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The Relevance of the Silica Metabolizing Enzyme Silicatein-Α to Biomineralization and the Formation of Biogenic Silica in Siliceous Sponges

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The Relevance of the Silica Metabolizing Enzyme Silicatein-Α to Biomineralization and the Formation of Biogenic Silica in Siliceous Sponges The relevance of the silica metabolizing enzyme silicatein-α to biomineralization and the formation of biogenic silica in siliceous sponges Dissertation zur Erlangung des Grades "Doktor der Naturwissenschaften" am Fachbereich Biologie der Johannes Gutenberg-Universität in Mainz vorgelegt von Ute Schloßmacher Mainz, 2012 Dekan: Erster Berichterstatter: Zweiter Berichterstatter: Tag der mündlichen Prüfung: 18.07.2012 This thesis is based on the following 4 research articles, which are presented as chapters 2 to 5. Müller W.E.G., Boreiko A., Schloßmacher U., Wang X.H., Tahir M.N., Tremel W., Brandt D., Kaandorp J.A. & Schröder H.C. (2007a) Fractal-related assembly of the axial filament in the demosponge Suberites domuncula: relevance to biomineralization and the formation of biogenic silica. Biomaterials 28: 4501- 4511 Müller W.E.G., Boreiko A., Schloßmacher U., Wang X.H., Eckert C., Kropf K., Li J. & Schröder H.C. (2008b) Identification of a silicatein(-related) protease in the giant spicules of the deep-sea hexactinellid Monorhaphis chuni. J Exp Biol 211, 300-309. Müller W.E.G., Schloßmacher U., Wang X.H., Boreiko A., Brandt D., Wolf S.E., Tremel W. & Schröder H.C. (2008g) Poly(silicate)-metabolizing silicatein in siliceous spicules and silicasomes of demosponges comprises dual enzymatic activities (silica-polymerase and silica-esterase). FEBS J 275, 362–370. Schloßmacher U., Wiens M., Schröder H.C., Wang X.H., Jochum K.P. & Müller W.E.G. (2011) Silintaphin-1: Interaction with silicatein during structure guiding biosilica formation. FEBS J 278, 1145-1155. Contents CHAPTER 1 ........................................................................................................... - 1 - 1.1 Porifera ....................................................................................................................... - 1 - 1.2 Aufbau der Porifera ................................................................................................... - 3 - 1.3 Potentielle Funktion der Nadeln: ein einzigartiges Nervensystem? ...................... - 6 - 1.3.1 Licht erzeugender Komplex: Luciferase-Luciferin: .................................................... - 6 - 1.3.2 Lichtdetektionssystem: Cryptochrom ........................................................................ - 8 - 1.4 Biosilikat: Schwammenzyme und ihre assoziierten Proteine ................................ - 9 - 1.4.1 Silicateine ................................................................................................................. - 9 - 1.4.1.1 Silikatchemie ....................................................................................................... - 14 - 1.4.1.2 Reaktionsmechanismus des Silicateins ............................................................... - 14 - 1.4.2 Biosilikatabbauendes Enzym: die Silicase .............................................................. - 16 - 1.4.3 Schwammspezifische Silicatein-assozierte Proteine: Die Silintaphine .................... - 18 - 1.4.3.1 Silintaphin-1 ......................................................................................................... - 18 - 1.4.3.2 Silintaphin-2: ........................................................................................................ - 18 - 1.5 Nadelwachstum ....................................................................................................... - 19 - 1.5.1 Axiales Nadelwachstum: ......................................................................................... - 21 - 1.5.2 Reifung und Härtung des Biosilikats ....................................................................... - 24 - 1.5.3 Zelluläres Zusammenspiel während der Nadelsynthese ......................................... - 29 - Referenzen ..................................................................................................................... - 31 - CHAPTER 2 ......................................................................................................... - 38 - FRACTAL-RELATED ASSEMBLY OF THE AXIAL FILAMENT IN THE DEMOSPONGE SUBERITES DOMUNCULA: RELEVANCE TO BIO- MINERALIZATION AND THE FORMATION OF BIOGENIC SILICA .................. - 38 - 2.1 Introduction ............................................................................................................. - 38 - 2.2 Experimental section............................................................................................... - 41 - 2.2.1 Chemicals, materials and enzymes ........................................................................ - 41 - 2.2.2 Sponges and spicules............................................................................................. - 41 - 2.2.3 Spicular extract ....................................................................................................... - 41 - 2.2.4 HF procedure ......................................................................................................... - 42 - Contents 2.2.5 Gel electrophoresis and immunoblotting ................................................................. - 42 - 2.2.6 Determination of proteinase activity ........................................................................ - 43 - 2.2.7 Zymogram gel ........................................................................................................ - 43 - 2.3 Results ..................................................................................................................... - 44 - 2.3.1 Analysis of spicule extract ...................................................................................... - 44 - 2.3.2 Proteinase activity of “HF-Extract-Urea” and “TG-Extract” ...................................... - 45 - 2.3.3 Proteolytic activity (zymogram) ............................................................................... - 46 - 2.3.4 Filament/aggregate formation from disassembled silicatein .................................... - 47 - 2.4 Discussion ............................................................................................................... - 51 - 2.5 References ............................................................................................................... - 57 - CHAPTER 3 ......................................................................................................... - 60 - IDENTIFICATION OF A SILICATEIN(-RELATED) PROTEASE IN THE GIANT SPICULES OF THE DEEP-SEA HEXACTINELLID MONORHAPHIS CHUNI .... - 60 - 3.1 Introduction ............................................................................................................. - 60 - 3.2 Experimental section............................................................................................... - 62 - 3.2.1 Spicules and spicule extracts.................................................................................. - 62 - 3.2.2 Analysis of proteolytic activity: zymogram analysis ................................................. - 62 - 3.2.3 Analysis of proteolytic activity: enzyme activity test ................................................ - 63 - 3.3 Results ..................................................................................................................... - 64 - 3.3.1 Proteolytic activity of silicatein ................................................................................ - 64 - 3.3.2 Cathepsin-like activity in giant basal spicules ......................................................... - 65 - 3.4 Discussion ............................................................................................................... - 67 - 3.5 References ............................................................................................................... - 69 - CHAPTER 4 ......................................................................................................... - 72 - POLY(SILICATE)-METABOLIZING SILICATEIN IN SILICEOUS SPICULES AND SILICASOMES OF DEMOSPONGES COMPRISES DUAL ENZYMATIC ACTIVITIES (SILICA-POLYMERASE AND SILICA-ESTERASE) ....................... - 72 - 4.1 Introduction ............................................................................................................. - 72 - Contents 4.2 Experimental section............................................................................................... - 76 - 4.2.1 Chemicals............................................................................................................... - 76 - 4.2.2 Sponges and primmorphs ....................................................................................... - 76 - 4.2.3 Silicatein-α .............................................................................................................. - 76 - 4.2.4 Electron immunogold labeling ................................................................................. - 77 - 4.2.5 Maldi analysis ......................................................................................................... - 77 - 4.2.6 Esterase activity ..................................................................................................... - 78 - 4.3 Results ..................................................................................................................... - 79 - 4.3.1 Presence of silicatein in spicules and in cell organelles, the silicasomes ................ - 79 - 4.3.2 Catalytic function of silicatein: silica-polymerase..................................................... - 81 - 4.3.3 Catalytic function of silicatein: silica-esterase activity.............................................
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