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Evolution of an Irreducibly Complex System – Lenski’s E. Coli

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On another thread we have been discussing abiogenesis in particular, but there was also some discussion about the evolution of an irreducibly complex system. Commenter CHartsil indicated that “we actually watched an IC system evolve” in reference to Lenski’s E. coli research. At my request, he has posted a brief summary of the research and his take, which I am now elevating to a new thread on this important topic.

For those who disagree with CHartsil’s take, strong objections on substantive grounds are of course welcome, whether relating to Lenski’s research or CHartsil’s interpretation of it, but not irrelevant personal attacks. Thank you.

—–

Guest Post by CHartsil:

This is a pro-ID board so I doubt I need to explain irreducible complexity. When arguing against it, most will bring up Ken Miller or Nick Matzke. They have great points but theirs are indirect and theoretical pathways for systems considered IC. That’s why I’m fond of Lenski’s cit* E. coli.

This particular strain of E. coli evolved the ability to metabolize citrate aerobically. While most E. coli can do this anaerobically, part of the definition of wild-type E. coli is actually the inablity to use citrate as a substrate aerobically. This may not have been a terribly fascinating addition of function if not for the frozen fossil records kept by Lenski et al.

These frozen generations allowed Lenski to determine that this trait was not acquired via a single mutation as it could only be repeated after generation 20,000. Given the distinct cladistic division amongst the populations at the border generation, it was determined that there were at least two potentiating mutations prior to the cit* event.

In this third clade a tandem duplication resulting in a novel regulatory module leading to the aerobic cit* could be repeated and verified. It has been noted since that the fitness of the population has been improving without notable upper limit, increasing based on the number of copies of the new regulatory module.

I find this to be sufficient in warranting the dismissal of the concept irreducible complexity. In Lenski’s E. coli, we observe the rise of a new function resulting from a new gene and new gene regulation. This function is comprised of now interdependent components which demonstrably did not exist in parent generations. It is by definition irreducibly complex and it was observed to evolve.

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Nota bene: for purposes of the above discussion, CHartsil is using the following definition of irreducible complexity: “a system comprised of interdependent parts, the removal of any of which will cause the system to cease functioning.”

Comments
Natural selection, ie evolution, [...]
Joe, natural selection and evolution are not synonyms.Hangonasec
March 1, 2015
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fmm
Any heritable change to an organism is evolution. Really?
Really. Now you're gettin' it! I would differ with Zachriel somewhat in that it does not have to become widespread, although it is of little interest (like quantum particles popping into and out of existence) until it does.Hangonasec
March 1, 2015
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Zac, Just to be super clear. Your position is that any heritable change to a population has to be the result of evolution whether it has any connection to RV/NS or not.fifthmonarchyman
March 1, 2015
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Natural selection, ie evolution, pertains to individuals...Joe
March 1, 2015
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fifthmonarchyman: Any heritable change to an organism is evolution. Evolution refers to the population, so the change has to become part of the population. (ETA: That might be encapsulated in the term heritable.)Zachriel
March 1, 2015
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Zac say, Horizontal gene transfer results in a heritable change to an organism, so, of course, it’s evolution. I say, eyes roll. Any heritable change to an organism is evolution. Really? peacefifthmonarchyman
March 1, 2015
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Zac says Then the public may need to be better educated, though most schools do cover sexual recombination at least. I say, Lets see, The un-evolutionarily process of two organisms in the same population combining to produce a third organism in the same population is some how relevant to a population stealing vital structures from a completely unrelated population at just the right time being the result of evolution. You kill me Zac peacefifthmonarchyman
March 1, 2015
