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Stuhrmann, T. Studying New Eukaryotic Niches of Bacterial Life

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Stuhrmann, T. Studying New Eukaryotic Niches of Bacterial Life Studying new eukaryotic niches of bacterial life Marine Biological Laboratory Microbial Diversity Course 2005 Torben Stührmann Max Planck Institute for Marine Microbiology Department of Molecular Ecology Celsisus Str. 1 28379 Bremen-Germany Abstract Biologists are becoming increasingly aware that a variety of animals and plants bears symbiotic microorganisms. The formation of alliances between animals and microorganisms contribute to their nutrition or defence against predators and parasites. This study has the aim to search for new eukaryotic niches of bacterial live (symbiosis or commensalisms). As study object two different eukaryotic species from varying environments were taken. For the first study a brown insect larva was obtained from the Sippewissett Salt marshes which was not further characterized. The second study object was the jellyfish Cyanea capillata (the Lion’s Mane). This species was highly abundant in the Cape Cod area during the course period (June-August 2005). Symbiotic interactions between the cnidarians and Cyanobacteria are known but there is nothing known about bacterial interactions among the Scyphozoa. Microscopic- (Scanning electron microscopy, Light microscopy), cultivation- and molecular- techniques (16S rRNA clone libraries, Fluorescence in situ hybridization FISH) were applied for studying bacterial-eukaryotic interactions. Only few indications were found with FISH suggesting a eukaryotic niche for bacteria within the Scyphozoa. A rod shaped morphotype detected with a FISH-probe specific for Gammaproteobacteria was detected in the mouth tissue. Other approaches were not applied or failed. The 16S rRNA library of brown larvae revealed a high predominance of Gammaproteobacteria (Thiomicrospira, Halomonas, Marinobacter and Vibrio spp.). Epsilon-, alpha- proteobacteria, Bacteriodetes and Cyanobacteria were only less frequent. These results were confirmed by FISH. A coccoid highly motile morphotype was enriched from the gut tissue with artificial seawater media, yielding an almost pure isolate. This is the first known report surveying bacterial interactions in C. capillata. To reveal the secret of bacterial interactions with jellyfish further attention to this topic is needed (e.g. 16S rRNA libraries, cultivation). The results from the brown larvae taken at Sippewissett marshes indicate a niche of bacterial live. For verification of a new eukaryotic niche for bacteria a more detailed comparison of environmental and intestinal clone libraries is needed. But the results are promising to get further insights into bacterial/host interactions. Introduction Inspired by J. Leadbetters research on bacterial interactions in the termite gut (e.g. Leadbetter, J.R. & J.A. Breznak, 1996), this study has the aim to search for new eukaryotic niches of bacterial live (symbiosis or commensalisms). As study object two different eukaryotic species from varying environments were taken. The first study object was found in the Sippewissett Saltmarshes. It is a brown insect larva which was not further characterized during this study (see figure 1). After a few days in the laboratory the Sippewissett marshes become highly sulfidic and the brown larvae were moving out of the sediment. These brown larvae were studied with molecular methods for bacterial associations. The second species was the Lion’s Mane Jellyfish (Cyanea capillata). This species was highly abundant in the Cape Cod area during the course period (June-August 2005). Cnidaria are among the earliest evolved metazoan animal phyla with representatives found in fossils from the Precambrian 550 million years ago (Chen et al., 2002; Wood et al., 2002). Since Scyphozoa often form high biomass in the oceans, the degradation of dead individuals contaminated with antimicrobial and neurotoxins is of special interest. Nothing is known about microorganisms degrading those jellyfish. Using molecular 16S rRNA based methods, microorganisms involved in this process were identified. The phylum Cnidaria comprises benthic and pelagic aquatic animals including the classes Anthozoa (hard corals, soft corals, sea pens, and sea anemones), Hydrozoa (hydroids, fire corals), Scyphozoa (jellyfish) and Cubozoa (box jellyfish). The body plan of Cnidaria is diploblastic, i.e. the body consists of two cell layers, the ectoderm (epidermis) covering the outer surface of the body and the endoderm (gastrodermis) lining the body cavity. There is homogeneous elastic material (mesoglea) between these layers. This gelatinous material is a distinct feature of the bell of jellyfish. Nematocysts are stinging capsules characteristic of Cnidaria. Nematocysts contain and fire harpoon-like microscopic