Zentralismus und Komplexität drucken

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Zentralismus versagt in der Kontrolle hochkomplexer Systeme. Ein eindrucksvolles Beispiel aus den Regulationsmechanismen der Natur. Die Ineffizienz zentraler Kontrollen ist uns allen aus unserem täglichen Leben bekannt. In der Natur ist das Problem der Regulation komplexer Systeme, wie der Genexpression in höheren Organismen, durch ein Zusammenspiel von zentraler und dezentralisierter Kontrolle gelöst.

Peter Schuster für den Science-Blog

Die Aussage, daß eine zentrale Kontrolle großer, komplexer Einheiten zum Scheitern verurteilt ist, stellt eine Binsenweisheit dar. Wirtschaft und Gesellschaft untermauern die Gültigkeit dieser Aussage durch zahllose Beispiele in der Vergangenheit und Gegenwart, welche beweisen: Systeme werden ineffizient, sobald sie eine kritische Größe überschreiten. Bereits in frühen Zeiten waren sich kluge Herrscher dieses Problems bewußt und haben dem eine Strategie des „divide et impera“ entgegengesetzt.

Die Natur scheint eine elegante Lösung für die Regulation ihrer großen, hochkomplexen Systeme gefunden zu haben, indem sie modulare Strukturen verwendet und die Module darin partielle Autonomie besitzen. Das beste Beispiel dafür ist der Vielzeller-Organismus, in welchem jede einzelne Zelle über gerade so viel Autonomie in ihrem Stoffwechsel verfügt, als toleriert werden kann ohne dem gesamten System Schaden zuzufügen: Ein wenig mehr Unabhängigkeit für somatische Zellen und unlimitiertes Wachstum – ein Tumor – kann entstehen.

Zentrale Kontrolle in Bakterien

Die Regulierung von Stoffwechsel und Vermehrung der einfachsten Lebensformen, der Bakterien, wird noch zentral gesteuert. Um möglichst effizient zu bleiben, sich möglichst effizient zu vermehren, ist den Bakterien in der Größe ihres Genoms aber eine Obergrenze gesetzt. Wie aus der Sequenzierung von mehr als tausend bakteriellen Genomen hervorgeht, ist es die darin enthaltene Zahl an regulatorischen Genen, welche die Größe des Genoms limitiert [1, 2]: Bakterien mit einem kleinen Genom von einigen hundert Genen besitzen nur wenige dieser regulatorischen Gene. Mit wachsender Größe des Genoms nimmt deren Zahl aber in einer quadratischen Funktion zu (Abbildung 1) und führt schließlich dazu, daß jedes der nicht-regulatorischen Gene – welche ja für die in Stoffwechsel und Vermehrung essentiellen Proteine kodieren - ein eigenes regulatorisches Gen zur Steuerung besitzt. Es ist evident, daß eine weitere Steigerung dieses „Überwachungs-/Kontrollsystems“ zu „kostspielig“ wird, eine maximale Größe erreicht ist.


Abbildung 1. Die Zahl der regulatorischen Gene im bakteriellen Genom wächst mit dessen Größe im quadratischen Ausmaß.
Die Zahl der Gene im Genom und die Zahl der Regulatorgene sind im logarithmischen Maßstab dargestellt. Meßwerte in Grün stammen von Archäbakterien, in Blau von Bakterien. Bild modifiziert nach Croft et al., [2]. Bitte beachten Sie, daß der Maßstab der senkrechten Achse logarithmisch ist.

Zusammenspiel von zentraler Kontrolle und dezentralisierter Regulation in höheren Organismen

Eukaryonten – das sind alle höheren Organismen von der Hefe bis zum Menschen – benutzen für die Regulation ihrer wesentlich komplexeren Systeme der Genexpression ein Zusammenspiel von zentraler Kontrolle und dezentralisierter Regulation:

  • Zentral ist dabei die genetische Kontrolle auf der DNA-Ebene, d.h. auf der Ebene der Übersetzung der in den Genen festgelegten, noch rohen Baupläne für Proteine.
  • Dezentralisiert erfolgt die Regulation der in die Vorläufer-Messenger RNA (pre-mRNA) kopierten (transkribierten) Baupläne durch alternatives Auseinanderschneiden (splicing) und Zusammenfügen von Abschnitten der pre-mRNA zu reifen Messenger-RNAs, den für die Synthese der Proteine verwendeten Vorlagen (siehe unten, Abbildung 2) und
  • Eine Reihe von epigenetischen Regulationsmechanismen greifen am Gen selbst oder in späteren Phasen der Übersetzung regulierend ein.

