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Deciphering Protein Evolution
The Scientist ^ | Nov 26, 2001 | Barry A. Palevitz

Posted on 01/14/2002 3:02:24 PM PST by Karl_Lembke

Deciphering Protein Evolution

Actin shares a common ancestor with a bacterial protein

By Barry A. Palevitz

One of the enduring questions in biology is how eukaryotic cells arose from prokaryotic ancestors at least 2 billion years ago. Besides differences in genome organization, eukaryotic animals, plants, and fungi possess a much higher degree of cellular compartmentation in the form of membrane bound organelles than their distant bacterial and Archaean cousins. But how did such a plethora of cellular domains, each with a discrete role in metabolism, evolve?

To the extent that science proves anything, it answered the question for two eukaryotic organelles a long time ago. Mitochondria and chloroplasts evolved from endosymbiotic associations between an ancestral host cell and smaller prokaryotic partners. In the case of chloroplasts, the symbiont was a photosynthetic cyanobacterium; for mitochondria, most likely it was ana-proteobacterium.

The cytoplasm of eukaryotic cells is like chicken soup-it's chock full of organelles suspended like chunks of assorted vegetables and noodles in cytosolic broth. The broth also contains filaments of various dimensions that collectively comprise the cell's cytoskeleton. Like the bones of a large animal, the cytoskeleton provides a structural framework lending shape to cells and against which enzymatic 'muscles' work to elicit movement. That's how amoebae migrate, algae swim, stem cells divide, and cytoplasm streams relentlessly up, down, and across plant cells.

While the cytoskeleton is as much a hallmark of eukaryoticity as any mitochondrion or chloroplast, the origin of its filaments in deep time is more mysterious. Biologists assumed that genes for cytoskeletal proteins arose from prokaryotic precursors, but evidence in favor of the hypothesis was scarce, until recently.

Tubulin First on Stage

Microtubules comprise one component of the cytoskeleton responsible for a variety of movements including mitosis and meiosis. The 25 nm tubes consist of dimerica- and b-tubulin subunits that share about 40 percent sequence homology. Another form,y-tubulin, functions in microtubule formation.

But where did microtubules come from? It now appears that tubulins share a common ancestor with a protein called FtsZ, a key player in bacterial cell division.1 FtsZ is also present in plants, where it functions in chloroplast division,2 and a similar protein associates with mitochondria, at least in one alga.3 FtsZ polymerizes into filaments in the test tube in a process dependent on GTP. The same nucleotide is required for tubulin assembly into microtubules.1

Tubulins and FtsZ are clearly related, judging from similarities in three-dimensional structure. And although the proteins share only about 15 percent amino acid sequence identity overall, they're much more similar at the local level, particularly at the domain responsible for binding and cleaving GTP.4,5

Actin Into the Fold

Like the tubulins, actin-another essential component of the eukaryotic cytoskeleton-is a globular protein that binds nucleotide, in this case ATP. As actin monomers polymerize into 6-nm-wide microfilaments consisting of two helically wound protofilaments, the ATP, situated in a deep enzymatic cleft between two halves of the protein, hydrolyzes to ADP and inorganic phosphate.

It turns out that actin shares its ATPase domain with a family of proteins including hexokinase, the enzymatic kick starter of glycolysis, and several bacterial proteins. One of them is called MreB, a protein essential for generating or maintaining the rod shape of many bacteria. By examining structural similarities between eukaryotic actin and MreB from Thermotoga maritima, a research team at the Medical Research Council in Cambridge, England recently concluded that the two proteins are more closely related to each other than to other members of the family and undoubtedly share a common ancestor.6

The group showed that the three-dimensional shapes of actin and MreB are so similar they can be superimposed. The analogy with tubulin/FtsZ goes even further. Both proteins share considerable amino acid homology at several key sequences surrounding the ATP binding site, again situated deep in a cleft between two halves of the folded polypeptide chain.

Under the right conditions, MreB polymerizes into protofilaments that pair up lengthwise. The protein subunits are spaced about the same distance apart along the filaments as in polymeric actin, but MreB double filaments aren't nearly as helical.

The similarity between MreB and actin doesn't stop at structure and sequence. In a paper published earlier in 2001, a research group led by Jeffrey Errington at the University of Oxford, U.K. visualized MreB in the rod shaped cells of Bacillus subtilis using fluorescence and electron microscopy.7 MreB forms filamentous bands that encircle the cell in low helices, like reinforcing hoops. In an essay accompanying the Cambridge group's article, Duke University cell biologist Harold Erickson calculated that each band contains 10 protofilaments.8

When Errington's team genetically deprived cells of functional MreB, they became spherical. A search of genome databases showed that MreB is present in bacteria with nonspherical shapes, including rods. It's absent in spherical cocci. In other words, MreB has a cytoskeletal function. "I think it is quite convincing that MreB is the actin progenitor," says Erickson. "A key step, still unknown, going from bacteria to vertebrates is to develop a mechanism to make the double-helical actin filament from the single MreB protofilament structure."

