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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.

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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
fifthmonarchyman: So if occurs in biology it is the result of evolution no mater what the process that gave rise to it. No. The argument based on irreducible complexity is that there are no plausible intermediate steps. We provided two plausible scenarios where irreducible complexity can evolve step-wise. fifthmonarchyman: Is there any natural process that you would not consider evolution? Only biological organisms are subject to biological evolution. Are you conflating this with other uses of the word evolution, as you did with rivers above? fifthmonarchyman: Think of an archway the individual stones can exist but a structure is not an archway until all the stones are present and interacting with each other. It is impossible for such a structure to arrive via evolution this should be obvious. Yet archways can form naturally. fifthmonarchyman: Part A can not “evolve” into system (a union B) by definition. Argument by definition is not an argument. fifthmonarchyman: This is not to say that it is not possible for system (A union B) is arise by natural processes only that it can’t arise by “evolution” Scare-quotes don't constitute an argument either. fifthmonarchyman: The more parts and the more intricate the union the less likely a system can arise by chance. It's not chance, but posited incremental adaptation leading to step-wise evolution of irreducible complexity. PaV: Here’s Michael Behe’s take on the Lenski experiment. 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."Zachriel
February 28, 2015
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End thread//Paleysghost
February 27, 2015
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"The Darwin camp has found a new mechanism, thanks to their supporter Christopher Hartsil (real name withheld to protect the identity of a minor) and his wonderful discovery of "Reducible Simplicity". 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!! ...as highlighted by Michael Behe in 1996 and 2007. Keep up the good work there, buddy."Paleysghost
February 27, 2015
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Mere details UBP. With sufficient imagination they can be overcome.Mung
February 27, 2015
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...and of course, there is that one other little tidbit where Darwinian evolution requires IC to even exist. :)Upright BiPed
February 27, 2015
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CHartsil: Here is the bottom line. First of all, thank you for taking time to post your thoughts about Lenski’s E. coli experiments. There has been much discussion, and several commenters have addressed the details of those experiments. Unfortunately you continue to persist in assertions that a new irreducibly complex system evolved right before our eyes and that, therefore, Lenski’s experiments somehow refute Behe’s notion of irreducible complexity specifically, and the design inference generally. At the risk of restating some of the points that have already been made in this thread, there are three key issues that need to be brought to the fore. 1. Your position appears to be based more on enthusiastic pro-evolutionary commentary than on the actual science. Again, I would refer you to Behe’s 2010 paper or at least the discussion posted by kairosfocus @2-3 above to get a more objective view as to what actually occurred via the evolutionary mechanisms. The adaptive changes were small in both number and scope, did not result in the creation of any information-rich structure, and likely resulted from a loss of function of pre-existing parts. 2. You are implying, if not directly stating, that Lenski’s experiments refute Behe. That is a misrepresentation of Behe’s arguments. Behe has certainly never argued that if a significant adaptive advantage can be obtained by making a small number of mutations to a pre-existing irreducibly complex structure, in a large population, over thousands of generations, under extreme selective pressure, that a trial-and-error process like random mutations plus natural selection could never produce such a change. Indeed, Behe himself has looked at similar situations with malaria and sickle-cell anemia, and such changes are well within what he has termed the “edge of evolution.” And that is setting aside for a moment the fact that subsequent research suggests that the adaptations in Lenski’s E. coli might well be instances of loss of pre-existing function – breakage of cellular mechanisms – rather than gain of additional function. Such a result would follow Behe’s “First Rule of Adaptive Evolution,” which says (roughly paraphrasing) that if a selective advantage can be obtained either by breaking a pre-existing functional structure or by building a new structure, then evolution will break the pre-existing structure. Thus, if anything, Lenski’s experiments support both Behe’s notion of irreducible complexity, as well as his rule of adaptive evolution. 