Die Erforschung des Zusammenwirkens dieser Mechanismen ist neuesten Datums, basierend auf der im Human Genome (HUGO) Projekt erfolgten Entschlüsselung des menschlichen Genoms, d.h. aller Gene, welche die Baupläne für die den Aufbau und Stoffwechsel unseres Organismus bestimmenden Proteine enthalten. Damals, im Jahr 2001, war die Verwunderung groß, als beim Menschen mit rund 22 000 für Proteine kodierenden Genen nicht viel mehr Gene als beispielsweise beim Wurm Caenorhabditis (rund 19000 Gene) festgestellt wurden und diese insgesamt nur rund 1,5 % des gesamten Genoms ausmachten.

Das ENCODE-Projekt

Welche Funktion haben nun die restlichen – nicht für Proteine kodierenden – 98,5 % der DNA, sind diese nicht – wie häufig angenommen - zum Großteil nur Müll (junk DNA)? Ein riesiges internationales Team von 442 Forschern hat sich mit dem „Encyclopedia of DNA Elements (ENCODE)-Projekt“ die systematische Erkundung aller funktionellen Elemente der DNA zum Ziel gesetzt und vor wenigen Tagen die ersten Ergebnisse in rund 30, öffentlich zugänglichen Publikationen veröffentlicht [2]. Demnach konnten bereits zu rund 80 % der DNA biochemische Funktionen zugeordnet werden, die direkt – zentral - am Chromosom oder indirekt über RNA-Transkripte, die nicht in Proteine übersetzt werden und über weitere Mechanismen regulieren wann, in welchen Zelltypen, unter welchen Bedingungen und in welchem Ausmaß Gene abgelesen und in Proteine übersetzt werden.

Für unsere Frage nach zentraler und dezentraler Regulierung kommt dabei besondere Bedeutung dem Mechanismus zu, mit dem die ursprünglich in der genetischen Information vorhandenen Baupläne in funktionelle Proteine übersetzt werden, dem Vorgang des alternativen Splicing der pre-mRNA (siehe oben).

Hier wendet die Natur erfolgreich ein Modul-Schema an (Abbildung 2): Dasselbe Stück DNA, das von einem Gen in eine pre-mRNA transkribiert wird, kann durch unterschiedliches Herausschneiden und Zusammenfügen diverser Abschnitte zur Bildung mehrerer reifer mRNA Moleküle und in der Folge zu unterschiedlichen Proteinen führen. Wie ENCODE zeigt, folgt diese Expression keiner minimalistischen Strategie: abhängig vom individuellen Organismus und den Zelltypen/Geweben in welchen die Expression stattfindet, entstehen aus einer pre-mRNA simultan bis zu 10 – 12 Isoformen, wobei jeweils zumindest zwei Isoformen vorherrschen.


Abbildung 2. Genexpression und Alternatives Spleißen in eukaryontischen Zellen:
Aus ein und demselben Gen, das in eine pre-mRNA kopiert (transkribiert) wurde, entstehen durch Alternatives Spleißen dieser pre-mRNA mehrere unterschiedliche reife mRNAs und aus diesen durch Proteinsynthese an den Ribosomen (Translation) eine Reihe verschiedener Protein-Isoformen.

Zweifellos ist das von ENCODE gezeichnete Bild noch ein sehr unvollständiges, und wir müssen auf Überraschungen gefasst sein. Was aber als gesichert angesehen werden kann, ist, daß es Eukaryonten gelungen ist, durch dezentrale Regulierungsmechanismen – einem „divide et impera“ - die Limitierung in der Anzahl ihrer Gene aufzuheben und höchstkomplexe Systeme zu steuern.

Zentrale – dezentrale Kontrolle: Versuch einer ökonomischen Betrachtung

Um eine grobe Abschätzung der unterschiedlichen „Kosten“ von zentraler versus dezentraler Kontrolle zu treffen, wollen wir ein zentral reguliertes, 10 000 Gene enthaltendes Genom eines Bakteriums vergleichen mit einem gleich großen, virtuellen System, das aber aus 10 lokal regulierten Genomen zu je 1000 Genen bestehen soll:

Im Fall der zentralen Organisation werden – wie aus Abbildung 1 ableitbar - rund 1200 Regulatorgene benötigt [1, 2]. Im dezentralen System reicht dagegen ein kleines Zentrum - beispielsweise mit 180 Regulatorgenen (jeweils 4 Gene für die insgesamt 45 paarweisen Wechselwirkungen) – aus um die Aktivitäten der 10 Untersysteme kontrollieren, je 12 Gene pro Untersystem sollten für dessen Regulation genügen und jeweils 2 „Kontrolloren“ für die Meldungen aus den Untersystemen an das Zentrum. In Summe könnte damit das skizzierte dezentralisierte Modell mit insgesamt 320 Regulatorgenen - einem Viertel des Bedarfs des zentralen Modells - sein Auslangen finden.

