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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
#33 @Piotr " ...it was an intelligently designed process?" For the long past changes we can't generally reconstruct who first came up with some linguistic variation and why. You would need to find who did it first and ask them how did they come up with it. But we can see the process going on today on the web where new words are routinely coined via deliberate process. Note also that intelligent process generating novelty need not be conscious in all its steps. In fact the most revolutionary ideas and the most productive intelligent processes are subconscious, in one's sleep or just ideas that work themselves out in the background and suddenly pop into one's consciousness while doing something else. But even in such cases, the discoverer had often spent great deal of time and effort on the problem, then left it alone, even gave up, then the solution came to him out of a blue as it were.nightlight
February 25, 2015
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Exactly the same kind of statistical analysis can be done on evolution of languages or technologies, even though we recognize that intelligent processes were guiding those changes.
Interesting. So when, for example, speakers of mainstream British English began to drop their final or preconsonantal /r/'s regularly in words like CAR and CARD in the eighteenth century, it was an intelligently designed process? Or when speakers of Old Latin began to rhotacise intervocalic /s/'s, which led to odd alternations like FLOS/FLORIS, was it deliberate design on their part?Piotr
February 25, 2015
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#29 @wd400 "It’s a pretty clear demonstration that benificial mutations can arise without design (unless, I guess, organisms can see into the uture!)" Nothing in that or any other experiment demonstrates that mutations were "random" or "aimless". Exactly the same kind of statistical analysis can be done on evolution of languages or technologies, even though we recognize that intelligent processes were guiding those changes. The evolution of languages is in fact reconstructed in the same way as evolution of biological organisms via measurements of distances, linguistic or genetic. As explained in the previous post, there is plenty of clearly evident intelligence in the cells to account for design of the needed biochemical changes in DNA. Cellular biochemical networks are far smarter in problem domain of molecular design and bioengineering that all the human science and technology put together.nightlight
February 25, 2015
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WD400: Have you read the paper? Or a summary of it? There are lots of mutations, so which have been fixed in different populations, some of those by selection. And there’s ongoing change in mean fitness. That’s evolution.
I'll offer you my summary: an already available gene needs to be next to a nearby promoter to express its protein. By way of gene duplication it gets in place. The end. Behold the power of evolution! How many generations did this take?Box
February 25, 2015
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What he’s saying is akin to claiming that because humans can breathe, humans breathing underwater would not be a new function
Land artiodactyls ---> protocetids ---> whales Whew! still mammals. Trivial post-flood variation within a kind. (I actualy heard this argument from Robert Byers.)Piotr
February 25, 2015
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Nighlight, Have you heard of the Luria Delbruk experiment? It's a pretty clear demonstration that benificial mutations can arise without design (unless, I guess, organisms can see into the uture!)wd400
February 25, 2015
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The Lenski experiment does nothing other than to show one particular preexisting strain became dominant when an artificial set of environmental conditions were forced on them. No evolution took place.
Have you read the paper? Or a summary of it? There are lots of mutations, so which have been fixed in different populations, some of those by selection. And there's ongoing change in mean fitness. That's evolution.wd400
February 25, 2015
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#15 CHartsil
"Where exactly did the experiment demonstrate that the mutations or up/down regulations needed were "random" or "aimless" i.e. that the mechanism was neo-Darwinian?" Where it could not be repeated prior to the 20,000 generation
How many generations of humans it took to intelligently design cars, airplanes, computers, smart phones...? Intelligence does not imply or require instant success or no failures. Some problems are hard and require many intelligent, yet unsuccessful tries before they are solved. Even mearly realizing that there is a solvable problem and formulating it can take many generations for otherwise intelligent beings. Hence, the number of generation before the needed capability is produced does not demonstrate neo-Darwinian mechanism for novelty generation i.e. that it was "random" or "aimless" mutation that generated it. The cellular biochemical networks may have well been seeking to optimize their biochemistry to the environment. It just took them that many generations to figure out how to get there from what they had initially. James Shapiro calls this intelligent process "natural genetic engineering" (NGE). More generally, the cellular biochemical networks are the same kind of distributed self-programming computers as the networks of neurons making human brains. From the research of neural networks, which are mathematical models of such adaptable networks, it is known that when exposed to punishments and rewards, the create and run anticipatory algorithms to optimize their net 'rewards - punishments' utility function. These algorithms look ahead, try various actions in their internal model space, follow them up and evaluate consequences, then pick the action in the real world that yields the optimum value for their utility function.