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fifthmonarcnhyman: We are talking about evolutionary verses other natural processes. Is this another case where you are using special fmm definitions? Horizontal gene transfer results in a heritable change to an organism, so, of course, it can result in evolution. fifthmonarcnhyman: The only way your comment makes any sense is if you are assuming that all natural biological processes are evolutionary. Sex is not evolution. Development is not evolution. Metabolism is not evolution. Ambulating is not evolution. Mastication is not evolution. A flower blooming is not evolution.Zachriel
March 1, 2015
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zac says, Natural horizontal mechanisms are observed. Just because humans insert genes ‘horizontally’ doesn’t mean that all horizontal change is artificial. I say, Who said anything about natural verses artificial? We are talking about evolutionary verses other natural processes. The only way your comment makes any sense is if you are assuming that all natural biological processes are evolutionary. Oh never mind you are assuming that ;-) I hope you see why such equivocation makes productive communication impossible peacefifthmonarchyman
March 1, 2015
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ZAchriel may need to be better educated as no one can say if blind watchmaker processes produced the changes Lenski observed.Joe
March 1, 2015
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PaV: Fine. So we have an example, according to Behe, of a gain of Functional Coded elemenT in a small experimental population of bacteria. PaV: To develop the Cit+ trait, somewhere between half-a-trillion and a trillion replications of the bacteria had to occur. Far fewer than happened in the typical human gut in the last few days. PaV: A gene duplication event (hence no new information) and a few SNP mutations afterwards (IOW, a few amino acids likely changed). Along with a potentiating mutation. PaV: How could such a process possibly account for large scale genomic changes? Let's see... An ocean of microbes over a billion years compared to a few lab specimens over a couple of decades. PaV: So, that’s 40 million “replications” per 100 years. The pachyderm genome is far more complex, even including a body plan toolkit. fifthmonarchyman: As a rule of thumb the more complex the occurrence the less likely that it happened by chance. Complexity can increase through selection, even if each step is happenstance. fifthmonarchyman: The only way to include things like HGT in “evolution” is to equivocate the term until it means any change that is biological. Natural horizontal mechanisms are observed. Just because humans insert genes 'horizontally' doesn't mean that all horizontal change is artificial. fifthmonarchyman: Of course it’s beneficial for the absorbing organism but it’s not evolution as it’s understood by the public. Then the public may need to be better educated, though most schools do cover sexual recombination at least.Zachriel
March 1, 2015
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Hey Hangonasec When I was a boy I used to play a game called superheroes in which my brothers and I would play act battles using superpowers it would often go something like this player one--- I have super speed I can catch you anywhere player two----I have super strength so you can't hold me when you catch me. player one----I now have superpower absorbing powers. Whatever power you have I can absorb just when I need to use it. player two----This game sucks. You are cheating you are guaranteed to win before we even start. That's what HGC does it lets one organism steal the "evolutionary powers" of other organisms without going through the actual evolutionary process necessary to acquire them. Of course it's beneficial for the absorbing organism but it's not evolution as it's understood by the public. It's cheating. peacefifthmonarchyman
March 1, 2015
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Hangonasec I don’t see why one should accept that a single base change (point mutation), or some rearrangement/duplication from within a genome, are evolution, yet a change involving bases of external source is not. I say, Actually I would not automatically accept rearrangement/duplication from within a genome to be a random occurrence. At very least this has not been established it's more likely they are not random at all,We know for example that duplication/rearrangement are more likely in stressful environments. As a rule of thumb the more complex the occurrence the less likely that it happened by chance. A fortuitous insertion of a complex vital structure from a external source is just too much to accept as luck of the draw. Especially given that a major priority of organisms is to keep these sorts of contaminants out. If traditional evolutionary mechanisms (RM/NS) were powerful HGC would not happen. The only way to include things like HGT in "evolution" is to equivocate the term until it means any change that is biological. Apparently that is just what is happening here peacefifthmonarchyman
March 1, 2015
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fmm @167:
Can you? [indicate permanent biological change that is not evolution], I say, Of course, Off the top of my head Things like HGT are not evolution. Combining two systems into a new third system is not evolution.
