Unser Haushund: Eine Spitzmaus im Wolfspelz? - Wolf-Ekkehard ...

Unser Haushund: Eine Spitzmaus im Wolfspelz? - Wolf-Ekkehard ... Unser Haushund: Eine Spitzmaus im Wolfspelz? - Wolf-Ekkehard ...

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164zwischen den Hunderassen bestätigen die Aussage Trumlers – und selbst das"Rauschen" auf der DNA-Ebene mit ihrer Vielzahl an SNPS und CNVs fügt sichdem Degenerationsthema ein.Und zu Dawkins wiederhole ich with some supplementary remarks andinferences sein Statement wie folgt: "Bear in mind this order of evolutionary change[from Wolf to Chihuahua due to a series of genetic and anatomical dysfunctions], andthen extrapolate backwards twenty thousand times as far into the past. It becomesrather easy to accept that evolution” [and now the correct inference] from a fish intoa human has never happened at all. And all the parallel losses offunctions/deterioration of integration on the genetical and anatomical levels in thefox domestication experiment reinforce this conclusion."Aber ist es denn wirklich so, dass durch die großen Zahlen an Duplikationenüberhaupt keine positiven Veränderungen für den betroffenen Organismus aufallen oder zumindest mehreren Ebenen (siehe oben) hervorgerufen werdenkönnen?" – wird man an dieser Stelle vielleicht einwenden.Sehen wir uns dazu einige als positiv betrachtete Duplikationen etwas näher an.Zunächst bei Bakterien. Erst kürzlich (September 2012) wurde das Paper von Z.D. Blount, J. E. Barrick, C. J. Davidson und R. E. Lenski: Genomic analysis of akey innovation in an experimental Escherichia coli population als bedeutenderBeitrag zum Verständnis von evolutionary novelties u. a. durch Genduplikationengefeiert (Nature 489: 513-518, 27 September 2012 288 ). Hier das Abstract derAutoren:"Evolutionary novelties have been important in the history of life, but their origins are usually difficult toexamine in detail. We previously described the evolution of a novel trait, aerobic citrate utilization(Cit(+)), in an experimental population of Escherichia coli. Here we analyse genome sequences toinvestigate the history and genetic basis of this trait. At least three distinct clades coexisted for more than10,000 generations before its emergence. The Cit(+) trait originated in one clade by a tandemduplication that captured an aerobically expressed promoter for the expression of a previously silentcitrate transporter. The clades varied in their propensity to evolve this novel trait, although genotypesable to do so existed in all three clades, implying that multiple potentiating mutations arose during thepopulation's history. Our findings illustrate the importance of promoter capture and altered gene regulationin mediating the exaptation events that often underlie evolutionary innovations."Michael J. Behe hat am 13. November 2012 zu dieser "evolutionarenInnovation" durch Genduplikationen, promoter capture and altered generegulation einen sehr gut durchdachten Kommentar unter dem Titel Rose-ColouredGlasses, Lenski, Citrate, and Biologos verfasst, aus dem ich im Folgenden einigeHauptpunkte zitieren möchte (zu weiteren Fragen vgl. man den Originalbeitrag). 289"….the Lenski lab observed a mutant strain in their experiments that could metabolize citrate in thepresence of oxygen, which unmutated E. coli cannot do. (Blount et al. 2008) (Importantly, however, thebacterium can metabolize citrate in the absence of oxygen.) This allowed the mutated bacterium tooutcompete its relatives, because the growth medium contained a lot of citrate, as well as oxygen. It wasan intriguing result, and was touted as a major innovation, but at the time Lenski's lab was unable to trackdown at the DNA level the exact mutations that caused the change.Now they have. In a recent publication in Nature (Blount et al. 2012) they report the multiple mutations288 http://www.ncbi.nlm.nih.gov/pubmed/22992527289 Siehe http://www.evolutionnews.org/2012/11/rose-colored_gl066361.html

