Interleukin-21

{{#if: | {{#if: | {{#if:6005| {{#if: 133 Aminosäuren | {{#if: | {{#if: | {{#if: (155 aa) | {{#if: IL-21, IL-21iso | {{#if: IL21Q9HBE4605384 | {{#if: IL21 | {{#if: Q9HBE4605384 | {{#if: | {{#if:| {{#if: | {{#if: | {{#if: | {{#if: | {{#if: | {{#if: | {{#if: | {{#if: | {{#if: Wirbeltiere{{#if: Q9HBE4 | {{#ifeq: | NV ||1}}|}}| {{#if: Q9HBE4 | {{#ifeq: | NV || | style="background:#C3FDB8; color:#000000;" | Homologie-Familie {{#if: Wirbeltiere | | style="background:#C3FDB8; color:#000000;" | Übergeordnetes Taxon {{#if: | | style="background:#C3FDB8; color:#000000;" | Ausnahmen {{#if: | }}

{{#if: | | Interleukin-21 }}
Interleukin-21 }}
Bändermodell vom Interleukin-21 des Menschen (von zwei Seiten), nach PDB 2OPQ
Interleukin-21 }}
Andere Namen	{{{Andere Namen}}} }}
Vorhandene Strukturdaten: }}
Eigenschaften des menschlichen Proteins }}
Masse/Länge Primärstruktur	133 Aminosäuren }}
Sekundär- bis Quartärstruktur	}}
Kofaktor	}}
Präkursor	(155 aa) }}
Isoformen	IL-21, IL-21iso }}
Bezeichner
{{#if: | Gen-Namen | Gen-Name}} | Gen-Name(n) }}	{{#if:6005 | IL21 | IL21}}{{#if: |, }} }}
Externe IDs	{{#if: | GeneCards: Vorlage:GeneCards}}{{#if:605384 | OMIM: 605384}}{{#if:Q9HBE4| {{#if: | | UniProt }} {{#if:|{{{titel}}}|Q9HBE4}}{{#if:|Vorlage:Abrufdatum}}}}{{#if: | MGI: [1]}}{{#if: | PubChem: {{#switch: {{{PubChem}}}		0 | }}	- = –	valid|1|2=/^[1-9]%d?%d?%d?%d?%d?%d?%d?%d?$/	0 | Wikipedia:Vorlagenfehler/Vorlage:PubChem}}	template= Vorlage:PubChem	format= }} }}{{#invoke:TemplatePar|check	all= 1=	opt= 2=	0 | Wikipedia:Vorlagenfehler/Vorlage:PubChem}}	template= Vorlage:PubChem	format= }}{{#if: {{#invoke:Wikidata|pageId}}||{{#ifeq: 0 | 0 | }} }}}}{{#if: | CAS-Nummer:  }}{{#if: | }} }} }}
Arzneistoffangaben
ATC-Code	}} {{#if:|
DrugBank	[2] }} {{#if:|
Wirkstoffklasse	}} }}
Transporter-Klassifikation
TCDB	{{{TCDB}}} {{#if: |
Bezeichnung	{{{TranspText}}} }} }}
{{#if:|Inhibitorklassifikation|{{#if:|Enzymklassifikationen|Enzymklassifikation}}}}
EC, Kategorie	{{#if:| {{{EC-Nummer}}}}}{{#if: |, [[{{{Kategorie}}}]]}} }}
MEROPS	{{{Peptidase_fam}}}}}
MEROPS	{{{Inhibitor_fam}}}}}
Reaktionsart	{{{Reaktionsart}}} }}
Substrat	{{{Substrat}}} }}
Produkte	{{{Produkte}}} }} }}
Vorkommen
{{#if:		{{{Homolog_fam}}}|Hovergen}}]	{{{Homolog_fam}}}|Hovergen}}]}} }} }}
Wirbeltiere }}
}} }}

{{#if:Interleukin-21-2OQP.png||}}

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Interleukin-21 (IL-21) ist ein Interleukin, welches bei Menschen durch das IL21-Gen codiert wird.<ref name="pmid11081504">{{#invoke:Vorlage:Literatur|f}}{{#if:

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Interleukin-21 ist ein Zytokin mit bedeutenden regulatorischen Effekten auf Zellen des Immunsystems, darunter natürliche Killerzellen (NK-Zellen), B-Zellen und zytotoxische T-Zellen, die mit Viren infizierte oder Krebszellen zerstören können.<ref name="pmid11081504" /><ref name="pmid12429707">{{#invoke:Vorlage:Literatur|f}}{{#if:

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  }}</ref> Dieses Zytokin induziert Zellteilung und -wachstum in seinen Zielzellen.