I don't see why one should accept that a single base change (point mutation), or some rearrangement/duplication from within a genome, are evolution, yet a change involving bases of external source is not. If a genetic fragment - say a retroviral insert, or a piece of Wolbachia - is incorporated into a genome, and then inherited down the main line, it is functionally contributing to the 'RV' part of the oversimplified 'ID evolution equation' RV+NS, just as much as point mutation. If 2 entire genomes combine, and the combination remains intact in subsequent generations, then again that is generating the 'RV' part.Hangonasec
March 1, 2015
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Done reading this garbage, lol. Send this OP claim over to a journal for peer-review and let your "peers' over at Nature tell you how ignorant it is after tossing your paper directly into the garbage bin. ////Paleysghost
February 28, 2015
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Since this thread has served its purpose, let me just note one thing before leaving. To develop the Cit+ trait, somewhere between half-a-trillion and a trillion replications of the bacteria had to occur. And this was, just as in the case of the malarial parasite, an instance where the "selection pressure" for adaptation was quite large. What do we see happening? A gene duplication event (hence no new information) and a few SNP mutations afterwards (IOW, a few amino acids likely changed). Well, this is, AGAIN, the "edge of evolution." So much replicating, and so little to show for it. How could such a process possibly account for large scale genomic changes? A Scientific American article said that in the 1930's there were perhaps 5 to 10 million elephants in Africa. So, with a generation time of 25 years, and with the population replacing itself---that is, a male and female elephant give rise to another male and female elephant, the population replaces itself every 25 years. So, that's 40 million "replications" per 100 years. So, it would take 25,000 of these "replications" to reach a trillion. This is equivalent to 2.5 million years. IOW, to get a gene duplication to end up in just the right place to help an African elephant, we need 2.5 million years. And, of course, there has been no new genomic information added. To me it seems like we either ought to pretend that we know nothing about coded information and its implication, or, we through out neo-Darwinism. Both can't continue to co-exist.PaV
February 28, 2015
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Apparently, the guy who responded didn't understand his own link to the paper. The mutations in this VERY REDUCIBLE pathway do not contribute as being parts of a new "IC apparatus", but are rather just STEPWISE contributions that contribute to a REDUCIBLE PATHWAY which leads to a specific function. You do not understand what the IC hypothesis means. You do not understand what your own links imply. Discussion on this article was DEAD and over a year ago when this troll first posted it to his FAKE ID group on facebook and the fact that is has 200 comments on UD is a testimony to how people really just feel a need to argue and type stuff on the internet for no produxctive reason.Paleysghost
February 28, 2015
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Piotr:
When you want to discuss physics, you ought to use words like “mass”, “power”, “wave” or “potential” in their scientific sense, not colloquially. Ditto for biology. Every discipline has its technical terminology for the sake of precise communication. If you don’t use it, you confuse the issue.
Let’s look at this two ways: (1) What do they mean when Darwinists say that “evolution” is a fact? They’re, of course, referring to the “fossil record.” But then when they talk about “evolution” they also mean RM+NS. So, which is it? (2) The “epicycles” of the Ptolomeic system were, I’m sure, “well-defined,” and, I’m sure, everyone knew what was meant by them. Did that make them correct? So, we have ‘equivocation,’ and we have ‘error’. If you want to avoid both of these problems, then talk about “change” and “adaptation.” And, if you should find instances of “evolution,” then call that evolution. Good luck finding them (and here I’m referring to “macroevolution,” which is the common sense understanding to the term)
Look at the millions of insect species, or the mollusc lineage, which has produced a variety of animals from sedentary mussles with no brain at all to octopuses with a complex brain as big as a cat’s and with ape-like intelligence.
I appreciate this insight. I think you’re right; I haven’t fully appreciated invertebrate diversity as much I should have. Good point.
No? How about spawn, tadpoles and adult frogs, an egg and a chicken, an acorn and an oak-tree? A single life cycle is not evolution, just like one rung is not a ladder and one note is not a musical piece.
