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Phylogenetische Analyse Der 18S Rrna Identifiziert Den ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2004 Band/Volume: 0013 Autor(en)/Author(s): Wylezich Claudia, Radek Renate, Schlegel Martin Artikel/Article: Phylogenetische Analyse der 18S rRNA identifiziert den parasitschen Protisten Nephridiophaga blattellae (Nephridiophagidae) als Vertreter der Zygomycota (Fungi) 435-442 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Denisia 13 | 17.09.2004 | 435-442 Phylogenetische Analyse der 18S rRNA identifiziert den parasitischen Protisten Nephridiophaga blattet I ae (Nephridiophagidae) als Vertreter der Zygomycota (Fungi)1 C. WYLEZICH, R. RADEK £t M. SCHLEGEL Abstract: Phylogenetic analysis of 18S rRNA identifies the parasitic protist Nephridiophaga blattellae (Nephrid- iophagidae) as a member of fungi. — The taxonomic affiliation of the spore-forming, unicellular nephridiophagids that thrive in the Malpighian tubules of insects is still completely unresolved. Their morphological and ultrastruc- tural characters do not closely resemble any known taxon of spore-forming protists. Some authors classed them with the Haplosporidia but proof is poor; e.g. the typical haplosporosomes are missing in nephridiophagids. In the present examination, the 18S rRNA of Nephridiophaga blatiellae (from the German cockroach Blauella germanica) was ampli- fied and sequenced in order to reconstruct the phylogenetic relationships of that group. Results exclude a close rela- tion to haplosporidia. Data rather support their classification within the fungi. Morphological findings such as the presence of chitin also argue in favour of this hypothesis. According to our analysis the exact position of nephridio- phagids seems to be near to the Zygomycota, as it is also reported for microsporidia. Key words: Nephridiophagidae, phylogeny, Zygomycota, Microsporidia, 18S rRNA. Einleitung STORCH 2000, die kürzlich erstmals aus der Totenkopf- schabe Blaberus craniifer BURMEISTER 1838 (Blabera) be- Die Nephridiophagidae SPRAGUE 1970 sind mikro- schrieben wurde. Nach RADEK & HERTH (1999) gehören skopisch kleine, einzellige, Sporen formende, heterotro- wie bereits von LANGE (1993) vorgeschlagen drei weite- phe Organismen, die in den Malpighischen Gefäßen von re Gattungen zu den Nephridiophagidae: Coleospora Insekten leben (z. B. RADEK & HERTH 1999), aber relativ GlBBS 1959, Oryctospora PURRINI & WEISER 1990 und harmlos sind. Im Entwicklungszyklus finden sich ver- Peltomyces LEGER 1909, welche ausschließlich in Käfern schiedene Lebensstadien. Die vielkernigen, vegetativen (Coleoptera) vorkommen. Dieses Taxon scheint eine Plasmodien kommen extrazellulär - oder bei einigen Ar- monophyletische Gruppe zu sein. Über ihre nächsten ten auch intrazellulär - im Lumen der Malpighischen Verwandten herrscht hingegen vollkommene Unklarheit Gefäße vor. Sporogonie findet ebenfalls im Wesentlichen (z. B. LANGE 1993; RADEK & HERTH 1999). Einige Auto- dort statt, wobei durch eine innere Abgrenzung in den ren (z. B. KOWALJOWA & ISSI 1973) beschrieben Mitglie- Sporen bildenden Plasmodien etwa 10-35 flache, ovale der der Nephridiophagidae