Fairerweise muß man dazu anmerken, daß bei einem Einsparen von rund ¾ der Kontrolloren im dezentralisierten Modell auch einiges an Information verloren gegangen ist. Der Großteil der Kontrollfunktionen wird lokal geregelt, das „Nachrichten-Hauptquartier“ erhält gerade soviel an Information, wie für das Management der Wechselwirkungen zwischen den Untersystemen absolut erforderlich ist und kann damit kaum Planungen für die Zukunft des gesamten Systems anstellen. Derartige Konzepte können ja kaum ohne ein umfassendes Bild des gesamten Umfeldes und der daraus abgeleiteten Schlußfolgerungen erstellt werden.

Der Darwin`sche Mechanismus einer Optimierung durch Variation und Selektion braucht sich aber um die Zukunft nicht zu kümmern, da er ja nach einem Prinzip von „trial and error“ vorgeht. Für sich allein existierende Vielzeller-Organismen können damit leicht mit einem Mindestmaß an zentraler Kontrolle und „Zukunftsplanung“ auskommen. Natürlich ändert sich die Situation, wenn es um Gesellschaften im Tier- und Menschenreich geht. Erziehung und Lernen des einzelnen Individuums verändern hier den Darwinischen Mechanismus in einen Lamarckschen: Information, die ein einzelnes Mitglied der Gesellschaft erhalten hat, kann schnell und weitgehend auch an zukünftige Generationen weitergegeben werden, zentralisierte Prognosen über zukünftige Entwicklungen liegen daher im Interesse der Gesellschaft. Ein Zuviel an Föderalismus, ein Zuwenig an zentraler Steuerung ermöglichen hier kaum eine erfolgreiche Planung für die Zukunft. Dazu möchte ich nur ein zur Zeit wichtiges Beispiel anführen: das Versagen einer weltweiten Reduktion der Kohlendioxyd-Emissionen trotz eindeutigen Nachweises von dessen Notwendigkeit.

Schlußfolgerung aus akademischer Sicht

Die Ineffizienz zentraler Kontrollen kennen wir alle aus unserem täglichen Leben. Wenn man an einer großen Universität mit mehr als 80 000 Studenten arbeitet, möchte man die Lehre aus dem Beispiel der Bakterien ziehen, daß die Größe eines Systems limitiert ist, das zentral verwaltet werden kann. Für eine dezentralisierte Universitätsverwaltung sprechen viele Gründe, einer davon liegt darin die Kosten für die „Overheads“ gering zu halten, da diese ja die ohnehin stark begrenzten Mittel der Forschungsgelder dramatisch reduzieren:

Wir sollten daher auf unsere Forschungstätigkeit fokussiert sein und nicht darauf unsere Dekane und Rektoren permanent über Ideen und Planungen zu unterrichten, die sich möglicherweise nicht realisieren lassen, wenn wir - wie es ja unsere Aufgabe ist - ins Neuland vorstoßen. Der Erfolg dieses Vorgehens läßt sich an den Veröffentlichungen in der wissenschaftlichen Literatur sehr gut ablesen und bietet damit die Basis für Evaluierungen durch die zentrale Verwaltung. In den meisten anderen Belangen sollte sich diese aber heraushalten und kann damit so klein wie möglich gehalten werden.

Einzelnachweise und Quellen:

[1] EV Koonin, YI Wolf (2008) Genomics of bacteria and archaea: the emerging dynamic view of the prokaryotic world. Nucleic Acids Res. 36:6688-6719.

[2] Croft, L.J.; Lercher, M.J.; Gagen, M.J.; Mattick, J.S.(2003) Is prokaryotic complexity limited by accelerated growth in regulatory overhead? Genome Biology, 5:P2

[3] The ENCODE Project Consortium. An integrated encyclopedia of DNA elements in the human genome. Nature 489: 57- 74. 6 September 2012

Glossar:

DNA: Desoxyribonucleinsäure; ein üblicherweise in Form eines Doppelstranges (Doppelhelix) vorliegendes Polymer, welches die primäre genetische Information aller Zellen enthält.