Either the point eludes you (unlikely), or you choose to not look at it too closely (more likely). The question I asked was not: are there other examples of large-scale phenotypic diversity? I asked was there an example of greater diversity. The examples you give are examples of straight-forward developmental biology, and nothing more, except in the case, perhaps, of tadpoles; but this is probably nothing more than arrested development. However, the correct analogy would be for an acorn to become an oak tree, then dissolve into liquid, and then give rise to a fig tree. Even this is not so much of a huge change in phenotype as caterpillar to butterfly; but, at least it gets us a little bit closer to the reality. Finally, let’s note that you relied on “developmental biology” and not on a “change in allele frequency” when giving examples of phenotypic change. IOW, we do better when we leave the “gene-centric” view behind. That is, we’ll be hearing a whole lot more about “Junk-DNA” in the near future. (I’m speaking cryptically here, and hope you can follow)
Yep, that’s why it’s developmental biology, not evolution (micro- or macro-).
Neo-Dawinism links phenotypic change to genotypic change. I don’t think you would disagree with this. I gave an example of a HUGE phenotypic change with NO genotypic change at all to underlie it. Well, then, isn’t changing “beak sizes” trivial compared to a caterpillar becoming a butterfly? It would seem that this should give a thinking person pause before they taut the wonders of what “changing allelic frequencies” can bring about. Is the putative non-sequitur now a little bit less of a non-sequitur? IOW, “microevolution” is no more than an organism “adapting” to an environment; and “macroevolution” cannot be demonstrated. Let’s look for a better way of explaining things. And let’s look for a better lexicon.PaV
February 28, 2015
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The problem, Piotr, is that there is a common sense notion of evolution, and then there is a “scientific” understanding of evolution. The supposed scientific definition, as used by wd400, is at odds with the “common sense” notion, and ends up confusing the issue.
When you want to discuss physics, you ought to use words like "mass", "power", "wave" or "potential" in their scientific sense, not colloquially. Ditto for biology. Every discipline has its technical terminology for the sake of precise communication. If you don't use it, you confuse the issue.
You say that worms “evolve.” That there are all kinds of phyla. wd400 says that “evolution” is changing allele frequencies (which, of course, in any given species, as in humans, can already be found, though there is no “evolution”). I don’t think anyone associated “evolution” with either of these notions.
Well, all biologists do.
Why not try to be clear in what we are saying: such as, “adaptation,” and “diversification.” That the chordate lineage produced such things as dinosaurs, birds, apes and humans—now that is “evolution.”
You clearly don't realise how much invertebrates differ from one another in terms of complexity, body plan, and even intelligence. Look at the millions of insect species, or the mollusc lineage, which has produced a variety of animals from sedentary mussles with no brain at all to octopuses with a complex brain as big as a cat's and with ape-like intelligence.
As I stated before, a caterpillar (I wonder the phyla) becomes a butterfly. Can you think of any greater phenotypic diversity in all of nature? No.
No? How about spawn, tadpoles and adult frogs, an egg and a chicken, an acorn and an oak-tree? A single life cycle is not evolution, just like one rung is not a ladder and one note is not a musical piece.
And it happens in a matter of weeks or days. And yet there are NO allelic frequencies changing because it’s the exact, same genome that produces both.
Yep, that's why it's developmental biology, not evolution (micro- or macro-).
With this simple observation, all of Darwinian theory should be thrown out.