lichtmikroskopisch als Mi- Sporen entstehen. Junge Sporoblasten mit zwei oder vier crosporidia. Ultrastrukturelle Untersuchungen zeigten Kernen kommen vor; reife Sporoblasten sind dagegen jedoch, dass den Nephridiophagidae ein Polfaden sowie fast immer einkernig (siehe Abbildungen 1-3). die charakteristische, posterior lokalisierte Vakuole der Aus Schaben sind bislang vier relativ wirtsspezifische Microsporidia fehlen. Andere Autoren (z. B. SPRAGUE Arten der Gattung Nephridiophaga bekannt: N. blattetiae 1970) stellten sie hingegen - teilweise ohne Angabe von CRAWLEY 1905 {Coebsporidium) aus Blattella germanica Gründen - zu den Haplosporidia (Alveolata). Die Ne- LINNAEUS 1767 (Blatta) (CRAWLEY 1905; WOOLEVER phridiophagidae weisen allerdings keine Haplosporoso- 1966; RADEK & HERTH 1999), N. periplanetae LUTZ & men auf; die Sporenwandbildung unterscheidet sich SPLENDORE 1903 (Plistophora) aus Blatta orientalis LlN- ebenfalls deutlich bei beiden Gruppen. Vielmehr zeigte NAEUS 1758, Periplaneta americana LlNNAEUS 1758 (Blat- die Einbeziehung der Elektronenmikroskopie deutlich, ta) und P. brunnea BURMEISTER 1838 (LUTZ & SPLENDO- RE 1903; LANGE 1993), N. (Coelosporidiwn) tangae PUR- ' Wir widmen diese Studie unserem hochverehrten Kollegen Herrn Univ.-Prof. Dr. Horst Aspöck anlässlich seines 65. Geburtstages in An- RINI, WEISER & KOHRING 1988 aus Blatta sp. (PURRINI et erkennung seiner hohen, überaus umfangreichen und breit gefächerten al. 1988; LANGE 1993) sowie N. blaberi (FABEL, RADEK & wissenschaftlichen Verdienste. 435 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Abb. 1-3: Stadien im Lebenszyklus von Nephridiophaga blattellae. (1) Reife Sporen üherschichtet. Der Reaktionsansatz wurde 5 min bei 95 im Sporenplasmodium (Interferenzkontrast, Maßstab lOum); (2) Sporoblast °C vorgeheizt und weiterhin für 35 Zyklen (92 °C 60 s, innerhalb eines Sporenplasmodiums (pi; n: Nucleus, mi: Mitochondrien, er: Endoplasmatisches Reticulum, sw: Sporenhülle; TEM, Maßstab 1um); (3) 40 °C 90 s, 72 °C 120 s) sowie abschließend eine Stunde Malpighischer Tubulus (Mt) mit mehreren Parasitenstadien: einkernige Merozoiten bei 72 °C inkubiert. Das PCR Produkt wurde gereinigt (me), vegetative Plasmodien (vpl), Sporenplasmodien (spl) mit Sporen (s; mi: (Microcon-30, AM1CON, Witten, Deutschland) und in Mitochondrien, n: Nuclei, mv: Microvilli; TEM, Maßstab lOum). den pGEM-T-Vektor (PROMEGA, Mannheim, dass die Sporen der Nephridiophagidae keine typischen Deutschland) ligiert. Mit den erhaltenen Plasmiden wur- Charakteristika anderer Sporen bildenden Taxa aufwei- den kompetente E. coli (JM 109) transformiert. Rekom- sen (z. B. LANUE 1993; RAOEK & HERTH 1999; siehe hier- binante Plasmide wurden mit einer Thermo Sequenase zu auch Tabelle 1). (AMERSHAM, Braunschweig, Deutschland) sequen- ziert und die DNA-Fragmente in einem automatischen Um nun die phylogenetischen Beziehungen der Ne- Sequenzierer (Ll-COR, MWG-Biotech, Ebersberg, phridiophagidae zu anderen Einzellern :u untersuchen, Deutschland) aufgetrennt. Beide Stränge wurden kom- wurde das 18S rRNA-Gen von Nephridiophaga blattellae plett in überlappender Weise mit markierten Ml3- und sequenziert und phylogenetisch