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Zinkfingernukleasen – Wikipedia Zum Inhalt springen

Zinkfingernukleasen

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(Weitergeleitet von Zinkfingernuklease)

Zinkfingernukleasen (ZFN) sind künstlich hergestellte Restriktionsenzyme. Sie enthalten eine Zinkfingerdomäne, die an DNA bindet, und eine Nukleasedomäne, welche die DNA schneidet.<ref>{{#invoke:Vorlage:Literatur|f}}{{#if: 

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  }}</ref> Die Zinkfingerdomäne kann so gebaut werden, dass sie eine bestimmte DNA-Sequenz erkennt.

Das bedeutet, dass man mit ZFN ein komplexes Genom an einer ganz bestimmten Stelle schneiden kann und so einen zielgerichteten Einbau von fremder DNA ermöglicht.

Nukleasedomäne

Zinkfingernukleasen enthalten in der Regel die unspezifisch schneidende Nukleasedomäne des Typ-IIS-Restriktionsenzyms FokI<ref name="kim1996">{{#invoke:Vorlage:Literatur|f}}{{#if: 

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  }}</ref>. Diese Restriktionsdomäne ist nur aktiv, wenn sie dimerisiert vorliegt<ref name="bitinaite1998">{{#invoke:Vorlage:Literatur|f}}{{#if: 
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  }}</ref>, weswegen zwei unterschiedlich gebaute ZFN-Monomere benötigt werden, um die DNA-Zielsequenz zu schneiden. Bei Standard-ZFN ist die Restriktionsdomäne über ihr N-terminales Ende an die DNA-bindende Zinkfingerdomäne gebunden. Damit die Nukleasedomänen dimerisieren und schneiden können, müssen die beiden unterschiedlichen ZFN-Monomere an die beiden unterschiedlichen Stränge der DNA binden, wobei ihre C-Termini einen bestimmten Abstand zueinander haben müssen.<ref name=Händel2008>Eva-Maria Händel, Stephen Alwin, Toni Cathomen: Expanding or Restricting the Target Site Repertoire of Zinc-finger Nucleases: The Inter-domain Linker as a Major Determinant of Target Site Selectivity. In: Molecular Therapy. 17, 2008, S. 104–111, {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}.</ref>

Mehrere verschiedene Protein-Engineering-Techniken werden eingesetzt, um einerseits die Enzymaktivität und andererseits Affinität und Spezifität der Zinkfingerdomäne zu steigern. So wurde beispielsweise „Gerichtete Evolution“ angewandt, um FokI-Varianten zu erzeugen, die eine erhöhte Nukleaseaktivität aufweisen.<ref name=Guo2010>Jing Guo, Thomas Gaj, Carlos F. Barbas III: Directed Evolution of an Enhanced and Highly Efficient FokI Cleavage Domain for Zinc Finger Nucleases. In: Journal of Molecular Biology 400, 2010, S. 96–107. {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}</ref> Zudem wurde strukturelles Design eingesetzt, um mittels Austausch geladener Aminosäuren im Dimerisierungsinterface der Nukleasedomäne sogenannte „obligat-heterodimere“ FokI-Varianten zu erzeugen, die eine deutlich erhöhte Restriktionsspezifität aufweisen.<ref name=Szczepek2007>Michal Szczepek, Vincent Brondani u. a.: Structure-based redesign of the dimerization interface reduces the toxicity of zinc-finger nucleases. In: Nature Biotechnology. 25, 2007, S. 786, {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}.</ref><ref name=Miller2007>Jeffrey C Miller, Michael C Holmes u. a.: An improved zinc-finger nuclease architecture for highly specific genome editing. In: Nature Biotechnology. 25, 2007, S. 778–785, {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}.</ref><ref name=Doyon2010>Yannick Doyon, Thuy D Vo u. a.: Enhancing zinc-finger-nuclease activity with improved obligate heterodimeric architectures. In: Nature Methods. 8, 2010, S. 74–79, {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}.</ref><ref name=Ramalingam2010>Sivaprakash Ramalingam, Karthikeyan Kandavelou, Raja Rajenderan, Srinivasan Chandrasegaran: Creating Designed Zinc-Finger Nucleases with Minimal Cytotoxicity. In: Journal of Molecular Biology 405, 2011, S. 630–641. DOI:10.1016/j.jmb.2010.10.043</ref>

DNA-Bindedomäne

Die DNA-Bindedomäne enthält typischerweise zwischen drei und sechs unterschiedliche Zinkfingermotive, wobei jedes individuelle Zinkfingermotiv 3 bp erkennt. Binden die Zinkfingerdomänen perfekt an ihre Erkennungssequenz, so reichen drei Zinkfinger pro ZFN-Monomer aus, um einen bestimmten Lokus in einem komplexen Genom spezifisch zu erkennen. Mehrere verschiedene Strategien wurden entwickelt, um Cys2His2-Zinkfinger herzustellen, die an gewünschte Sequenzen binden.<ref> {{#invoke:Vorlage:Literatur|f}}{{#if: 

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  }}</ref> Diese Methoden umfassen sowohl das modulare Zusammenbauen (siehe unten) als auch Selektionsstrategien, wie das Phagendisplay, das Yeast-1-Hybrid-Systeme, das Bacterial one-hybrid System, das Bacterial two-hybrid System oder zelluläre Selektionssysteme.