Non sequitur.Piotr
February 28, 2015
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PaV says, The supposed scientific definition, as used by wd400, is at odds with the “common sense” notion, and ends up confusing the issue. It is my impression that evolutionary biologists, otherwise known as Darwinists, like it just this way; however, the general public, and science itself, is not served by using equivocal terms. I say, exactly!! quote: "By 2050—earlier, probably—all real knowledge of Oldspeak will have disappeared. The whole literature of the past will have been destroyed. Chaucer, Shakespeare, Milton, Byron—they'll exist only in Newspeak versions, not merely changed into something different, but actually contradictory of what they used to be..........The whole climate of thought will be different. In fact there will be no thought, as we understand it now. Orthodoxy means not thinking—not needing to think. Orthodoxy is unconsciousness." end quote: George Orwell 1984 peacefifthmonarchyman
February 28, 2015
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Innovation or Renovation? By Ann Gauger - Sept. 24, 2012 Excerpt: But how significant was this innovation (citrate; Lenski)? In his paper in Quarterly Review of Biology, Dr. Michael Behe pointed out that E. coli was already capable of using citrate for anaerobic growth (when no oxygen was available). He postulated that a change in gene regulation could turn on citrate transport and permit growth on citrate under aerobic conditions. After an enormous amount of work, having sequenced the genomes of many clones along the lineages that led to the ability to use citrate, as well as lineages that never did, and testing the phenotypes of identified mutations, Blount et al. have now reported that Behe was largely right. The key innovation was a shift in regulation of the citrate operon, caused by a rearrangement that brought it close to a new promoter. http://www.biologicinstitute.org/post/32246480851/innovation-or-renovation?og=1 Rose-Colored Glasses: Lenski, Citrate, and BioLogos - Michael Behe - November 13, 2012 Excerpt: In my own view, in retrospect, the most surprising aspect of the oxygen-tolerant citT mutation was that it proved so difficult to achieve. If, before Lenski's work was done, someone had sketched for me a cartoon of the original duplication that produced the metabolic change, I would have assumed that would be sufficient -- that a single step could achieve it. The fact that it was considerably more difficult than that goes to show that even skeptics like myself overestimate the power of the Darwinian mechanism. http://www.evolutionnews.org/2012/11/rose-colored_gl066361.html Bacterial 'Evolution' Is Actually Design in Action by Brian Thomas, M.S. - Dec. 2012 Excerpt: At that time, the mechanism underlying the citrate-eating phenotype was unknown. Behe wrote, "If the [Cit+] phenotype is due to one or more mutations that result in, for example, the addition of a novel genetic regulatory element, gene-duplication with sequence divergence, or the gain of a new binding site, then it will be a noteworthy gain-of-FCT [Functional Coded elemenT] mutation."2 So, the big question is: Did E. coli evolve into a Cit+ strain by natural selection? Or did mutations construct new and functional coded elements to its DNA? If so, it would be the first in recorded biological history. If not, then it would be just another loss or modification of a pre-existing piece. In Lenski's experiment, the bacteria (both Cit+ and wild-type) already possessed a gene named citT. It encodes a protein that transports a range of citrate-like chemicals. The recent results showed that the bacteria made extra copies of citT and a neighboring sequence—a process called gene amplification. More copies of the gene should translate to higher amounts of the transporter protein that it encodes. With enough transporters, the bacteria could access enough citrate. But oxygen deactivates citT, and having many copies of a gene that is turned off is not very useful! But the bacteria solved this problem when the amplification event also moved the gene sequence to a different place in the bacterial chromosome, where a different but pre-existing promoter could regulate it. Unlike the original one, it appears that the new promoter does not have an "oxygen off" switching mode. Instead, it allowed expression of citT in the presence of oxygen so that the bacteria successfully imported enough citrate to grow. The study authors wrote, "The structure of the cit amplification led us to propose that the Cit+ trait arose from an amplification-mediated promoter capture."1 Further investigation confirmed the proposal. So, the bacteria solved the problem of accessing an alternative food source by generating extra copies of the critical gene and by placing those copies under the control of an appropriate promoter. Does any of this resemble natural, undirected Darwinian evolution? Not at all. This amazing mechanism invented no new functional coded elements. It merely modified pre-existing elements. Therefore, not only did the Cit+ bacteria not evolve in the molecules-to-man direction, but they showed what could only be ingenious DNA rearrangement mechanisms. What mainstream headlines portrayed as evidence for evolution is actually the opposite.3 http://www.icr.org/article/bacterial-evolution-actually-design/bornagain77
February 28, 2015
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Piotr:
And, please, whatever the merits of Webster’s dictionary, it isn’t the right reference book for checking up scientific terminology.