165that confer and increase the ability to transport citrate in an atmosphere containing oxygen. They divide themutations conceptually into three categories: 1) potentiation; 2) actualization; and 3) refinement. "Actualization"is the name they give to the mutation that first confers a weak ability to transport citrate into the laboratory E.coli. (It turns out that the bacterium is lacking only a protein to transport citrate into the cell in the presence ofoxygen; all other enzymes needed to further metabolize citrate are already present.) The gene for the citratetransporter, citT, that works in the absence of oxygen is directly upstream from the genes for two other proteinsthat have promoters that are active in the presence of oxygen. A duplication of a segment of this regionserendipitously placed the citT gene next to one of these promoters, so the citT gene could then beexpressed in the presence of oxygen. 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.Over time the mutant got better at utilizing citrate, which the authors called "refinement." Further workshowed this was due to multiple duplications of the mutant gene region, up to 3-9 copies. Again, geneduplication is a fairly common process, so again it is unsurprising. In another experiment Lenski and coworkersshowed that increasing the concentration of the citrate transporter gene was sufficient in itself toaccount for the greater ability of E. coli to grow on citrate. No other mutations were needed.A more mysterious part of the whole process is what the group called "potentiation." It turns out that theoriginal E. coli they began with decades ago could not benefit from the gene duplication that broughttogether a citT gene with an oxygen-tolerant promoter. Before it could benefit, a preliminary mutationhad to occur in the bacterium somewhere other than the region containing the citrate-metabolismgenes. Exactly what that mutation was, Lenski and coworkers were not able to determine. However, theyexamined the bacterium for mutations that may contribute to potentiation, and speculated that "A mutationin arcB, which encodes a histidine kinase, is noteworthy because disabling that gene upregulates thetricarboxylic acid cycle." (They tried, but were unable to test this hypothesis.) In other words, the"potentiation" may involve degradation of an unrelated gene.D. h. auch in den Untersuchungen des Lenski-Labors mit Billionen von Bakterienüber mehr als 50.000 Generationen konnten keine (durch Zufallsmutationenentstandenen) neuen funktionalen DNA-Sequenzen (Gene) für die Bildung völligneuer Proteine nachgewiesen werden – wie viel geringer ist dann erst dieAussicht, dass sich durch diesen Mechanimus völlig neue Gene im Zuge derKurzzeit-Domestikation des Grauen Wolfs gebildet hätten!Abschließend stellt Behe in seinem Beitrag fest:"In my own view, in retrospect, the most surprising aspect of the oxygen-tolerant citT mutation was that itproved so difficult to achieve. If, before Lenski's work was done, someone had sketched for me a cartoon ofthe 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 toshow that even skeptics like myself overestimate the power of the Darwinian mechanism."Aber immerhin: Durch diese Genduplikationen ist eine metabolische Veränderung(metabolic change) für ein neues environment induziert worden 290 . Das zeigt, dasseine Anpassung an veränderte Umweltbedingungen durch diesen (Zufalls-)Mechanismus möglich zu sein scheint (siehe Einschränkung in der Fußnote) – wennauch erst durch den Einsatz gewaltiger Zahlen von Individuen und Generationen 291 ,die bei Säugetieren sowie bei der Domestikation des Grauen Wolfs [“a few wolves gaveup their freedom in exchange for our garbage”] in wenigen tausend Jahren gar nichtgegeben sind und damit für diese entfällt.Unter der bisher unbewiesenen Voraussetzung, dass vielleicht einige wenigeCNVs mit Genduplikationen beim Hund (und zwar CNVs, die beim Grauen Wolf290 - Eine Umwelt zwar, die – soweit ich das bisher verstehen kann –, so in der Natur so überhaupt nicht existiert und durch einen Prozessüber mehrere Stufen, der ohne die ganz gezielt eingesetzte menschliche Hilfe nie zustande gekommen wäre. Mit anderen Worten: So theyprepared E. coli for a totally artificial environment in which the bacteria will never occur and - even if these surroundings existed in the wild -would never survive inasmuch as several steps with billions of bacteria over thousands of generation would be necessary to cope with that strangehabitat/milieu which would consist of extraordinary amounts of citrate and oxygen. (Auf die Frage "Is their any natural environment where theability to transport citrate in an atmosphere containing oxygen would be useful for E. coli?” Antwortete mir Michael J. Behe am 20. 11. 2012: "Ifthere is such an environment, I haven't heard of it and Lenski hasn't mentioned it. He has taken pains in his papers to say that the inability to usecitrate in the presence of oxygen is a species-defining trait for E. coli.”)291 Insbesondere auch die Zahl der Generationen extrapoliert für die letzten 3 Milliarden Jahre.