Gen

Das menschliche IL-21-Gen ist etwa 8,43 kb groß, ist auf dem Chromosom 4 lokalisiert und ist an 180 kb des IL-2-Gens gebunden. Das mRNA-Produkt ist 616 Nukleotide lang.<ref name="pmid11081504" /><ref name="entrez" />

Verteilung in Geweben und Zellen

IL-21 findet sich in aktivierten menschlichen CD4⁺ T-Zellen, aber nicht in den meisten anderen Geweben.<ref name="pmid11081504" /> Zudem ist IL-21 das Hauptzytokin für T follikuläre Helferzellen hat wichtige Effekte in der B Zellproliferation, Antikörperinduktion und -maturation.<ref name="pmid15210760">{{#invoke:Vorlage:Literatur|f}}{{#if:

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    | Vorlage:Cite book/Meldung
  }}</ref> Weiterhin wird IL-21 in natürlichen Killer-T-Zellen (NK-T-Zellen) gebildet und reguliert deren Funktion.<ref>{{#invoke:Vorlage:Literatur|f}}{{#if: 
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Interleukin-21 wird auch von Krebszellen des Hodgkin-Lymphoms (HL) gebildet. Diese Beobachtung könnte einen Großteil des Verhaltens von klassischen Hodgkin-Lymphomen erklären, einschließlich der Ansammlungen von anderen Immunzellen um HL-Zellen in Zellkulturen. IL-21 steht damit im Fokus bei der Erforschung von potenziellen Behandlungsansätzen oder zumindest der Entwicklung eines Tests auf HL.<ref name="blood-112-8">{{#invoke:Vorlage:Literatur|f}}{{#if:

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Klinische Bedeutung

Allergien

IL-21R-Knockout-Mäuse zeigen höhere Niveaus von IgE und geringere von IgG1 als Wildtyp-Mäuse, nachdem sie dem Antigen ausgesetzt waren. Die IgE-Niveaus gingen nach Injektion von IL-21 zurück. Dies ist bedeutend für die Rolle, die IL-21 bei der Kontrolle allergischer Reaktionen spielt, und zwar wegen der Rolle von IgE bei Typ-1-Überempfindlichkeitsreaktionen.<ref name="pmid12446913">{{#invoke:Vorlage:Literatur|f}}{{#if:

       | {{#if: Vorlage:Cite book/ParamBool
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    | Vorlage:Cite book/Meldung
  }}</ref> Mit IL-21 wurden Therapieversuche durchgeführt, um allergische Reaktionen zu vermindern. Es konnte gezeigt werden, dass damit proinflammatorische Zytokine, die von T-Zellen produziert werden, vermindert werden konnten, und bei an allergischer Rhinitis (allergischem Schnupfen) leidenden Mäusen konnten ebenfalls IgE Niveaus gesenkt werden.<ref name="pmid17982108">{{#invoke:Vorlage:Literatur|f}}{{#if: 
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  }}</ref> Eine Studie an Mäusen mit Erdnuss-Allergie zeigte, dass die systemische Behandlung mit IL-21 der allergischen Reaktion effektiv entgegenwirkte.<ref name="pmid18056403">{{#invoke:Vorlage:Literatur|f}}{{#if: 
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       | Vorlage:Cite book/Meldung
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  }}</ref> Dies hat große Bedeutung für die pharmakologische Entwicklung von IL-21-basierten Medikamenten gegen lokale und systemische Allergien.

Immuntherapie bei Krebs

IL-21 wurde für klinische Versuche der Phase 1 bei Patienten mit metastasierenden Melanomen (MM) und Nierenzellkarzinom (renal cell carcinoma, RCC) freigegeben. Es wurde dafür den Behörden gegenüber mit lediglich grippeartigen Nebenwirkungen als sicher genug dargestellt. Dosis-limitierende Toxizität wurde über geringe Anzahlen von Lymphozyten, Neutrophilen und Thrombozyten sowie anhand der Leberwerte ermittelt. Nach den Kriterien für Behandlungseffekte bei festen Tumoren (Response Evaluation Criteria in Solid Tumors, RECIST) zeigten 2 von 47 MM-Patienten und 4 von 19 RCC-Patienten umfassende oder teilweise Effekte. Weiterhin wurden Anstiege von Perforin, Granzym B, IFN-γ und CXCR3 mRNA in peripheren NK-Zellen und CD8⁺ T-Zellen festgestellt. Dies legt nahe, dass IL-21 die CD8⁺-Effektor-Funktionen fördert und hierdurch zu einer Anti-Tumor-Antwort führt. Mit IL-21 wurden danach zu Klinische Studien der Phase 2 durchgeführt, wo es alleine oder im Verbund mit Medikamenten wie Sorafinib und Rituximab verabreicht wurde.<ref name="pmid19845910">{{#invoke:Vorlage:Literatur|f}}{{#if:

