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Metagenomic Analysis of Bacterial Communities on Dokdo Island

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Metagenomic Analysis of Bacterial Communities on Dokdo Island J. Gen. Appl. Microbiol., 60, 65‒74 (2014) doi 10.2323/jgam.60.65 ©2014 Applied Microbiology, Molecular and Cellular Biosciences Research Foundation Full Paper Metagenomic analysis of bacterial communities on Dokdo Island (Received September 11, 2013; Accepted December 27, 2013) Ye-Eun Kim,1,† Hyeokjun Yoon,1,† Miae Kim,1 Yoon-Jong Nam,1 Hyun Kim,1 Yeonggyo Seo,1 Gyeong-Min Lee,1 Young Ja Kim,2 Won-Sik Kong,3 Jong-Guk Kim,1,＊ and Young-Bae Seu1,＊ 1 Department of Life Sciences and Biotechnology, Kyungpook National University, South Korea 2 Korea Environmental Industry & Technology Institute, South Korea 3 National Institute of Horticultural & Herbal Science, RDA, South Korea Dokdo, located east of the mainland of South Ko- dition, the percentage of the genus Mycobacterium rea, is a volcanic island designated as a natural mon- (of the phylum Actinobacteria) was nearly three ument of South Korea due to its ecological value. times higher in Seodo than Dongdo, and the propor- Dokdo is divided into Dongdo and Seodo, islands tion of the genus Gaiella was about 3.7 times higher with geological differences. The soil bacterial com- in Dongdo than Seodo. Overall, through the metage- munities on Dokdo (Dongdo and Seodo) were ana- nomic analysis, the number of species identified in lyzed using the pyrosequencing method. There were Dongdo and Seodo was 1,239 and 1,055, respective- 1,693 and 1,408 operational taxonomic units (OTU) ly. This information on the numerous culturable from Dongdo and Seodo, respectively. The statistical and unculturable bacteria is expected to help in the analyses (rarefaction curves as well as Chao1, Shan- screening of new species in Dokdo. non, and Simpson indices) showed that bacterial di- versity was slightly higher in Dongdo than Seodo. Key words: bacterial community; Dokdo Island; From results of a BLASTN search against the Ez- metagenomics Taxon-e database, the validated reads (obtained af- ter sequence preprocessing) were almost all classi- fied at the phylum level. From the phylum level Introduction down to the species level, the number of classified reads considerably decreased due to the absence of Dokdo, located 217 km (37°14′12″ N, 131°52′07″ E) direct- information concerning unculturable or unidenti- ly east of the mainland regions of Korea, is a volcanic island fied bacteria to date. Among the 36 phyla identified, formed by lava eruptions (Kim et al., 2013; Sohn, 1995). three phyla (Proteobacteria, Actinobacteria and Ac- This island has been preserved as a natural monument (since idobacteria) accounted for around 74.64%. The May 9, 2000) due to its unique ecological significance taxonomic composition was similar at the higher (http://jikimi.cha.go.kr/english). Because of frequent turbu- ranks (family and above) between Dongdo and Seo- lent flow, there are typical oceanic climatic features in this do, but a little different at the genus level. There region, with an average temperature of around 12°C and were also various differences in the relative abun- 85% air moisture throughout the year due to continuous rain dance of taxonomic ranks between Dongdo and Se- or snow (Chang et al., 2002). Dokdo consists of two main odo. In particular, the proportion of the genus Ac- islets, Dongdo and Seodo, but the whole island is one mass idobacterium (of the phylum Acidobacteria) was under the ocean. Dongdo is made up of andesitic volcanic about six times higher in Seodo than Dongdo. In ad- rocks, while Seodo is composed of andesite and basalt rocks. Geographically, most of the land is covered under a 30 cm ＊Corresponding author footnote: Drs. Kim and Seu are co-corresponding authors. Jong-Guk Kim, Microbial-Genomics Laboratory, 427, Biology Building, Dept. of Life Sciences and Biotechnology, Kyungpook National Univer- sity, 702‒701, Daegu, South Korea. Tel: +82‒53‒950‒5379 Fax: +82‒53‒955‒5379 E-mail: [email protected] Young-Bae Seu, Bio-Organic Chemistry Laboratory, 422, Biology Building, Dept. of Life Sciences and Biotechnology, Kyungpook National University, 702‒701, Daegu, South Korea. Tel: +82‒53‒950‒5380 Fax: +82‒53‒955‒5522 E-mail: [email protected] †Both authors contributed equally to this work. None of the authors of this manuscript has any financial or personal relationship with other people or organizations that could inappropriately influence their work. 66 KIM et al. soil layer, and Dongdo and Seodo have a 60° steep slope ing massive sequence data (Handelsman, 2004; Shokralla et (Sohn, 1995). Due to their poor soil layer, rainwater is al., 2012). Among the NGS technologies, Roche 454 GS- immediately drained; consequently, these islets lack water. FLX pyrosequencing is widely used in metagenomics (Gilles Thus, only a few (approximately 50‒60) plant species exist et al., 2011; Li et al., 2013). In this study, metagenomic on this island. In contrast, approximately 4,596 kinds of analysis was performed using the GS-FLX pyrosequencing plant species exist on the mainland regions (Kim et al., 2007; platform. Bacterial communities from soil samples of Park et al., 