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Evolution Der Muskelentwicklung Der Malacostraca (Crustacea

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Evolution Der Muskelentwicklung Der Malacostraca (Crustacea Evolution der Muskelentwicklung der Malacostraca (Crustacea) – Vergleichende Untersuchung ausgewählter Vertreter vor dem Hintergrund embryonaler und larvaler Transformationen Kumulative Dissertation zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. nat.) Am Lehrstuhl für Allgemeine und Spezielle Zoologie des Instituts für Biowissenschaften an der Mathematisch-Naturwissenschaftlichen Fakultät der Universität Rostock vorgelegt von Günther Joseph Jirikowski, geboren am 24.11.1983 in Ulm / Baden-Württemberg Rostock, Februar 2015 1 2 Gutachter: 1. Gutachter: Prof. Dr. Stefan Richter, Allgemeine und Spezielle Zoologie, Institut für Biowissenschaften, Universität Rostock 2. Gutachter: Prof. Dr. Gerhard Scholtz, Vergleichende Zoologie, Institut für Biologie, Humboldt-Universität zu Berlin, Datum der Einreichung: 20. Februar 2015 Datum der Verteidigung: 17. April 2015 3 4 Inhaltsverzeichnis 1 Einleitende Zusammenfassung ................................................................................................................................... 7 1.1 Übersicht .............................................................................................................................................................................. 7 1.2 Die Malacostraca .............................................................................................................................................................. 9 1.3 Einauplius und kryptische Larvalentwicklung .................................................................................................... 10 1.4 Entwicklungsmodi der Malacostraca ..................................................................................................................... 12 1.4.1 Larvalentwicklung ....................................................................................................................................................... 12 1.4.1.1 Entwicklung über eine Naupliuslarve .............................................................................................................. 12 1.4.1.2 Entwicklung über eine Zoea-ähnliche Larve ............................................................................ 12 1.4.2 Pseudodirektentwicklung ........................................................................................................... 13 1.4.3 Direktentwicklung ...................................................................................................................... 14 1.5 Heterochronie und Entwicklungsmodus....................................................................................... 14 1.6 Embryogenese der Malacostraca und Entwicklung des Mesoderms ........................................... 16 1.7 Myogenese .................................................................................................................................... 18 1.8 Ziele der Dissertation .................................................................................................................... 19 1.9 Ergebnisse ..................................................................................................................................... 20 1.9.1 Untersuchte Arten ...................................................................................................................... 20 1.9.2 Beschreibung der Myogenese .................................................................................................... 20 1.9.3 Topographie embryonaler Muskulatur ....................................................................................... 21 1.9.4 Vergleichende Analyse des Entwicklungsverlaufes.................................................................... 23 1.9.4.1 Kodierung von Entwicklungssequenzen .................................................................................. 23 1.9.4.2 Heterochronieanalyse ............................................................................................................. 24 1.9.5 Ergebnisse der Heterochronienalyse .......................................................................................... 25 1.10 Evolution der Entwicklung der Malacostraca ............................................................................. 27 1.10.1 Evolution des Entwicklungsmodus ........................................................................................... 27 1.10.2 Evolution der Naupliuslarve und des Einaupliusstadiums ........................................................ 33 1.10.3 Die evolutionäre Bedeutung von Mesoderm und Muskulatur .................................................. 34 1.11 Alternative phylogenetische Hypothesen ................................................................................. 