3. Finally, and perhaps most importantly, we need to step away from the angels-on-the-head-of-a-pin claim that we are dealing with a “new” IC system, rather than a minor adaptation of an existing IC system. Although the latter is clearly the case, I will defer for a moment, just for purposes of discussion. Let’s assume, as you vocally insist, that Lenski’s E. coli experiments show a “new” IC system, just for argument’s sake. What does that teach us? Ultimately, what we are dealing with is this: According to the Darwinian storyline, complex, functional, information-rich structures are supposed to arise by evolution all the time. Yet after decades of dedicated research to find the smoking gun of evolution and after billions of dollars collectively spent by the evolutionary community on the effort, what does evolutionary theory have to show for it? Well, for one thing we have Lenski's results: A small number of beneficial mutations to a pre-existing biological system that eventually occurred after thousands of generations in a large population that was subject to extreme selective pressure. This is an interesting result, to be sure. But in the context of the larger claims of evolution’s grand creative power, it is a pittance, a rounding error, the exception that proves the uncomfortable rule. It is well within the edge of evolution that Behe has proposed, and does nothing to suggest that the Darwinian mechanism has any great creative power. Conclusion: When viewed objectively, without the blinders of materialistic philosophy, Lenski’s experiments are strong evidence for, and a powerful vindication of, Behe’s arguments in particular, and the skeptics of Darwinism in general. At the end of the day, Lenski’s E. coli are just another in a long line of Darwinian claims that, upon closer inspection, fail to live up to their hype: from moth coloration to finch beaks to antibiotic resistance in bacteria. In each case, when these examples are carefully studied, instead of supporting the grander claims of the evolutionary creation story, they inadvertently demonstrate the pathetic impotence of Darwinian mechanisms and underscore the need for an alternative source of innovation in the history of life.Eric Anderson
February 27, 2015
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Joe:
Rivers do not evolve . . .
Sure they do. You just have to use one of the ever so many definitions of that slippery word: "change over time." The word "evolution" has so many disparate definitions in practice (and I'm talking about the literature, not just blogs and lay commentary), that it is a real challenge just to keep track of them.Eric Anderson
February 27, 2015
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pav
Citrate is a suitable source for metabolism in organisms that can metabolize them. But in a minimal growth medium, a “limited” growth medium, where glucose serves as the carbon source, citrate is present at a very low level, and is not the carbon source.
In the Lenski expt glucose levels in the culture media were 25 ppm yet, contrary to your claim, citrate levels were over 350 ppm. Hardly low levels by any consideration. So, pav, you were wrong on this issue as well as the other details that have been pointed out to you. One gets the feeling you aren't really familiar with the Lenski expt(s) or the 'animal' model being used. At a concentration of >350 ppm the citrate represents a significant food resource if it can be utilized by the bacteria. pav
When the E. coli attains the ability to metabolize the citrate, then, and only then, does it become an advantage to the E. coli with this ability, an advantage it exercises over the E. coli that can’t metabolize the citrate. Hence the relative fitness increases.
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.franklin
February 27, 2015
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Here's Michael Behe's take on the Lenski experiment. Why hasn't this been talked about?PaV
February 27, 2015
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wd400:
Well, I guess if you want to make up your own definitions of words that’s fine.
And I guess if you want to define meaningless statements as actually having meaning, you can do that, too.PaV
February 27, 2015
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That’s interesting, because I think your understanding/definition of evolution is wrong.
I'm certainly very confident that one of us doesn't know very much about evolution. You are just wrong to think evolution means "progress" or enough progress to change a species name(!). 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.wd400
February 27, 2015
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franklin:
Why would you suspect that? Citrate is a suitable source for metabolism and being able to utilize both glucose and citrate would be advantageous over a single food source…….and it appears you don’t understand why citrate is included in bacteria culture media…hint it is to supply necessary iron to the bacteria.
Citrate is a suitable source for metabolism in organisms that can metabolize them. But in a minimal growth medium, a "limited" growth medium, where glucose serves as the carbon source, citrate is present at a very low level, and is not the carbon source. When the E. coli attains the ability to metabolize the citrate, then, and only then, does it become an advantage to the E. coli with this ability, an advantage it exercises over the E. coli that can't metabolize the citrate. Hence the relative fitness increases. Now, I've already noted that the growth medium being used was "limited" in the glucose available to all the bacteria experiencing growth, so that, in effect, there is some advantage to be able to metabolize citrate. Now, throw all kinds of glucose at the organism that can metabolize citrate, and, I would suspect the E. coli will return to its wild type. Again, evolution? Run the experiment I'm proposing. Prove me wrong. The rest of your post is just obvious.PaV
February 27, 2015
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wd400:
Your definition of evolution is… wrong.