"But in all cases where the novelty generating mechanism is understood, it is a purposeful, intelligent process." False equivocation and affirming the consequent. In technology, objects that are not subject to replication with variation are designed or improved upon with known, testable mechanisms of design.
Merely calling new E.coli tries "replication with variation" does not demonstrate that the process was "aimless" or "random". The early PCs had 2MHz single core processors and 4K of RAM, 160K floppy disk. Today, billions of "copies with variation" (if you wish to call it that way) of PCs later, they have 4 or more cores per PC, each running several thousand times faster with tens of thousands more RAM and permanent storage (also accessible thousands times faster). There is no essential difference in the requirement and availability of intelligent processes for novelty generation between these "copies with variation" of PCs and "copies with variation" of E. coli. The main difference is the particular implementation of the distributed computers (cellular biochemical networks for E. coli vs networks of neurons forming human brains for PC evolution) and their anticipatory algorithms. But each process was intelligent (in the sense of creating and running anticpatory problem solving algorithms) in its own problem domain.
In technology, objects that are not subject to replication with variation are designed or improved upon with known, testable mechanisms of design.
You are defending "randomness of gaps" (ROG) theory -- just because our present science hasn't yet figured out how to "decompile" (reverse engineer) the algorithms used by the cellular biochemical networks to generate novelty, then the process is declared "random" and "aimless". That is as anti-scientific as 'god of gaps' since it amounts to saying -- move on, nothing to see beyond this point, it is just some "random" and "aimless" chemical reactions that did it, end of story. In fact, neo-Darwinian dogma was repeatedly the major impediment and science stopper for decades, among others in the research of epigenetics and "junk" DNA. While we don't know yet how to decompile (reverse engineer) anticipatory algorithms for most real world adaptable networks, such as brain or cellular biochemical networks, that is possible with our mathematical models (abstractions) of such distributed computers, neural networks. Such decompilation of internal algorithms produced by neural networks (when exposed to abstract punishments & rewards) shows that the adaptable networks create internal models of their environment, with self actor (representing the 'system itself' in the model space) and the model environment, which may also include other intelligent actors. The algorithms (which are self-programmed by the network in the unsupervised learning mode) then run these self actors in the model space forward in model time, just like chess player moving chess pieces several moves forward in his mental model of the chess position, and evaluating resulting final positions in order to decide which move to play on the real chess board. Note that J. Shapiro's NGO represents the decompilation only of few low level disjoint "subroutines" of the algorithms performed by the cellular biochemical networks. Over time, the higher level functions that "call" these "subroutines" will be figured out, then gradually the entire anticipatory algorithms tying them all together will be decompiled as well. The neo-Darwinist priesthood will fight this progress every step of the way, just as they fought epigenetic and "junk" DNA research and discoveries. Eventually, though, as always they will quietly submit while morphing their marvelously flexible language, finally declaring, yes, of course, that's what we actually meant all along.
"Why is that well demonstrated intelligence dogmatically excluded (without any empirical evidence)" That which is asserted without evidence can be summarily dismissed without evidence.