165that confer and increase the ability to transport citrate in an atmosphere containing oxygen. They divide themutations conceptually into three categories: 1) potentiation; 2) actualization; and 3) refinement. "Actualization"is the name they give to the mutation that first confers a weak ability to transport citrate into the laboratory E.coli. (It turns out that the bacterium is lacking only a protein to transport citrate into the cell in the presence ofoxygen; all other enzymes needed to further metabolize citrate are already present.) The gene for the citratetransporter, citT, that works in the absence of oxygen is directly upstream from the genes for two other proteinsthat have promoters that are active in the presence of oxygen. A duplication of a segment of this regionserendipitously placed the citT gene next to one of these promoters, so the citT gene could then beexpressed in the presence of oxygen. 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.Over t<strong>im</strong>e the mutant got better at utilizing citrate, which the authors called "refinement." Further workshowed this was due to multiple duplications of the mutant gene region, up to 3-9 copies. Again, geneduplication is a fairly common process, so again it is unsurprising. In another exper<strong>im</strong>ent Lenski and coworkersshowed that increasing the concentration of the citrate transporter gene was sufficient in itself toaccount for the greater ability of E. coli to grow on citrate. No other mutations were needed.A more mysterious part of the whole process is what the group called "potentiation." It turns out that theoriginal E. coli they began with decades ago could not benefit from the gene duplication that broughttogether a citT gene with an oxygen-tolerant promoter. Before it could benefit, a prel<strong>im</strong>inary mutationhad to occur in the bacterium somewhere other than the region containing the citrate-metabolismgenes. Exactly what that mutation was, Lenski and coworkers were not able to determine. However, theyexamined the bacterium for mutations that may contribute to potentiation, and speculated that "A mutationin arcB, which encodes a histidine kinase, is noteworthy because disabling that gene upregulates thetricarboxylic acid cycle." (They tried, but were unable to test this hypothesis.) In other words, the"potentiation" may involve degradation of an unrelated gene.D. h. auch in den Untersuchungen des Lenski-Labors mit Billionen von Bakterienüber mehr als 50.000 Generationen konnten keine (durch Zufallsmutationenentstandenen) neuen funktionalen DNA-Sequenzen (Gene) für die Bildung völligneuer Proteine nachgewiesen werden – wie viel geringer ist dann erst dieAussicht, dass sich durch diesen Mechan<strong>im</strong>us völlig neue Gene <strong>im</strong> Zuge derKurzzeit-Domestikation des Grauen <strong>Wolf</strong>s gebildet hätten!Abschließend stellt Behe in seinem Beitrag fest:"In my own view, in retrospect, the most surprising aspect of the oxygen-tolerant citT mutation was that itproved so difficult to achieve. If, before Lenski's work was done, someone had sketched for me a cartoon ofthe 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 toshow that even skeptics like myself overest<strong>im</strong>ate the power of the Darwinian mechanism."Aber <strong>im</strong>merhin: Durch diese Genduplikationen ist eine metabolische Veränderung(metabolic change) für ein neues environment induziert worden 290 . Das zeigt, dasseine Anpassung an veränderte Umweltbedingungen durch diesen (Zufalls-)Mechanismus möglich zu sein scheint (siehe Einschränkung in der Fußnote) – wennauch erst durch den Einsatz gewaltiger Zahlen von Individuen und Generationen 291 ,die bei Säugetieren sowie bei der Domestikation des Grauen <strong>Wolf</strong>s [“a few wolves gaveup their freedom in exchange for our garbage”] in wenigen tausend Jahren gar nichtgegeben sind und damit für diese entfällt.Unter der bisher unbewiesenen Voraussetzung, dass vielleicht einige wenigeCNVs mit Genduplikationen be<strong>im</strong> Hund (und zwar CNVs, die be<strong>im</strong> Grauen <strong>Wolf</strong>290 - <strong>Eine</strong> Umwelt zwar, die – soweit ich das bisher verstehen kann –, so in der Natur so überhaupt nicht existiert und durch einen Prozessüber mehrere Stufen, der ohne die ganz gezielt eingesetzte menschliche Hilfe nie zustande gekommen wäre. Mit anderen Worten: So theyprepared E. coli for a totally artificial environment in which the bacteria will never occur and - even if these surroundings existed in the wild -would never survive inasmuch as several steps with billions of bacteria over thousands of generation would be necessary to cope with that strangehabitat/milieu which would consist of extraordinary amounts of citrate and oxygen. (Auf die Frage "Is their any natural environment where theability to transport citrate in an atmosphere containing oxygen would be useful for E. coli?” Antwortete mir Michael J. Behe am 20. 11. 2012: "Ifthere is such an environment, I haven't heard of it and Lenski hasn't mentioned it. He has taken pains in his papers to say that the inability to usecitrate in the presence of oxygen is a species-defining trait for E. coli.”)291 Insbesondere auch die Zahl der Generationen extrapoliert für die letzten 3 Milliarden Jahre.

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