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Morphologies and Phylogenetic Classification of Cellulolytic Myxobacteria

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Morphologies and Phylogenetic Classification of Cellulolytic Myxobacteria http://www.paper.edu.cn System. Appl. Microbiol. 26, 104–109 (2003) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/sam Morphologies and Phylogenetic Classification of Cellulolytic Myxobacteria Zhang-Cai Yan, Bing Wang, Yue-Zhong Li, Xun Gong, Huai-Qiang Zhang, and Pei-Ji Gao State Key Laboratory of Microbial Technology, College of Life Science, Shandong University, Jinan, China Received: September 15, 2002 Summary The evolutionary distances of the 16S rDNA sequences in cellulolytic myxobacteria are less than 3%, which units all the strains into a single genus, Sorangium. The size of myxospores and the shape of spo- rangioles, rather than fruiting body colors or swarm morphologies are consistent with the changes of the 16S rDNA sequences. It is suggested that there are at least two species in the genus Sorangium: one in- cludes strains with small myxospores and spherical sporangioles, and the color of the fruiting bodies is normally orange or brown, though sometimes yellow or black. The second species has large myxospores, polyhedral sporangioles with many inter-cystic substrates, and normally deep brown to black color. Key words: cellulolytic – myxobacteria – Sorangium – morphologies – 16S rDNA – Classification Introduction Myxobacteria are unique among bacteria for their lulosum, but suggested that there were at least two complicated multicellular morphogenesis, which also species in the genus considering the differences of their forms the primary basis for myxobacterial taxonomy fruiting body colors. [7, 10]. Phylogenetically, the Myxobacterales are located In 1992, Shimkets and Woese [12] phylogenetically in the δ-division of Proteobacteria depending on the analyzed 12 representative strains from 10 myxobacteri- analysis of their 16S rDNA sequences [8, 14], but classi- al genera, including one cellulose-degrader. After com- fied into 12 genera and about 40 species by their mor- prehensive comparison of the 16S ribosomal RNA se- phological characteristics [7, 10]. Based on their special- quences, they confirmed that myxobacteria form three izations in degradation of biomacromolecules, myxobac- major sublines, matching the prior conclusion from anal- teria, except the unculturable genus Haploangium, are ysis of the RNase T1-resistant fragments of five divided into two groups. One is bacteriolytic, efficiently myxobacterial 16S ribosomal RNAs [5]. Of the three, lysing whole living cells of other microorganisms, and Sorangium (Polyangium cellulosum), together with the other is cellulolytic, decomposing cellulose instead of Chondromyces and Polyangium forms the Chon- living cells [9]. Out of all the classified myxobacterial dromyces subline. Later, Spröer et al. [13] compared species, only one is included in the latter. In the 8th edi- many more myxobacterial strains by 16S rDNA se- tion of Bergey’s Manual of Systematic Bacteriology, cel- quences and discussed the correlation between morpho- lulolytic myxobacterial strains were united with genus logical and phylogenetic classification of myxobacteria. Polyangium due to their similar morphological charac- However, no cellulolytic myxobacteria were included in ters and named Polyangium cellulosum [6]. In the 9th the 54 strains they assayed. In the present paper, ten cel- edition of the manual, they were relocated in an individ- lulolytic myxobacterial strains, selected by their nuances ual but deliberately undetermined genus Sorangium and in morphological characters from our myxobacterial li- renamed S. cellulosum [7]. In the 2nd edition of The brary [2, 3] are assayed of their 16S rDNA sequences Prokaryotes (1992), Reichenbach and Dworkin [10] also and the correlation between morphologies and phyloge- classified the cellulolytic myxobacteria as Sorangium cel- netic classification is discussed. 转载 中国科技论文在线 http://www.paper.edu.cn Morphologies and Phylogenetic Classification of Cellulolytic Myxobacteria 105 Fig. 1. The morphologies of the cellulolytic myxobacteria. Panels A to I are the fruiting bodies of So9882-1, So9735-22, So9857, So0089-1, So9721-1, So9733-1, So9881, So ce26 and So9741 respectively; J to L the swarm edges of So9733-1, So9881 and So9741; and M to O the cells moving, disposing and degrading the cellulose fibers. The bar in B is 100 µm for A–L and in N 10 µm for M–Q. Pictures in panels A to L were obtained by phase contrast microscopy and pictures in panels M to O by scanning electron mi- croscopy. 中国科技论文在线 http://www.paper.edu.cn 106 Z.