There is vast amount of evidence for intelligence of cellular biochemical networks. Start for example with engineering and synthesizing of new live cells from 'scratch' (from simple molecules used as 'food'). Why don't you try doing it in the lab without using live cells or their components, but just from basic 'food' and 'light' that say photosynthetic bacteria use to synthesize new cells. Not only you couldn't do it, but you could take all the world's best biochemists and molecular biologists together, in one all star team, give them all the resources they ask for, and they won't be able to synthesize from scratch one live organelle, let alone the whole live cell from scratch. All you will get for the investment will be hundreds of conferences, thousands of papers and millions of words, but not a single live cell to show. Yet, while you were reading the above paragraph, your own cellular biochemical networks have achieved such mind-boggling feats of molecular scale bioengineering thousands of times. One cell in your little toe knows more physics, chemistry and biochemistry needed for molecular scale bioengineering than all the world's scientists and sciences put together. Of course, needless to mention the vastly greater intelligence needed to construct the live cells plus whole live organism such as animal, plant or human. Our science is light years behind of even dreaming about such feats of molecular engineering. So, why is this plainly evident (e.g. from cell reproduction to ontogeny) intelligence in the cells excluded by the neo-Darwinian dogma as the designer and engineer of the genetic novelty behind evolutionary phenomena? Note that existence of biological evolution is the strongest argument for Intelligent Design since it takes far more underlying intelligence to engineer and build a system that not only functions well in the initial environment, but that changes over time to adapt to new environments.nightlight
February 25, 2015
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The Lenski experiment does nothing other than to show one particular preexisting strain became dominant when an artificial set of environmental conditions were forced on them. No evolution took place. If this is the best you've got you're in trouble.humbled
February 25, 2015
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"What exactly was “removed” from the system?" The potentiating mutations prior to the 20,000th generation. Also, if either the duplicate gene or the new regulator is removed, the cit* ceases functioning. "if the bacteria “cease[d] functioning,” then how did it “evolve”? That is, if it ceased functioning, then it would die. Dead things don’t “evolve.”" It's talking about individual systems, not the organisms in which the systems reside.CHartsil
February 25, 2015
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CHartsil: This is NOT an IC system. In the Discovery response, they point this out. Here's the definition being used: "“a system comprised of interdependent parts, the removal of any of which will cause the system to cease functioning.” So, two questions: (1) What exactly was "removed" from the system? and (2) if the bacteria "cease[d] functioning," then how did it "evolve"? That is, if it ceased functioning, then it would die. Dead things don't "evolve."PaV
February 25, 2015
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"That is a correct statement which obviously does not imply that it takes only one mutation to get there. In fact it doesn’t say anything about the number of mutations needed." >Only http://www.merriam-webster.com/dictionary/only >alone in a class or category : existing with no other or others of the same kindCHartsil
February 25, 2015
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CHartsil,
Box: What do you mean ‘wrong’? So it took two mutation to get next to the promoter. And? Where did Behe (or anyone) deny that?”
CHartsil: >only needs a promoter that functions in the presence of oxygen.
That is a correct statement which obviously does not imply that it takes only one mutation to get there. In fact it doesn't say anything about the number of mutations needed.Box
February 25, 2015
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"What do you mean ‘wrong’? So it took two mutation to get next to the promoter. And? Where did Behe (or anyone) deny that?" >only needs a promoter that functions in the presence of oxygen.CHartsil
February 25, 2015
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CHartsil: At least two potentiating mutations were also necessary as a genetic drift building block to the duplication and new regulation. Not that it matters. If you have a Roman arch built with a scaffold, then remove the last support, it's only a single step, but the final product is irreducibly complex. The question isn't how many steps are involved, but whether the final product is both complex and irreducible.Zachriel
February 25, 2015
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CHartsil #18,
CHartsil: Again, wrong. At least two potentiating mutations were also necessary as a genetic drift building block to the duplication and new regulation.