That's interesting, because I think your understanding/definition of evolution is wrong.
There’s not much point talking abut the mistakes that stem from it.
Do you mean that I deviate from the orthodoxy that evolutionary biologists impose? Is that a correct understanding of what you wrote?
I’m pretty sure this is also misleading:
“The “potentiating” mutations quoted above had the effect of “increasing” the mutation rate”
Think again. I didn't say that the mutation rate of cit+ went up all by itself; I said that the mutation rate went up, which, of course, would mean E. coli could find the necessary mutations for the cit+ easier/faster.PaV
February 27, 2015
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How was it determined that gene duplications are blind watchmaker processes?Joe
February 27, 2015
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Hangonasec:
And if they are already viable and competent, those immediate needs are met, by iterated replication of the current blueprint.
So now you are arguing against evolution. LoL!. How did they get to be viable and competent? Those IC systems that make them so had to come about some way. And the rest of your post is evidence-free wishful thinking.Joe
February 27, 2015
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Piotr:
You start with a primitive function, add more parts, and get a more complex system with new secondary functions (exaptations) which may be selected for and fine-tuned.
And if it takes 5 parts before some primitive function exists? What about systems that require thousands of parts- systems that cannot be built up slowly? Bacterial flagella only have 30-50 different proteins but some of those have thousands of subunits. And you have to avoid cross reactions during formation.Joe
February 27, 2015
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Well, I guess if you want to make up your own definitions of words that's fine. But it makes it hard to talk to others when you do that.wd400
February 27, 2015
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Wd400 asks, Can you?, 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. Cope's rule is not evolution if the changes are the result of innate tendencies. peacefifthmonarchyman
February 27, 2015
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fmm
Are you honestly claiming that because Myoglobin and hemoglobin both handle oxygen that they are the same system?
In order to understand my example you would have to know something about how myoglobin and hemoglobin function. Once you understand that you will clearly see how that example answers your question:
More successful at what? We need to define what we are talking about System(A union B) is not the same system as part A alone
franklin
February 27, 2015
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If a change is not ultimately the result of random variation filtered by natural selection then it is not evolution. Would you claim otherwise?
It may still be evolution, though not evolution by natural selection. (There were once many theories of evolution, after all).
Can you think of any permanent biological change that is not evolution
Can you? The only "permanent" biological change I can think of is extinction, and that's certainly a change in allele frequencies (to zero!)wd400
February 27, 2015
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franklin says, Might be. Myoglobin/hemoglobin and the loading/unloading of oxygen. I say, Are you honestly claiming that because Myoglobin and hemoglobin both handle oxygen that they are the same system? Are a donkey and a pickup truck the same system because they both can carry cargo? peacefifthmonarchyman
February 27, 2015
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Wd400 says, Would you seriosuly argue a change in genotype leading to a change in phenotype is not as example of evolution? I say If a change is not ultimately the result of random variation filtered by natural selection then it is not evolution. Would you claim otherwise? you say, The changes brought about by the same genes expressed in a different environment (so called phentypic plasticity) aren’t evolutionary changes. I say, So at least we agree on one process that is not evolution. That is something I guess. Can you think of any permanent biological change that is not evolution? peacefifthmonarchyman
February 27, 2015
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fmm
More successful at what? We need to define what we are talking about System(A union B) is not the same system as part A alone.
Might be. Myoglobin/hemoglobin and the loading/unloading of oxygen.franklin
February 27, 2015
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I’ll ask you the same question I asked Zac. Is there any natural process that you would not consider evolution?