More Acts to Follow

The story doesn't end with MreB; there's more to find out. Scientists want to know if MreB is also present in eukaryotes-associated with mitochondria and chloroplasts-as is FtsZ. According to Katherine Osteryoung, a plant biologist at Michigan State University in East Lansing who identified two FtsZ genes in the mustard plant Arabidopsis,2 "there's no obvious indication of MreB in plants that I've found or am aware of."

Actin normally functions along with the motor enzyme myosin to produce cellular motion, while microtubules utilize two other motor families called dynein and kinesin related proteins. Researchers now wonder whether MreB and FtsZ work in conjunction with bacterial motors. According to Erickson, "none have been turned up in genetic screens for cell division (or other activities), and none have been identified by sequence gazing. My bet is that kinesin and myosin evolved in eukaryotes, after the evolution of microtubules and eukaryotic actin filaments."

Still, Osteryoung is pleased with the latest results: "To someone interested in these issues, establishment of the prokaryotic origins of two major eukaryotic cytoskeletal proteins is enormously satisfying. I look forward to the day when evolutionary intermediates... from MreB to actin and FtsZ to tubulin, perhaps awaiting discovery in some obscure and primitive eukaryote, will more fully reveal the evolutionary steps by which key components of the eukaryotic cytoskeleton acquired their present-day structures and functions."

Barry A. Palevitz (palevitz@dogwood.botany.uga.edu) is a contributing editor for The Scientist.

References

  1. H.P. Erickson, "FtsZ, a tubulin homologue in prokaryotic cell division," Trends in Cell Biology, 7:362-7, 1997.
  2. K.W. Osteryoung, "Organelle fission: Crossing the evolutionary divide," Plant Physiology, 123:1213-6, 2000.
  3. P.L. Beech et al., "Mitochondrial FtsZ in a chromophyte alga," Science, 287:1276-9, 2000.
  4. E. Nogales et al., "Structure of the alpha-beta tubulin dimer by electron crystallography," Nature, 391:199-203, 1998.
  5. J. Lowe, L.A. Amos, "Crystal structure of the bacterial cell-division protein FtsZ," Nature, 391:203-6, 1998.
  6. F. Van den Ent et al., "Prokaryotic origin of the actin cytoskeleton," Nature, 413:39-44, Sept. 2, 2001.
  7. L.J.F. Jones et al., "Control of cell shape in bacteria: helical, actin-like filaments in Bacillus subtilis," Cell, 104:913-22, 2001.
  8. H.P. Erickson, "Evolution in bacteria," Nature, 413:30, Sept. 6, 2001.


TOPICS: Culture/Society; News/Current Events
KEYWORDS: crevolist; evolution
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To: tallhappy
This is only to be expected from evolution

Or from intelligent design. It's a wash if that's what you want to argue.

True enough. However, there are lots of patterns that evolution would not explain or allow for. Intelligent design can be made to account for anything at all. As a result, it's not a scientific theory.

41 posted on 01/14/2002 5:56:41 PM PST by Karl_Lembke
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To: lexcorp
Yes, Yahweh *may* have said "Man," and man was so. But there is equal evidense pointing to the universe being the dream of Brahma or having been sneezed out the nose of the Great Green Arkleseizure.

That is not quite what the account in Genesis one says. I must also respectfully disagree with your point about their being "equal" evidence for those other two ideas you name (realizing that the last was a humor attempt). There is far more evidence for the account in Genesis one than for the Hindu cosmology.

Heck, there is far more evidence for the Genesis account just in VERSE ONE than the whole Hindu cosomology. Genesis 1:1 states that the universe had a BEGINNING. That is in contrast to any STEADY STATE type theories which postulate that the universe has no beginning or end. These 'eternal universe' ideas have been discredited in favor of a cosmology consistent with the otherwise absurd idea that the entire universe was created from nothing. So teachers Genesis 1:1 and the Big Bang theory. Other religions have creation stories where the universe is made out of some eternal substance, or ideas that are even more discredited.

I understand that you are not claiming this evidence is a 'home run'. I am a bit sensitive on this point because so many otherwise intelligent evolutionists cannot seem to understand the difference between what the evidence plainly shows and the filter of their interpretation of that evidence.

A few threads ago I had an otherwise clever fellow seriously claim that evolution was well documented throughout the fossil record. His "proof" was that the fossil record showed that changes in the forms of organisms had occured!

He seemed impenatrably oblivious to the fact that the very focus of the debate was deciding WHAT FORCE WAS RESPONSIBLE for the changes shown in the fossil record. TO him, the very fact that new creatures were introduced to the biosphere was primie facie evidence that macroevolution must be true.

I'm glad you are not like that. YOu know that it is not a 'home run'. To even be a hit for either side, something about this data must show more support for evolution than for the Creationist subset of ID. THe fact that the two types of bacteria have building blocks that are in some ways similar is for the most part a wash.