What do you mean 'wrong'? So it took at least two mutations to get next to the promoter. And? Where did Behe (or anyone) deny that?Box
February 25, 2015
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"It might have been more accurate for Behe to have stated that the gene for the citrate transporter, citT, only needs a promoter that functions in the presence of oxygen." Again, wrong. At least two potentiating mutations were also necessary as a genetic drift building block to the duplication and new regulation. "Did we witness an evolutionary miracle? A whole new protein? Are we talking irreducible complexity? Of course not" Did a system evolve that serves a function which requires multiple interacting components? Yes Define IC for us againCHartsil
February 25, 2015
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bFast: By the most minimal definition of IC, I think this qualifies. Is this a 2 or a 3 mutational step event? In any case, it is within the range of what Behe describes as “The Edge of Evolution.” Irreducible complexity is normally defined as a complex and irreducible system. The derived citrate system fits this definition. The question is whether such a system can evolve. And it clearly can.Zachriel
February 25, 2015
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CHartsil #12,
Behe: It turns out that the bacterium is lacking only a protein to transport citrate into the cell in the presence of oxygen; all other enzymes needed to further metabolize citrate are already present.
CHartsil: He also gets this wrong. The metabolization actually comes from the new regulator conferred by the head-to-tail duplication.
It might have been more accurate for Behe to have stated that the gene for the citrate transporter, citT, only needs a promoter that functions in the presence of oxygen. IOW the gene for the citrate transporter protein is already present it just lacks a promotor - which is also present as we will see later. In fact Behe is too kind, for the gullible reader it may seem as if a whole new protein is needed.
Blount et al: One reason that E. coli cannot grow aerobically on citrate is its inability to transport citrate24,31,32. The origin of the Cit+ phenotype therefore required expression of a citrate transporter.
Behe: The gene for the citrate transporter, citT, that works in the absence of oxygen is directly upstream from the genes for two other proteins that have promoters that are active in the presence of oxygen.
Okay, so the gene for the citrate transporter needs to get near one of those promotors. How?
Behe: A duplication of a segment of this region serendipitously placed the citT gene next to one of these promoters, so the citT gene could then be expressed in the presence of oxygen.
Did we witness an evolutionary miracle? A whole new protein? Are we talking irreducible complexity? Of course not:
Behe: Gene duplication is a type of mutation that is known to be fairly common, so this result, although requiring a great deal of careful research to pin down, is unsurprising.
Box
February 25, 2015
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"Where exactly did the experiment demonstrate that the mutations or up/down regulations needed were “random” or “aimless” i.e. that the mechanism was neo-Darwinian?" Where it could not be repeated prior to the 20,000 generation "But in all cases where the novelty generating mechanism is understood, it is a purposeful, intelligent process." False equivocation and affirming the consequent. In technology, objects that are not subject to replication with variation are designed or improved upon with known, testable mechanisms of design. "What is the essence of the distinction between the E.coli experiment and all other well understood instances of evolution that demonstrates “aimlessness” or “randomness” in the case of E. coli experiment?" Can you refine this question? "Why is that well demonstrated intelligence dogmatically excluded (without any empirical evidence)" That which is asserted without evidence can be summarily dismissed without evidence.CHartsil
February 25, 2015
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There are four points that I would like to make in this conversation. By the most minimal definition of IC, I think this qualifies. Is this a 2 or a 3 mutational step event? In any case, it is within the range of what Behe describes as "The Edge of Evolution." It was noted that the precursor to this success offered the organism advantage. When the precursor offers disadvantage, that's when the IC case gets interesting. The e-coli in this case were offered EXTREME reward for pulling off this example of minimal IC. It would appear to me that most natural systems would not be so generous with the size of the reward. It only took 50,000 generations to get there. If the LCA of human and chimp was about 5 million years ago, and if the human lineage averaged 10 years per generation (close enough) then there are about 500,000 generations in in the human lineage from the LCA of human and chimp. So this small IC event took 1/10 that generational time. The HAR1F took on 18 mutations in that period of time. As the HAR1F is ultra-conserved, It is clear that no individual point mutation in the edited region of the HAR1F offered even neutral effects. In fact, based upon the folding of the HAR1F it is difficult to imagine any less than 6 deleterious mutations were required before a positive one could have come about. This one transaction alone is far beyond the scope of the Darwinian prowess demonstrated in Lenski’s E. Coli.bFast