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Virusinfektionen

IL-21 ist möglicherweise ein kritischer Faktor bei der Kontrolle persistenter Virusinfektionen. IL-21- (oder IL-21R-) Knockout-Mäuse, die mit dem Virus der chronischen Lymphozytischen Choriomenginitis (LCM-Virus, LCMV) infiziert waren, konnten diesen im Gegensatz zu Wildtyp-Mäusen nicht loswerden. Außerdem zeigten diese Knockout-Mäuse mit beschädigtem IL-21-Signalweg weitaus dramatischere Ausschüttungen von LCMV-spezifischen, zytotoxischen T-Zellen, was nahelegt, dass von CD4+-T-Helferzellen produziertes IL-21 für nachhaltige CD8+-Effektoraktivität der T-Zellen nötig ist.<ref>{{#invoke:Vorlage:Literatur|f}}{{#if:

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    | Vorlage:Cite book/Meldung
  }}</ref> Es ist somit wahrscheinlich, dass IL-21 am Mechanismus beteiligt ist, mit dem CD4+-T-Helferzellen die Antwort des Immunsystems auf virale Infektionen orchestrieren.

Von HIV-Infizierten wurde berichtet, dass IL-21 die HIV-spezifische, zytotoxische Antwort der T-Zellen, sowie die NK-Zellen-Funktion verbessert.<ref name="white et al">{{#invoke:Vorlage:Literatur|f}}{{#if:

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     || 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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     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:Iannello A, Boulassel MR, Samarani S, Tremblay C, Toma E, Routy JP, Ahmad A.|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
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  }}</ref> Es konnte auch gezeigt werden, dass HIV-spezifische CD4-T-Zellen von Langzeitstabilen (seltene Fälle, bei denen sich die AIDS-Erkrankung nicht ausbildet, obwohl sie nicht behandelt werden) in der Lage sind, signifikant mehr IL-21 zu produzieren als die Mehrzahl der Menschen.<ref name="chevalier et al" /> Zudem wurden bei den Langzeitstabilen häufiger IL-21-produzierende, virusspezifische CD8-T-Zellen gefunden.<ref name="williams et al">{{#invoke:Vorlage:Literatur|f}}{{#if: 
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     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:Williams L.D., Bansal A., Sabbaj S., Heath S.L., Song W., Tang J., Zajac A.J., Goepfert P.A.|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
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  }}</ref> Diese Daten und die Tatsache, dass IL-21-stimulierte CD8- oder NK-Zellen in der Lage waren, die Virusreplikation von HIV in vitro zu unterbinden<ref name="chevalier et al" /><ref name="Iannello et al" /> zeigen, dass dieses Zytokin möglicherweise für Anti-HIV-Therapeutika genutzt werden kann.

Einzelnachweise

<references />

Weitere Literatur

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

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     | Vorlage:Cite book/Meldung
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        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
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               | Vorlage:Toter Link/archivebot
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        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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    | Vorlage:Cite book/Meldung
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• {{#invoke:Vorlage:Literatur|f}}{{#if:

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               | Vorlage:Toter Link/archivebot
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        | Vorlage:Cite book/Meldung
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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
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:Sivori S, Cantoni C, Parolini S, et al.|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}

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

       | {{#if: Vorlage:Cite book/ParamBool
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  }}{{#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
  }}{{#if: Vorlage:Cite book/ParamBool
     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:Pène J, Gauchat JF, Lécart S, et al.|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | 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/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: J. Leukoc. Biol.
     || 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:Strengell M, Julkunen I, Matikainen S|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | 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/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: Xi Bao Yu Fen Zi Mian Yi Xue Za Zhi
     || 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:Zhang SQ, Chen B, Luo X, Xu CZ|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
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• {{#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: Genome Res.
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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:^^|^^||+1}}{{#ifeq:Gerhard DS, Wagner L, Feingold EA, et al.|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
  }}

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

       | {{#if: Vorlage:Cite book/ParamBool
               | Vorlage:Toter Link/archivebot
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  }}Vorlage:Cite book/URL{{#if:  | Vorlage:Cite book/Meldung }}{{#if:        | Vorlage:Cite book/Meldung }}{{#if: J. Immunol.
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  }}{{#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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    | 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
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        | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
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     | Vorlage:Cite book/Meldung
  }}Vorlage:Cite book/Meldung2{{#ifexpr: 0{{#ifeq:^^|^^||+1}}{{#ifeq:^^|^^||+1}}{{#ifeq:Mehta DS, Wurster AL, Weinmann AS, Grusby MJ|^^||+1}}{{#ifeq:^^|^^||+1}} > 1
    | Vorlage:Cite book/Meldung
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