I think Archeabacteria have proteins and DNA quite different from the other two types of cells. I could tout this as evidence that Creationism is true, since it is clear now that prokaryotes did not evolve from Archeans. You could retort that evolution happened twice. Another wash.

Discovering the truth in this matter is going to take a lot more objective thought than most are willing to put forth.

42 posted on 01/14/2002 6:46:48 PM PST by Ahban
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To: Karl_Lembke
" Intelligent design can be made to account for anything at all."

Probability theory and information theory are unscientific? Have you read The Design Inference: Elimenating Chance Through Small Probabilities? If not, you should do so before saying what can and cannot be accounted for by these disciplines.

43 posted on 01/14/2002 7:08:05 PM PST by Bonaparte
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To: Ahban
An excellent post, Ahban. My one qualm is your characterizing ID as a "subset of creationism." The variations in theory found in the school of "scientific creationism" all have one thing in common -- they maintain that God created the universe and purport to show that there is a scientific basis for this conclusion. ID does not do this. In essence, it states that any outcome is either the result of chance or of intention, and that there are valid scientific ways of determining which one applies to an observed outcome.

The article at the heading of this thread is a perfect example of unscientific method. It repeatedly takes as a premise (macroevolution) the very thing it hopes to establish as a conclusion (macroevolution). This is circular reasoning.

If a preponderance of the evidence suggests that life was brought into being by conscious agency and not by chance, what can one say of a "scientist" who refutes what the best evidence points to?

44 posted on 01/14/2002 7:38:58 PM PST by Bonaparte
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To: lexcorp
Correct me if I'm wrong, lexcorp, but I have the impression that you haven't actually read much in the literature of ID -- in particular, The Design Inference (William Dembski, Cambridge University Press, 1998).
46 posted on 01/14/2002 7:52:16 PM PST by Bonaparte
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To: lexcorp
Not having detailed knowledge regarding protein sythesis and evolution MUST make me ignorant of even the basics of biological theory and knowledge!

Yes.

Whether or not creationism is bunk has nothing to do with your lack of knowledge and understanding fo basic biology. But it is clear you also lack basic logic skills.

47 posted on 01/14/2002 7:53:04 PM PST by tallhappy
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To: PatrickHenry
In every thread I am correct.

Thus ad hominums are your response.

48 posted on 01/14/2002 7:54:36 PM PST by tallhappy
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To: Bonaparte
Thanks for the support! And let me ease your one qualm, I think I said that Creationism is a subset of ID, not the other way around. That is my position, that Creationism is a major subset of ID, not that ID is a part of Creationism.
49 posted on 01/14/2002 7:56:50 PM PST by Ahban
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To: VadeRetro
I suspect he isn't tall, either.

LOL! (lucky my mouth was not full of pop...)

50 posted on 01/14/2002 7:57:35 PM PST by jennyp
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To: RightWhale
A common structure or composition implies one is the ancestor of the other, or the other is more evolved.

Or that both have a common ancestor. Or (less likely) convergent evolution.

51 posted on 01/14/2002 8:00:43 PM PST by Doctor Stochastic
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To: Doctor Stochastic
The idea is that all this is in context of an evolution model. It's a strong model, but still just a model.
53 posted on 01/14/2002 8:06:34 PM PST by RightWhale
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To: RightWhale
So? Some models are more useful than others.
54 posted on 01/14/2002 8:07:58 PM PST by Doctor Stochastic
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Comment #55 Removed by Moderator

To: Doctor Stochastic
Or (less likely) convergent evolution.

If it was convergent evolution, I think we'd expect the actual amino acid sequences of the proteins to differ just as much at the functional points as at the purely structural points, instead of showing more sequence similiarity like they do.

(Convergent evolution is where 2 unrelated species evolve a superficially similar solution to the same engineering problem, isn't it? In that case, the similarities should break down upon closer analysis, such as in the amino acid or the DNA sequences.)

56 posted on 01/14/2002 8:10:03 PM PST by jennyp
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To: longshadow
Bttt for me too! :)
57 posted on 01/14/2002 8:11:05 PM PST by RadioAstronomer
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To: Ahban
"... than for the Creationist subset of ID."

Thank you for clarifying, Ahban. Looking at your original phraseology, it could be read the other way, eg. the 'English cities' subset of 'Birmingham.' Had you said, "Creationism as a subset of ID" there would have been no ambiguity.

58 posted on 01/14/2002 8:11:22 PM PST by Bonaparte
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To: lexcorp
And in any event, the Heisenberg Uncertainty principle points out that things CAN pop into existence out of nothing

Just a quick comment. Quantum events are based on probabilities within given intervals of time; before the big bang there was no time. In my opinion that rules out quantum mechanics as the cause of the Universe.

59 posted on 01/14/2002 8:12:42 PM PST by week 71
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Comment #60 Removed by Moderator


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