structures (cnida) that penetrate the surface layer of the victim and deliver a mixture of highly toxic substances. Cnidarians are known for their antimicrobial activity (Bhosale et al. 2002). However, there are also known symbiotic interactions between the cnidarians (i.e. the Caribbean coral M. cavernosa) and cyanobacteria (Lesser et al., 2004). This study addresses the question of whether these interactions are limited to corals or if interactions are also detectable in Scyphozoa. For this different types of tissues from Jellyfish were studied. Both topics are not very well studied; in fact there is no publication dealing with bacterial interactions with Scyphozoa. This study wants to contribute new insights into the symbiotic interactions between Scyphozoa and bacteria. Further time was spent on Scanning electron microscopy (SEM) of different cultures enriched during the course. Materials and Methods Sample collection Cyanea capillata (Lion’s mane) species were caught in waters of the Cape Cod area. The C. capillata species were maintained at room temperature with a continuous flow of seawater. The brown larvae were obtained from Sippewisset Marshes Mats and were not further characterized. Samples for Scanning Electron microscopy were enriched during the course from Sippewisset marshes (GSB-, H2/CO2-Enrichment, and Azotobacter-media) enrichments mats as described in the course manual (Microbial Diversity course 2005). Sample preparation Samples from C. capillata were washed three times in filtered seawater (0.2 µm), for relaxation of species a 0.15 M MgCl2 was used. Alternatively a 4% agarose solution was used to preserve the structure of the Jellyfish, but is not recommended for sectioning. Two different samples of jellyfish were analyzed. One species died during maintenance and was fixed in agarose after several hours. The formaldehyde fixation of these samples was directly carried out on the filter sections. The second species was killed during the fixation process. The brown larvae were washed 3 times in 1x PBS. They were sectioned and opened, the white internal substance was diluted in 1x PBS. The brown capsule was discarded. The sample was fixed with 2% Formaldehyde for 30 min and filtrated through a 0.2 µm Millipore GTTP Filter. Otherwise the sample was diluted in 1 ml 1x PBS for the enrichment of intestinal bacteria and spread on agar plates in different dilutions (see Media and Growth conditions). FISH Fluorescence in-situ-hybridization (FISH) was performed as described in the course manual (Microbial Diversity course 2005). For investigation of different tissues the following probes were used with 35% Formamide at 46°C: Table 1: Conditions for the FISH probes used in the experiments probe specificity sequence formamide target competitor ALF968 alpha-group of GGTAAGGTTCTGCGCGTT 35 16S Proteobacteria beta-group of BET42a GCCTTCCCACTTCGTTT 35 23S Gam42a Proteobacteria Cytophaga/Flavo- CF319a TGGTCCGTGTCTCAGTAC 35 16S bacterium cluster most Bacteria, no EUB338 GCTGCCTCCCGTAGGAGT 0-35 16S Planctomycetales Gamma-group of GAM42a GCCTTCCCACATCGTTT 35 23S Bet42a Proteobacteria nonsense probe, for Non338 detection of non ACTCCTACGGGAGGCAGC 0-35 16S specific binding Light Microscopy The samples were observed under phase-contrast and epifluorescence on a Zeiss AXIOPlan Imager.M1 compound light microscope with AxioVision v4.4 capture software. Dissected jellyfish tissue was stained in DAPI (1 µg/mL) for 10 minutes and washed with sterile MQ water prior to observation. Scanning Electron Microscopy Samples were fixed from the different enrichments (2% glutaraldehyde, 1.5% formaldehyde in 0.1M sodium cacodylate buffer, pH 7.0) for 4 hours, dehydrated in acidulated 2,2- dimethoxypropane, and kept in absolute ethanol. Specimens were critical-point dried with liquid CO2 and sputter-coated with gold. Images and micrographs were captured on a JEOL JSM-840 scanning electron microscope. Media and growth conditions E. coli used in clone libraries was grown at 37° C on Luria-Bertani (LB) with kanamycin supplemented at 50 µg/mL and 40 µg/mL X-gal as appropriate. LB was solidified with 1.8% (wt/vol) Bacto agar. Intestinal larvae bacteria were cultivated on artificial seawater (ASW) at 30° C containing 1 mM Na2SO4, 1.5 mM K2PO4, 5 mM NaNO3, 5 mM MOPS (pH 7.2), and 1 mL of HCl- dissolved trace elements solution per liter seawater base (20 g NaCl, 3 g MgCl2•6H2O, 0.15 g CaCl2•2H2O per liter distilled water) and 1-mL of the 12-vitamin solution. ASW-media was solidified with 1.8% (wt/vol) prewashed Bacto agar. HCl-dissolved trace elements solution contained 2.1 g FeSO4•7H2O, 30 mg H3BO3, 100 mg MnCl2•4H2O, 190 mg CoCl2•6H2O, 24 mg NiCl2•6H2O, CuCl2•2H2O, 144 mg ZnSO4•7H2O, 36 mg Na2MoO4•2H2O, 25 mg NaO3V and 6 mg Na2SeO3•5H2O in 987 mL distilled water. The 12-vitamin solution contained 10 mg riboflavin, and 100 mg each
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