The problem, Piotr, is that there is a common sense notion of evolution, and then there is a "scientific" understanding of evolution. The supposed scientific definition, as used by wd400, is at odds with the "common sense" notion, and ends up confusing the issue. It is my impression that evolutionary biologists, otherwise known as Darwinists, like it just this way; however, the general public, and science itself, is not served by using equivocal terms. Adaptation can be demonstrated; but not evolution. We would be better served reserving the word "evolution" for a kind of progressively more complex development of phenotype. You say that worms "evolve." That there are all kinds of phyla. wd400 says that "evolution" is changing allele frequencies (which, of course, in any given species, as in humans, can already be found, though there is no "evolution"). I don't think anyone associated "evolution" with either of these notions. Why not try to be clear in what we are saying: such as, "adaptation," and "diversification." That the chordate lineage produced such things as dinosaurs, birds, apes and humans---now that is "evolution." As I stated before, a caterpillar (I wonder the phyla) becomes a butterfly. Can you think of any greater phenotypic diversity in all of nature? No. And it happens in a matter of weeks or days. And yet there are NO allelic frequencies changing because it's the exact, same genome that produces both. With this simple observation, all of Darwinian theory should be thrown out.PaV
February 28, 2015
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Zachriel:
Behe, Experimental Evolution, Loss-of-Function Mutations and ‘The First Rule of Adaptive Evolution’,Quarterly Review of Biology 2010: “If the phenotype is due to one or more mutations that result in, for example, the addition of a novel genetic regulatory element, gene-duplication with sequence divergence, or the gain of a new binding site, then it will be a noteworthy gain-of-FCT mutation.”
Fine. Yes, this IS the "edge of evolution." Alas. As Eric points out, this is a pittance of neo-Darwinian evolution needs to be. And, certainly, it's "not" an IC situation.PaV
February 28, 2015
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Worms evolving into worms fits in with baraminology. And perhaps Piotr is closely related to worms and he should speak for himself.Joe
February 28, 2015
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See Blount et al., Genomic analysis of a key innovation in an experimental Escherichia coli population, Nature 2012: “The Cit(+) trait originated in one clade by a tandem duplication that captured an aerobically expressed promoter for the expression of a previously silent citrate transporter. The clades varied in their propensity to evolve this novel trait, although genotypes able to do so existed in all three clades, implying that multiple potentiating mutations arose during the population’s history.”
No one knows if that means the mutations were accidents, mistakes and errors, or not.Joe
February 28, 2015
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#186, PaV
With all due respect, no, I am not wrong. To say a worm “evolved” into a worm makes no sense.
Of course it does. There are about a dozen different phyla of "worms". "Worms" have been diversifying for more than half a billion years, evolwing into hundreds of thousands of species. You are just about as closely related to a sea urchin as one worm (e.g. an earthworm) is to another (e.g. the nematode Caenorhabditis elegans). And, please, whatever the merits of Webster's dictionary, it isn't the right reference book for checking up scientific terminology.Piotr
February 28, 2015
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Pav
If this is so, then they didn’t use a “minimal growth medium,” but, instead, should have said they used a “minimal growth medium with a high citrate concentration in the form of a chelating agent.” They didn’t say that. I looked around. I don’t know where you get the figure for citrate, I can only find a pnas paper where they say the citrate concentration was “high,” giving no ppm. [Blount-Borland-Lenski, 2008] That actually makes more sense of the experiment, and why the fitness increased so greatly once the citrate transport protein was activated.