analysiert. internen Primern (WYLEZICH et al. 2002) sequenziert. Material und Methoden Die mit den verschiedenen Primern erhaltenen Se- quenzfragmente wurden mit dem Programm DNASIS, Sporen von Nephridiophaga blattellae wurden aus Kul- Version 7.0 (HITACHI Software / Pharmacia) ver- turen der Deutschen Schabe Blattella germanica gewon- knüpft. Die ermittelte Sequenz wurde mit Hilfe von nen (Herkunft: Umweltbundesamt Berlin). Um die Spo- ClustalX, Version 1.81 (THOMPSON et al. 1999) zu Se- ren zu erhalten, wurde der Darm aus den dekapitierten quenzen verschiedener Eukaryoten-Taxa einschließlich Schaben gezogen und die Malpighischen Gefäße auf Ob- der Fungi aligniert. Die Datensätze wurden auf konser- jektträgern in destilliertem Wasser zerzupft. Sporen wur- vierte, gut homologisierbare Bereiche reduziert (1274 den unter mikroskopischer Kontrolle (inverses Zeiss Positionen im Eukaryoten-Aligment, 1600 Positionen Mikroskop) mittels einer feinen Glaspipette gesammelt im Fungi-Alignment). Phylogenetische Stammbäume und in 80 % Ethanol bis zur weiteren Bearbeitung aufbe- wurden unter Anwendung der Maximum-Likelihood- wahrt. Ebenso wurden Sporen von Nephridiophaga blähen Methode (Tree-Puzzle, Version 5.0, STRIMMER & VON aus der Totenkopfschabe Blaberus aaniifer isoliert. HAESELER 1996) nach dem Modell HKY85 (HASEGAWA Der Alkohol wurde entfernt und die genomische et al. 1985) rekonstruiert. Weiterhin wurden die Distanz- DNA nach dem Protokoll von KAVENOFF & ZiMM (1973, Matrix- und Minimum-Evolution-Methode (Mega, Ver- modifiziert nach STEINBRUCH & SCHLEGEL 1983) isoliert. sion 2.1 KUMAR et al. 2001) mit jeweils 1000 Bootstrap- Das 18S rRNA-Gen wurde mittels PCR amplifiziert. Da- Replikationen angewandt (Kimura-2-Parameter-Modell, zu wurden 1-20 ng DNA mit je 0,1 uM der Eukaryoten- KlMURA 1980). Dabei wurden verschiedene Taxa als spezifischen Primer (MEHLIN et al. 1988), 200 uM dNTPs Außengruppenvertreter verwendet. Zusätzlich wurden sowie Ix PCR-Puffer und 1 unit Taq-DNA Polymerase verschiedene alternative Stammbaumtopologien anhand (SIGMA, München, Deutschland) gemischt. Das End- des im Programm Puzzle enthaltenen Kishino-Hasegawa- volumen von 50 ul wurde mit einem Volumen Mineralöl Tests (KlSHlNO & HASEGAWA 1989) untersucht. 436 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Taxon Den Nephidiophagidae Wirtsspektrum Chitin Referenzen für Zuordnung der fehlende Merkmale Nephridiophagidae zu entsprechendem Taxon Apicomplexa Apikaikomplex, Pellicula Invertebraten (Arthropoda), Keine Vertebrata Haplosporidia Haplosporosomen, Sporenwand- Invertebraten (Mollusken) PURRINI & WEISER 1990 bildung durch umwachsendes SPRAGUE 1970 Episporen-Cytoplasma WOOLEVER 1966 Microsporidia Polfaden, Polaroplast, posteriore Invertebraten, Vertebrata + KOWAUOWA & Issi 1973 Vakuole PERRIN 1905 Myxozoa Polkapseln, mehrzellige Sporen Vertebrata / Anneliden + Keine Zygomycota (Fungi) Hyphen, Haustorien, asexuelle (teilweise) Invertebraten, + Keine (Sporangiosporen) und sexuelle Fungi 1 Sporen (Zygosporen) i Tab 1: Charakterisierung der Sporen formenden Taxa (Sporen meist unbeweglich, Ergebnisse parasitisch lebend), zu denen die
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