RNA: Ribonucleinsäure. Polymere, welche unterschiedliche biochemische Funktionen ausüben können, in der Umsetzung von genetischer Information (Genexpression) ebenso wie als katalytisch aktive Moleküle.

Pre-mRNA: primäres Transkript (Umschreibung) eines zu einem Gen gehörenden Abschnitts der DNA in ein mRNA-Molekül, durch alternatives Splicen entsteht daraus die messenger-RNA (mRNA).

Alternatives Splicen: Abschnitte der pre-mRNA, die keine kodierende Informationen enthalten (Introns), werden entfernt und die verbleibenden Abschnitte (Exons) in unterschiedlicher Weise miteinander verbunden. Damit können aus ein und demselben Gen/derselben pre-mRNA mehrere verschiedene reife mRNA-Moleküle entstehen und aus diesen mehrere unterschiedliche Proteine.

Weiterführende links:

ENCODE: Encyclopedia Of DNA Elements (video 4,4 min)

ENCODE, the Encyclopaedia of DNA Elements, is the most ambitious human genetics project to date.

ENCODE: Encyclopedia of DNA Elements (video 6,5 min)

Ewan Birney of EMBL-EBI, Tim Hubbard of the Wellcome Trust Sanger Institute and Roderic Guigo of CRG talk about ENCODE, an international project which revealed that much of what has been called 'junk DNA' in the human genome is actually a massive control panel with millions of switches regulating the activity of our genes.

Read more about this at: http://www.embl.org/press/2012/120905_Hinxton.

 

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  437. Markus Trullus (kein Partner)
    25. Dezember 2012 16:24

    Die Frage nach Zentralismus oder subsidiärer Struktur ist in wirtschaftlichen Organisationen nicht eindeutig zu beantworten. In Stadien der Ideenfindung/ Innovation ist es das dezentrale, bei Preisgestaltung und Fertigung eher noch die der zentralen Steuerung, also eine Anpassung an die jeweiligen Rahmenparameter, die höhere Effizienz bringt- wobei ich "systemisch" schon alle Parameter beiziehen möchte. Ich frage micht,ob das nicht auch in evolutionären Strukturen ein wichtiges Teil sein kann. Oder noch anders: Kontrolle zentral, Regelung dezentral? Für alle Fälle? Sicher nicht!

  438. settembrini
    12. Oktober 2012 23:42

    Die zentrale Organisation hat uns heute einen Erfolg beschert, wie dies nur von zentraler Stelle möglich ist:

    Wir alle sind zu Nobelpreisträgern geworden. Jeder von uns hat einen 2 Milliardsten Anteil an dem Preis.

    Das nenne ich wahre Verteilungsgerechtigkeit!

    Pace, pace e gioia!

  439. Romig
    20. September 2012 10:56

    Allen Respekt, Herr Professor Schuster!
    In der Soziologie, Politologie und Nationalökonomie sprechen wir vom "Gesetz der kleinen Gemeinschaften" (Othmar Spann, Walter Heinrich, Leopold Kohr, E. Schuhmacher u.v.a.). Für die Kath. Soziallehre wurde tragend der Begriff der "Subsidiarität", der sich scharf gegen den Zentralismus wendet.
    Es gehört zu den großen Geheimnisssen und Wundern der Schöpfung, dass zwischen Natur und Gesellschaft, wie Schuster nachweist, "Entsprechungen" oder Korrelationen herrschen, welche auf "überwölbende"Prinzipien und Normen hinweisen. Alles "Sein" ist also normgeprägtes oder "gesolltes Sein" Nach Kelsen entstehen Normen nur durch einen "Willensakt" eines Legislators, mithin eines Schöpfers der Normen oder der lex aeterna. Benedikt XVI. konnte in seiner berühmten Rede vor dem Deutschen Bundestag vor fast genau einem Jahr auf das wissenschaftliche Ethos Kelsens hinweisen, der noch im Alter von 84 Jahren ("auch da kann man noch etwas Vernünftiges sagen") die Lehre vom Dualismus von Sein und Sollen endgültig aufgegeben habe. Deshalb, schreibt Waldstein (der vom Papst in seiner Rede gleich dreimal zitiert wird)!, "hat Kelsen Recht, dass Naturrecht nur als der Sinn des Willens einer Gottheit verstanden werden kann" (Wolfgang Waldstein: Ins Herz geschrieben. Naturrecht als Fundament eier menschlichen Gesellschaft, St. Ulrich Verlag, Augsburg 2010, S. 19). Die angesprochene Korrelation zwischen Natur und menschlicher Gesellschaft läßt also den Schluß auf die Existenz (oder Überexistenz) Gottes zu, ein Schluß, der allerdings nur besagt, "dass Gott ist", nicht "was er ist". Die Ausformung der ursprünglichen Gotterkenntnis zum "Was" bestimmt nach Samuel Huntington die unterschiedlichen Kulturen.