2010; Shin et al., 2004). Dongdo and Seodo regions were analyzed and comparative The flora and fauna of Dokdo have been consistently analyses (bacterial composition and their relative abundance) studied, while the microflora has only been very recently were performed between Dongdo and Seodo samples. researched. Between 2005 and 2013, 36 species of 4 phyla (Firmicutes, Proteobacteria, Actinobacteria, and Bacteroide- Materials and Methods tes) were studied (National Center for Biotechnology Information (NCBI)). The phylum Firmicutes, a group of Collection of soil samples. The sampling sites were in low-GC-content gram-positive bacteria, produces endospores. Dongdo (37°14′26.8″ N, 131°52′10.4″ E) and Seodo In this phylum, the new species Virgibacillus dokdonensis (37°14′30.6″ N, 131°51′54.6″ E) islets of Dokdo, Uljin-gun, was identified for the first time, in 2005 (Yoon et al., 2005b). Gyeongsangbuk-do, South Korea. The sample soils were The phylum Proteobacteria, a group of gram-negative collected by using cores (depth: 15 cm, diameter: 2 cm) at bacteria, includes various nitrogen fixers and some these sites and were homogenized separately for experi- pathogens. This phylum is divided into the five classes: ments. Alphaproteobacteria, Betaproteobacteria, Deltaproteobacte- DNA extraction and pyrosequencing. For pyrosequenc- ria, Epsilonproteobacteria, and Gammaproteobacteria. The ing analysis, metagenomic DNA was extracted from soil class Alphaproteobacteria includes various symbiotic samples with the FastDNA SPIN Kit for Soil (MP Biomedi- bacteria of plants and animals. The new species Porphyro­ cals, Santa Ana, CA). The resulting DNA was amplified bacter dokdonensis and Sphingomonas dokdonensis of the using barcoded fusion primers targeting the V1/V3 regions class Alphaproteobacteria were identified (Yoon et al., of 16S rRNA genes (Wang and Qian, 2009). The fusion 2006c, e). The new species Variovorax dokdonensis of primers contained 454 specific adapters, a key sequence Betaproteobacteria (which includes nitrogen fixers and (4 bp), a barcode (7‒11 bp unique sequence), a linker pathogens) has also been newly identified (Yoon et al., sequence (2 bp), and universal primers (518R and 27F). The 2006b). For the class Epsilonproteobacteria (the species of forward primer sequence was 5′-CCTATCCCCTGTGT this class are known to inhabit the digestive tract of animals GCCTTGGCAGTC-TCAG-AC-GAGTTTGATCMTG and serve as symbionts or pathogens), the species Arcobacter GCTCAG-3′ and the reverse primer sequence was 5′-CCA marinus has been reported (Kim et al., 2010). The new TCTCATCCCTGCGTGTCTCCGAC-TCAG-X-AC- species Lysobacter dokdonensis, Shewanella dokdonensis, WTTACCGCGGCTGCTGG-3′; the “X”, representing the and Stenotrophomonas dokdonensis of the class Gammapro- barcode sequence, was used to separate samples. The PCR teobacteria (which includes various pathogens) were identi- reaction was carried out using the PTC-200 Peltier thermal fied (Oh et al., 2011; Sung et al., 2012; Yoon et al., 2006d). cycler (MJ Research, Waltham, MA). All PCR products The phylum Actinobacteria, a group of high-GC-content were purified with the QIAquick PCR Purification Kit gram-positive bacteria, produces external spores. In this (Qiagen, Valencia, CA) and the DNA products were quanti- phylum, a total of nine species of six genera has been report- fied using both the QuantiT TM PicoGreen® dsDNA Assay ed, and among the nine species, four (Isoptericola dokdonen­ Kit (Invitrogen, Carlsbad, CA) and the TBS-380 Mini sis, Nocardioides dokdonensis, Nocardioides hankookensis Fluorometer (Turner BioSystems, Sunnyvale, CA). The and Phycicoccus dokdonensis) are new species (Park et al., same amounts of the resulting PCR products from each 2008; Yoon et al., 2006f, 2008a, b). The phylum Bacteroide- sample were pooled and electrophoresed to select the DNA tes is a group of gram-negative and anaerobic bacteria. Three samples longer than 300 bp. The length of DNA fragments new species (Croceitalea dokdonensis, Maribacter was checked again by the Agilent 2100 Bioanalyzer (Agilent dokdonensis, and Polaribacter dokdonensis) of the class Technologies, Santa Clara, CA) (Panaro et al., 2000). The Flavobacteria have also been identified (Lee et al., 2008; pyrosequencing was performed by Chunlab, Inc. (Seoul, Yoon et al., 2005a, 2006a). South Korea) using the 454 GS FLX Titanium Sequencing To investigate these microflora, several methods, such as System (Roche, Branford, CT). All pyrosequencing data the culture-based assay, denaturing gradient gel electropho- were submitted to the EMBL Sequence Read Archive (SRA) resis (DGGE) and next generation sequencing (NGS)-based database under the study accession number PRJEB4517 metagenomics, have been applied (Handelsman, 2004; Hong (http://www.ebi.ac.uk/ena/data/view/PRJEB4517). et al., 2011; Islam and Sar, 2011; Shokralla et al., 2012; Taxonomical identification and statistical data analysis. Streit and Schmitz, 2004; Zhalnina et al., 2012). The culture- All preprocessed reads were identified by performing a based assay has limitation in terms of the study of uncultur- BLASTN search against the EzTaxon-e database (Altschul able microbes from environmental samples, such as soil.
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