Die einfachste Methode, neue Zinkfingerarrays zu erzeugen, ist die Kombination von Zinkfingern mit bekannter Spezifität. Der am weitesten verbreitete modulare Zusammenbauprozess ist das Kombinieren von drei unterschiedlichen Zinkfingern, die jeweils 3 bp erkennen, zu einem neuen Zinkfingerarray, der 9 bp erkennt. Der Hauptnachteil dieser Methode ist, dass sich die Spezifität eines Zinkfingers je nach benachbartem Zinkfinger im Array verändern kann, weshalb "kontextabhängige" Selektionsstrategien üblicherweise Zinkfingerarrays mit einer höheren Spezifität erzeugen.<ref name=Ramirez2008>Cherie L Ramirez, Jonathan E Foley u. a.: Unexpected failure rates for modular assembly of engineered zinc fingers. In: Nature Methods. 5, 2008, S. 374–375, {{#invoke:Vorlage:Handle|f|scheme=doi|class=plainlinks|parProblem=Problem|errCat=Wikipedia:Vorlagenfehler/Parameter:DOI|errClasses=error editoronly|errHide=1|errNS=0 4 10 100}}.</ref>

Anwendungen

Zinkfingernukleasen sind nützlich, um die Genome vieler Pflanzen- und Tierarten zielgerichtet zu verändern, einschließlich der Genome von Arabidopsis<ref name=Zhang2010>{{#invoke:Vorlage:Literatur|f}}{{#if: 

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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Plant Molecular Biology
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Cai|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref><ref name=Townsend2009>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>,

Soja<ref name=Curtin2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 

       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
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{{#invoke:Vorlage:Literatur|f}}{{#if: 

       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Fruchtfliege<ref name=Bibikova2003>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
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               | Vorlage:Webarchiv/archiv-bot
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Bibikova|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Fadenwurm<ref name=Wood2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Wood|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Platynereis dumerilii<ref name=Guehmann2015>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#invoke:TemplatePar|check
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Current Biology
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Seeigel<ref name=Ochiai2010>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Genes to Cells
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Seidenraupe<ref name=Takasua2010>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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   |errNS    = 0
   |template = Vorlage:Cite journal
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Insect Biochemistry and Molecular Biology
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Takasu|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Zebrafisch<ref name=Ekker2008>

{{#invoke:Vorlage:Literatur|f}}{{#if: 

       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
         }}
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Frosch<ref name=Young2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Young|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Maus<ref name=Goldberg2010>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Cell
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Goldberg|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Ratte<ref name=Geurts2009>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Kaninchen<ref name=Flisikowska2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#invoke:TemplatePar|check
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: PLoS ONE
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  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, Schwein<ref name=Hauschild2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#invoke:TemplatePar|check
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Proceedings of the National Academy of Sciences
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  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Hauschild|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>, Rind<ref name=Yu2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
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  }}{{#invoke:TemplatePar|check
   |all    = title=
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Cell Research
     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Yu|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref> und verschiedener Typen von Säugerzellen<ref name=Carroll2008>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
         }}
  }}{{#invoke:TemplatePar|check
   |all    = title=
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     || Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:D. Carroll|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>. Darüber hinaus wurden ZFN in vivo in einem Mausmodell für Hämophilie angewandt<ref name=Li2011>{{#invoke:Vorlage:Literatur|f}}{{#if: 
       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
               | Vorlage:Webarchiv/archiv-bot
         }}
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  }}{{#if: Vorlage:Cite book/ParamBool
        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
  }}</ref>, und eine klinische Studie hat ergeben, dass autologe CD4-positive T-Zellen mit durch Zinkfingernukleasen ausgeschaltetem CCR5-Gen sicher sind, um als Behandlung für HIV/AIDS dienen zu können<ref name=Tebas2014>{{#invoke:Vorlage:Literatur|f}}{{#if: 
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     || Vorlage:Cite book/Meldung
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        | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
       | Vorlage:Cite book/Meldung
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:Tebas|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}</ref>. Alternativ zu Zinkfingernukleasen können Transcription Activator-like Effector Nucleases und die CRISPR/Cas-Methode verwendet werden.

Weblinks

Einzelnachweise

<references />

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