-C. Yan et al. Materials and Methods Bacterial strains and culture conditions The ten cellulolytic myxobacterial strains reported in this paper are listed in Table 1. Among them, strain So ce26 was kindly furnished by H. Reichenbach (Gesellschaft für Biotech- nologische Forschung mbH, Braunschweig, Germany), and oth- ers were isolated and kept by the authors. Strains were routinely cultured on CNST medium containing (w/v) KNO3 0.05%; Na2HPO4·12H2O 0.025%; MgSO4 ·7H2O 0.1%; FeCl3 ·6H2O 0.001%; agar 1.5%; and trace elements solution [10] 1 ml l–1. The pH of the medium was adjusted to 7.0–7.2 before autoclav- ing. To provide the carbon source, small pieces of sterilized filter paper were placed on the medium surface. The bacteria were in- oculated on the paper and incubated at 30 °C. Morphologies The size of vegetative cells was the average of 200 cells in 3-day cultures, measured under a phase-contrast microscope. Fruiting bodies were scraped form plates and transferred to glass slide for observation. The mature myxospores were ex- truded from the aged fruiting bodies. Scanning electron mi- croscopy was employed to detect the cellular behaviors on cellu- lose fibers after the samples were fixed in 2.5% glutaraldehyde, dehydrated in a graded ethanol series to 100% ethanol, dried at the critical point and coated with gold. 16S rDNA sequencing The vegetative cells in 3-day cultures were harvested, com- pletely scattered with glass beads (3 mm in diameter), cen- trifuged, washed with distilled water twice and suspended in STE solution containing 0.1 mol l–1 NaCl, 10 mmol l–1 Tris-Cl (pH8.0) and 1 mmol l–1 EDTA. The genomic DNA was extract- ed from the cells, and used for PCR amplification of 16S rDNA segments [4]. The length and the access numbers of 16S rDNA segments in the GenBank nucleotide sequence database are pre- sent in Table 1. Phylogenetic analysis 16S rDNA sequences of the cellulolytic myxobacterial strains Fig. 2. The phylogenetic tree based on the comparison of 16S were compared with the known myxobacteria, especially rDNA sequences showing the correlation of the cellulolytic Polyangium spp. and Chondromyces spp. reported by Shimkets strains with other myxobacteria. The bar indicated the evolu- and Woese [12] or Spröer et al. [13]. The referenced myxobacte- tionary distance. The numbers on branch nodes were percent- rial strains and their GenBank access numbers are Polyangium ages of bootstrap supports from 1000 samples. cellulosum, M94282; Angiococcus disciformis An d4, AJ233910; Archangium gephyra Ar g1, AJ233912; Corallococ- cus exiguus Cc e100, AJ233932; Cystobacter fuscus Cb f10, AJ233898; Melittangium boletus Me b8, AJ233908; Myxococ- cus fulvus Mx f2, AJ233917; Nannocystis exedens Na e1, Results AJ233946; Stigmatella aurantiaca Sg a15, AJ233936; Polyangium sp., M94280; P. vitellinum Pl vt1, AJ233944; Morphologies of cellulolytic P. thaxteri Pl t3, AJ233943; Chondromyces crocatus, M94275; myxobacteria C. robustus Cm a22, AJ233941; C. robustus Cm a13, AJ233942; C. pediculatus Cm p17, AJ233940; C. lanuginosus Table 1 presents the major morphological characters Sy t2, AJ233939; C. apiculatus, M94274; and C. apiculatus Cm of the ten cellulolytic myxobacterial strains. All the a14, AJ233938. The evolutionary distance matrix of 16S rDNA strains were nearly the same in the shape of vegetative sequences was obtained by Neighbor-joining method [11] after cells or myxospores, i.e. cylindrical, rigid rods, and with they were completely aligned using ClustalX (1.81) [15]. The bluntly round ends. The sizes of myxospores could be topologies of the phylogenetic tree were constructed using the divided into two groups: 3–4 × 1–1.2 µm and Dnadist/Neighbor programs in the PHYLIP package (v3.6) [1]. 2–3 × 1 µm. Normally, myxospores were embedded in Likelihood and parsimony programs in the same package were sporangioles, however, strain So9987-5 had no any spo- also employed to testify the distance tree. The bootstrapping supports for the tree were calculated from 1000 samples. rangiole structure at all, even when it was initially detect- Bdellovibrio starrii (access number AF084852) and Sulfate-re- ed on a soil sample. The strain was isolated and purified ducing bacterium Desulfotomaculum reducens (access number by its typical myxobacterial film-like swarm. Another ex- U95951) were used to root the tree. ception was strain So9881-1, which quickly lost the ca- 中国科技论文在线 http://www.paper.edu.cn Table 1. The cellulolytic myxobacterial strains, access numbers of 16S rDNA sequences, and the major morphological characteristics Strains 16S rDNA Length of Vegetative Myxospores Fruiting bodies Swarms Access No. 16S rDNA cells (µm) (µm) ––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––––
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