February 25, 2015
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@CHartsil -- Where exactly did the experiment demonstrate that the mutations or up/down regulations needed were "random" or "aimless" i.e. that the mechanism was neo-Darwinian? Evolution is not just a characteristic of biological systems. It is even more obvious in human technologies, sciences, languages, religions, cultures,... But in all cases where the novelty generating mechanism is understood, it is a purposeful, intelligent process. What is the essence of the distinction between the E.coli experiment and all other well understood instances of evolution that demonstrates "aimlessness" or "randomness" in the case of E. coli experiment? Note that you can't say that there is no "intelligence" in the cells. After all cellular biochemical networks know/understand physics, chemistry and biochemistry deeply enough to be able to engineer new cells or organisms from scratch (from simple atoms and molecules), something that human best and brightest scientists are not even remotely capable of. Hence, we do have plentiful, well demonstrated intelligence of the cellular biochemical networks which is far beyond anything human scientists could show. Why is that well demonstrated intelligence dogmatically excluded (without any empirical evidence) as the most likely intelligence that generated the observed novelties and the novelty mechanism is declared to be "random" or "aimless" mutation?nightlight
February 25, 2015
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He appears to have just made up the idea of 'functional coded elements' as an ad hoc rationalization to explain away the clear cut case of the addition of information by Lenski's E. coli He also gets this wrong "It turns out that the bacterium is lacking only a protein to transport citrate into the cell in the presence of oxygen; all other enzymes needed to further metabolize citrate are already present" The metabolization actually comes from the new regulator conferred by the head-to-tail duplication. What he's saying is akin to claiming that because humans can breathe, humans breathing underwater would not be a new functionCHartsil
February 25, 2015
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Here is Behe's response in 2012. http://www.evolutionnews.org/2012/11/rose-colored_gl066361.htmlCollin
February 25, 2015
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"Does Lenski’s experiment tell us anything about how rarely IC systems come about by chance in nature and the probabilities involved for changes to species that multiply far slower than bacteria?" Well, it could be repeated pretty much at will after the generation containing the (at least) two potentiating mutations.CHartsil
February 25, 2015
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CHartsil, That's very interesting. I'm not sure how others view IC, but to me it has always been a probabilities argument. I never thought that IC systems were impossible, but merely improbable, and when a system's complexity increased, so did the likelihood that it could not have evolved by chance. So perhaps an arch, an IC system with two legs and a keystone, may have evolved over a given period of time, 10 arches on top of each other might not have done so. Does Lenski's experiment tell us anything about how rarely IC systems come about by chance in nature and the probabilities involved for changes to species that multiply far slower than bacteria? Perhaps like hominids to humans in a relatively brief number of generations?Collin
February 25, 2015
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"The system must have multiple co-dependent PARTS. " It does, if either the new regulatory module or the new duplicate gene are gone, it can't metabolize citrate aerobically.CHartsil
February 25, 2015
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"It was dependent on multiple changes in multiple systems and cannot function without all of said changes. It’s IC" I don't think that's the exact definition of an IC system. The system must have multiple co-dependent PARTS. Not multiple, perhaps linear, changes that must occur before some system functions. They must be concurrently functioning codependent PARTS that, the removal of which, would cause the system to cease functioning. Is that the case here?Collin
February 25, 2015
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"Evolution is not directional? Just the other day we read about research that says evolution likes em getting bigger" Certain traits are favored in a given environment. That does not mean evolution has a goal in mindCHartsil
February 25, 2015
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Evolution is not directional? Just the other day we read about research that says evolution likes em getting bigger....Andre
February 25, 2015
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