Of course the authors used a minimal growth media for the species they were workiing with, i.e., E coli. The authors did an excellent job of providing the details of their methodology. That you aren't familiar with what a 'DM25' culture media consists of is not their fault. You could have, after all, looked up what the DM25 media consists of....it is information that is readily available and to have them spell it out would be a waste of $$ in additional page costs. The funny thing is that you've gotten everything wrong about the details of the expt. pav
If this is so, then they didn’t use a “minimal growth medium,” but, instead, should have said they used a “minimal growth medium with a high citrate concentration in the form of a chelating agent.” They didn’t say that
They did say that quite explicitly in their Mat. and Met section as Zachriel has also pointed out. It is only your ignorance of the contents of the paper and your lack of understanding about the methodologies and purpose of the expt. that is getting in the way of any fruitful discussion. Instead 'we' are left with pointing out your lack of knowledge of this expt evidenced by all the details you've gotten wrong. Of course the results make more sense once you understand the contents of the paper outlining the expt.franklin
February 28, 2015
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PaV: So the problem is that the authors did not do a good job—at least in the papers I looked at briefly and searching for citrate concentration—of telling people what the “minimal growth medium” consisted of. They discuss the medium in their paper on contingency under Methods and Materials, Blount et al., Historical contingency and the evolution of a key innovation in an experimental population of Escherichia coli, Proceedings of the National Academy of Sciences 2008. You can read more about protocols on their website. http://myxo.css.msu.edu/ecoli/standprot.htmlZachriel
February 28, 2015
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wd400:
You are just wrong to think evolution means “progress” or enough progress to change a species name(!).
With all due respect, no, I am not wrong. To say a worm "evolved" into a worm makes no sense. Here's the Webster Dictionary definition (quibble with them):
1. the process by which different kinds of living organisms are thought to have developed and diversified from earlier forms during the history of the earth. [synonyms: Darwinism, natural selection "his interest in evolution"] 2. the gradual development of something, especially from a simple to a more complex form.
If you want to talk about "microevolution" and "macroevolution," then that's something different. But why equivocate when it comes to the meaning of "evolution"? wd400:
I don’t know why I bothered filling in the gaps in your knowledge of the Lenski experiment, it’s obvious you have no need of just details in creating your opinions.
Again, with all due respect, I know of no "gaps" that you filled in. Your concern is with, and only with, allele frequencies. Well, if you're a population geneticist, and you are, then this has been the object of your adult professional life. Does this alone give you the right to call it "evolution," and then say that this is what science says? I say, "no." For example: in the case of Cit+ the "allele" frequency of the transport protein for citrate has increased. Is this 'change' in "allele frequency" the cause of aerobic citrate metabolism? No. It's the position of the duplicated allele. This should just be obvious. All of this should clearly indicate that it is the regulatory mechanisms of an organism that are running the show, and that "allele frequencies" play more of a secondary role; not a primary one. franklin:
All e coli have the ability to metabolize citrate. The fitness increase in the Lenski expt(s) was not due to the bacteria developing the ability to metabolize citrate…it already had that ability! Wrong again, PaV.
Why the bluster? You know darn well that what we're talking about here is aerobic citrate metabolism, which E. coli does NOT have; it must acquire. Per your statement, Lenski's experiment showed nothing. Again, why the bluster? franklin:
In the Lenski expt glucose levels in the culture media were 25 ppm yet, contrary to your claim, citrate levels were over 350 ppm
If this is so, then they didn't use a "minimal growth medium," but, instead, should have said they used a "minimal growth medium with a high citrate concentration in the form of a chelating agent." They didn't say that. I looked around. I don't know where you get the figure for citrate, I can only find a pnas paper where they say the citrate concentration was "high," giving no ppm. [Blount-Borland-Lenski, 2008] That actually makes more sense of the experiment, and why the fitness increased so greatly once the citrate transport protein was activated. So the problem is that the authors did not do a good job---at least in the papers I looked at briefly and searching for citrate concentration---of telling people what the "minimal growth medium" consisted of. Maybe you can point this out to the authors.PaV
February 28, 2015
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Paleysghost: Lenski’s Cit. observations involve an extremely simple biochemical pathway which is formed from 2 consecutive (easily reducible) mutations, thus proving Darwinian evolution is true!! Um, no. See Blount et al., Genomic analysis of a key innovation in an experimental Escherichia coli population, Nature 2012: "The Cit(+) trait originated in one clade by a tandem duplication that captured an aerobically expressed promoter for the expression of a previously silent citrate transporter. The clades varied in their propensity to evolve this novel trait, although genotypes able to do so existed in all three clades, implying that multiple potentiating mutations arose during the population's history."Zachriel
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