35 1.12 Schlussfolgerungen .................................................................................................................... 35 1.13 Literaturverzeichnis der einleitenden Zusammenfassung ......................................................... 37 2 Erklärung über den Eigenanteil an den Manuskripten ............................................................... 43 3 Selbstständigkeitserklärung ....................................................................................................... 45 4 Danksagung ................................................................................................................................. 46 5 5 Jirikowski GJ, Kreissl S, Richter S, Wolff C. 2010. Muscle development in the marbled crayfish - insights from an emerging model organism (Crustacea, Malacostraca, Decapoda). Development, genes and Evolution, 220:89-105. ....................................................................................................... 47 6 Jirikowski GJ, Richter S, Wolff C. 2013. Myogenesis of Malacostraca – the “egg-nauplius” concept revisited; Frontiers in Zoology, 10:76-103. ............................................................................ 67 7 Jirikowski GJ, Wolff C, Richter S. 2015. Evolution of Eumalacostracan development – new insights into loss and reacquisition of larval stages revealed by heterochrony analysis, Evo Devo, 6:4. .................................................................................................................................................... 97 6 Abbildung 1. Kernfärbung an einem Metanauplius von Artemia franciscana (Branchiopoda, Anostraca) (a) und dem Einaupliusstadium von Neocaridina heteropoda (Malacostraca, Decapoda, Caridea) (b). Maßstabsbalken: 100µm. 1 Einleitende Zusammenfassung 1.1 Übersicht Die Malacostraca (Crustacea) beeindrucken neben der enormen Artenvielfalt und Disparität in Bezug auf adulte Tiere auch durch eine immense Diversität von Entwicklungsmodi, die sich in der Ausbildung unterschiedlichster embryonaler Morphologien oder Larvenformen zeigt. Die phylogenetischen Beziehungen innerhalb der Malacostraca konnten mit den Methoden der Molekularsystematik, trotz der rasanten technischen Fortschritte, bis zum heutigen Tage nicht mit ausreichender Zuverlässigkeit aufgeklärt werden (Jenner et al., 2009). Die gängigen Verwandtschaftshypothesen begründen sich auf morphologischen Merkmalen (Richter & Scholtz, 2001; Wirkner & Richter, 2010; Watling et al., 2000) und wurden wiederholt angezweifelt. Der ursprüngliche Entwicklungsmodus der Gruppe spielt dabei eine zentrale Rolle. Die Mehrzahl der 7 `Crustacea´ entwickelt sich über eine planktische Larve mit drei Extremitätenpaaren (erste Antenne, zweite Antenne und Mandibel), die als Nauplius bezeichnet wird (Abbildung 1a) (Walossek & Müller, 1990; Anderson, 1973; Lauterbach, 1986). Es ist unklar, ob die Naupliuslarve Teil des Grundmusters der Tetraconata (Pancrustacea) ist (Richter, 2002). Innerhalb der Malacostraca besitzen nur zwei Teilgruppen eine Naupliuslarve: die Dendrobranchiata (Decapoda) (Martin et al., 2014) und die Euphausiacea (Martin & Gomez-Gutierrez, 2014). Die Ergebnisse neuerer phylogenetischer Analysen (Edgecombe et al., 2000; Regier et al., 2010; Reumont et al., 2012) deuten darauf hin, dass die Naupliuslarve in der Stammlinie der Malacostraca verlorengegangen ist. Nach der von Richter & Scholtz vorgeschlagenen Phylogenie der Malacostraca (Richter & Scholtz, 2001; Wirkner & Richter, 2010) ist die Naupliuslarve beider Gruppen sekundär innerhalb der Malacostraca entstanden. Scholtz (Scholtz, 2000) diskutiert in diesem Zusammenhang die mögliche Bedeutung des Einauplius (Abbildung 1b), eines Embryonalstadiums, das für das Grundmuster der Malacostraca anzunehmenden ist. Der Einauplius spiegelt die Morphologie der Naupliuslarve insofern wider, dass er nur Anlagen der ersten Antenne, der zweiten Antenne und der Mandibel besitzt. Scholtz (Scholtz, 2000) argumentiert, dass der Einauplius auf ein plesiomorphes Entwicklungsprogramm zurückzuführen ist, das von Vorfahren mit Naupliuslarve übernommen wurde. Zudem führt er aus, dass der Einauplius die Entstehung einer sekundären Naupliuslarve ermöglicht hat und dass für diese Transformation Heterochronien – Veränderungen im zeitlichen Verlauf der Entwicklung – eine fundamentale Rolle gespielt haben. Die vorliegende Arbeit hat das Ziel, die Bedeutung des Einauplius für die Evolution des Entwicklungsmodus der Malacostraca zu prüfen. Dazu soll der Entwicklungsprozess als Ganzes und unter Einbeziehung der zugrundeliegenden zellulären Prozesse der Morphogenese einer vergleichenden Analyse unterzogen werden. Die Entwicklung des Mesoderms, insbesondere der Muskulatur, ist dabei von besonderem Interesse, da sie in der Diskussion
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