Countless. If we restrict ourselves to biology and changes in populations (not individuals) the changes brought about by the same genes expressed in a different environment (so called phentypic plasticity) aren't evolutionary changes. Likewise some animals responses to climate change (migration, change in mating effort/time) are probably not evoltionary changes either (just the same genes in new environment). Would you seriosuly argue a change in genotype leading to a change in phenotype is not as example of evolution?wd400
February 27, 2015
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Hangonasec says, If you mean ‘true IC’, a pair of parts neither of which could have ever existed independently of the other, then of course evolution can’t go there. I say, That is not the definition of IC. A system is IC if the system can't exist with out all the parts. The individual parts may or may not exist on their own. Think of an archway the individual stones can exist but a structure is not an archway until all the stones are present and interacting with each other. It is impossible for such a structure to arrive via evolution this should be obvious. you say, Consider the case where part A exists, then part B subsequently arises linked to it, and the pairing proves more successful than A alone. I say More successful at what? We need to define what we are talking about System(A union B) is not the same system as part A alone. This should be obvious. Part A can not "evolve" into system (a union B) by definition. This is not to say that it is not possible for system (A union B) is arise by natural processes only that it can't arise by "evolution" The more parts and the more intricate the union the less likely a system can arise by chance. But an IC system can never arise by evolution. This is unless you define evolution as anything that happens in biology and render intelligent communication impossible. peacefifthmonarchyman
February 27, 2015
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wd400 says, A change in genotype leading a change in phenotype is evolution. I say, Well there you have it. GMOs are evolution. HGC is evolution transposons are evolution. Apparently everything biological is evolution. I'll ask you the same question I asked Zac. Is there any natural process that you would not consider evolution? peacefifthmonarchyman
February 27, 2015
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Zac says, If you mean biological evolution, then anything not biological. and We showed where irreducible complexity can occur in biology through two different processes. I say, So if occurs in biology it is the result of evolution no mater what the process that gave rise to it. There you have it in black and white. According to Zac if it occurs in biology it must be the result of evolution. It should be obvious that given these presuppositions "evolution" is the only game in town. Given this mindset IC structures must arise by evolution because obviously they exist. Talk about circularity peacefifthmonarchyman
February 27, 2015
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wd400
Not the mutation rate generally, just the rate at which the cit+ phenotype arose. In other words, the potentiating mutations don’t change the rate at which the cit+ mutations arise, they just create the background in which cit+ mutations can work so when the turn up they are selected for. Contingency.
If I recall correctly the mutations that are being referenced are the ones that occurred to the DNA repair mechanisms in this specific linage.franklin
February 27, 2015
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PAV
Everything that happened looks to me to have happened in some kind of a relatively ‘ordered’ way.
yes, like point mutations at every possible nucleotide....sometimes more than once...;ordered way;...not so much. pav
The growth medium was “minimal”. What if they grew the Cit+ populations on a glucose rich medium, then what would happen? I suspect the E. Coli would lose the cit+ ability. So, then, we’re back where we started from. And this is evolution?
Why would you suspect that? Citrate is a suitable source for metabolism and being able to utilize both glucose and citrate would be advantageous over a single food source.......and it appears you don't understand why citrate is included in bacteria culture media...hint it is to supply necessary iron to the bacteria. Which is why, in Lenski's expt., the evolution of the ability to transport citrate across the cell membrane, in the presence of oxygen, conveyed an advantage to the cultured bacteria allowing them to utilize the additional food source, i.e., citrate. pav
There was NO selective advantage here because there was enough glucose available and because the mutations, as the authors say, were “nonadaptive.”
there was enough glucose available for the cultures to survive between passages but replication was resource-limited. The evolution of the ability to utilize citrate in the presence of oxygen promoted increased replication due to the ability to utilize citrate as a food resources..franklin
February 27, 2015
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Your definition of evolution is... wrong. There's not much point talking abut the mistakes that stem from it. I'm pretty sure this is also misleading:
"The “potentiating” mutations quoted above had the effect of “increasing” the mutation rate"
Not the mutation rate generally, just the rate at which the cit+ phenotype arose. In other words, the potentiating mutations don't change the rate at which the cit+ mutations arise, they just create the background in which cit+ mutations can work so when the turn up they are selected for. Contingency.wd400
February 27, 2015
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