I know that under certain conditions, eg change in temp, some bacteriophage viruses enter into a dormant lysogenic state which is when the viral DNA becomes integrated into the host genome but is not expressed. Not until conditions like temp improve again. Maybe something like this happened in responce to environmental changes but did not change back again. Perhaps the virus lost the ability to become lytic again?
Good question Sami.
Colin mentioned looking into this some more for the seminar if your interested :-)
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