    • darwin (kein Partner)
      28. September 2012 09:11

      Die Entwicklung unterschiedlichster Systeme, von einfachen zu immer komplexeren Strukturen verläuft offensichtlich ungerichtet - also "ungesollt" -, hätte durch minimale Änderungen der Rahmenbedingungen auch anders ablaufen oder gar nicht eintreten können.

      Dieser Prozess eines "trial and errors" in physikalischen, biologischen und darüber hinaus gesellschaftlichen Systemen läßt sich an Hand gleicher mathematischer Formulierungen modellieren. Derartige Modelle zeigen eindrucksvoll das "Wie", sind aber zweifellos ungeeignet darüber hinaus gehende philosophische Fragen zu beantworten.

  440. terbuan
    15. September 2012 18:01

    Lieber Peter Schuster,
    es war faszinierend und gleichzeitig erschreckend für mich als Laien, diese Erkenntnisse aus den Naturwissenschaften mit den Strukturen unserer Gesellschaft zu vergleichen, die Ähnlichkeiten sind verblüffend.
    Ein zentralistisches System wie beispielsweise in der Sowjetunion ist genauso zum Tode verurteilt wie in der Natur!
    Und so ein System wollen sie uns jetzt gerade in der EU aufzwingen, das kann nicht gut enden!

  441. settembrini
    13. September 2012 23:09

    "Die Ineffizienz zentraler Kontrollen ist uns allen aus unserem täglichen Leben bekannt"

    Nun werden wir dazu vermutlich ein Lehrbeispiel sehen, wenn die EZB sich anschickt 6000 Banken zu kontrollieren. Ob diese dann einen Kontrollor pro kleinere Bank und mehrere Kontrollore pro größere Bank einsetzt?

    Wie uns der Blog-Artikel zeigt, würde dies bereits bei den einfachsten Lebensformen, den zentralistisch organisierten Bakterien, nicht funktionieren. Diese wären mit einer solchen Zahl an Regulatoren heillos überfordert, ihre daraus entstehende völlige Ineffizienz würde in einem Desaster enden.

    Es ist keine Frage, warum die im Laufe der Evolution entstandenen, immer komplexeren Lebewesen nicht zu einer alleinseligmachenden zentralen Kontrolle zurückgekehrt sind.

    Wagen wir eine Prognose, wie das EZB-Projekt enden wird?

    • vermiculus (kein Partner)
      15. September 2012 11:28

      Ein beklemmendes Beispiel, das Sie da bringen, Hr. settembrini!

      Um mir unter diesem Licht ein etwas besseres Einschlafen zu ermöglichen, möchte ich einen Gedanken entgegen halten: nämlich den Begriff der "Kontrolle". (Ob die Wörter "Kontrolle" und insbesondere "Kontrolleure" wohl mit "Kon-" und "Troll" zu haben? Aber Spaß beiseite.)

      Solange nur kontrolliert im Sinne von "überwacht" (zB zum Estellen von Ratings oder, haha, weiterer Stresstests) wird, aber kein Versuch unternommen, auf laufende Aktionen des Sysems Einfluss zu nehmen, muss nicht unbedingt ein großes Risiko von so einer Kontrollbank ausgehen.

      Denn ein Regelkreis besteht ja aus den Elementen Messstrecke und Regelstrecke, und aus dem unvermeidlichen zeitlichen Versatz zwischen Messung und Regelung kann ein Überschwingen bzw. Umkippen des Systems entstehen. Wird auf die Regelstrecke verzichtet bzw. ist keine vorgesehen, so bleibt de facto ein dezentrales Bankensystem erhalten.

      Bleibt also nur zu hoffen, dass das, was da entstehen soll, eine Kontrollbank im Sinne einer "Überwachungsbank", also eines Alarmsystems, und nicht einer "Steuerungsbank" ist.






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