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Comparative Neuroanatomy Within the Context of Deep Metazoan Phylogeny

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Comparative Neuroanatomy Within the Context of Deep Metazoan Phylogeny Comparative Neuroanatomy within the Context of Deep Metazoan Phylogeny Von der Fakultät für Mathematik, Informatik und Naturwissenschaften der Rheinisch-Westfälischen Technischen Hochschule Aachen zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften genehmigte Dissertation vorgelegt von Diplom-Biologe Carsten Michael Heuer aus Düsseldorf Berichter: Herr Universitätsprofessor Dr. Peter Bräunig Herr Privatdozent Dr. Rudolf Loesel Tag der mündlichen Prüfung: 08.03.2010 Diese Dissertation ist auf den Internetseiten der Hochschulbibliothek online verfügbar. Summary Comparative invertebrate neuroanatomy has seen a renaissance in recent years. Highly conserved neuroarchitectural traits offer a wealth of hitherto largely unexploited characters that can make valuable contributions in inferring phylogenetic relationships in cases where phylogenetic analyses of molecular or morphological data sets yield trees with conflicting or weakly supported topologies. Conversely, in those cases where robust phylogenetic trees exist, neuroanatomical features can be mapped onto the trees, helping to shed light on the evolution of the central nervous system. This thesis aims to provide detailed neuroanatomical data for a hitherto poorly studied invertebrate taxon, the segmented worms (Annelida). Drawing on the wealth of investigations into the architecture of the brain in different arthropods, the study focuses on the identification and description of possibly homologous brain centers (i.e. neuropils) in annelids. The thesis presents an extensive survey of the internal architecture of the brain of the ragworm Nereis diversicolor (Polychaeta, Annelida). Based upon confocal laser scanning microscope analyses, the distribution of neuroactive substances in the brain is described and the architecture of two major brain compartments, namely the paired mushroom bodies and the central optic neuropil, is characterized in detail. It is concluded that the central optic neuropil cannot be confidently homologized with similarly unpaired neuropils in the arthropod brain, but that annelid and arthropod mushroom bodies are probably homologues. This proposed homology is further explored by comparing 3D reconstructions of mushroom body neuropils and associated structures in the polychaete species Nereis diversicolor, Harmothoe areolata, and Lepidonotus clava with a 3D model of the mushroom bodies in the insect representative Leucophaea maderae. The neuropils are found to share a common principal organization and a similar neuroarchitectural integration. Lastly, the occurrence of unpaired midline neuropils, mushroom bodies, and associated structures is investigated in a broad taxonomic survey, including more than 20 representatives from major groups of the annelid radiation. Considerably complex brains, sometimes comprising mushroom bodies and other subcompartments, are only observed in errant polychaetes but not in sedentary polychaete species, nor in clitellates representatives. The implications of an assumed homology between annelid and arthropod mushroom bodies are discussed in light of the ‘new animal phylogeny’. It is concluded that the homology of mushroom bodies in distantly related groups has to be interpreted as a plesiomorphy, pointing towards a considerably complex neuroarchitecture inherited from the last common ancestor, Urbilateria. Within the annelid radiation, the lack of mushroom bodies in certain groups is explained by wide-spread secondary reductions owing to selective pressures unfavorable for the differentiation of elaborate brains. Evolutionary trajectories of mushroom body neuropils in errant polychaetes remain enigmatic. Contents Introduction ................................................................................................. ..9 Deep Metazoan Phylogeny – old views and new hypotheses ...................................... .11 The contribution of neurophylogeny ........................................................................... .13 Arthropod neuroarchitecture outlined .......................................................................... .15 Thesis aims .................................................................................................................. .18 Neuroanatomy of the polychaete Nereis diversicolor ............................... 23 Introduction ................................................................................................................. .25 Material and Methods .................................................................................................. .26 Immunohistochemistry ........................................................................................ .26 Retrograde staining ............................................................................................. .27 Results ......................................................................................................................... .27 Immunoreactivity ................................................................................................. .29 Central optic neuropil ......................................................................................... .33 Mushroom bodies ................................................................................................ .34 Discussion .................................................................................................................... .37 Mushroom bodies ................................................................................................ .37 Central optic neuropil ......................................................................................... .39 3D reconstruction of annelid mushroom body neuropils ........................ 43 Introduction ................................................................................................................. .45 Material and Methods .................................................................................................. .46 Immunohistochemistry ........................................................................................ .46 Image acquisition and Computer reconstruction ................................................ .46 Assembling and Embedding the 3D model .......................................................... .48 Results .......................................................................................................................... .48 Nereis diversicolor ............................................................................................... .48 Harmothoe areolata ............................................................................................. .51 Lepidonotus clava ................................................................................................ .54 Discussion .................................................................................................................... .55 Annelid neuroanatomy: tracing higher brain centers in segmented worms ..................................................................................... 59 Introduction .................................................................................................................. .61 Material and Methods ................................................................................................... .62 Results .......................................................................................................................... .62 Arenicola marina ................................................................................................. .62 Ophelia limacina .................................................................................................. .64 Scalibregma inflatum ........................................................................................... .64 Eupolymnia nebulosa ........................................................................................... .64 Sabella penicillus ................................................................................................. .66 Tomopteris helgolandica ..................................................................................... .66 Nereis diversicolor ............................................................................................... .66 Phyllodoce maculata ............................................................................................ .68 Nephtys hombergii ............................................................................................... .68 Eunice torquata .................................................................................................... .68 Lumbrineris cf. fragilis ........................................................................................ .70 Odontosyllis cf. fulgurans .................................................................................... .70 Hesione pantherina .............................................................................................. .72 Harmothoe areolata ............................................................................................. .72 Lumbricus terrestris ............................................................................................. .72 Hirudo medicinalis .............................................................................................. .73 Discussion .................................................................................................................... .74 Mushroom bodies ................................................................................................
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