Open Access

Complete genome sequence of Cellulophaga algicola type strain (IC166T)

  • Birte Abt
  • , Megan Lu,
  • , Monica Misra,
  • , Cliff Han,
  • , Matt Nolan
  • , Susan Lucas
  • , Nancy Hammon
  • , Shweta Deshpande
  • , Jan-Fang Cheng
  • , Roxane Tapia,
  • , Lynne Goodwin,
  • , Sam Pitluck
  • , Konstantinos Liolios
  • , Ioanna Pagani
  • , Natalia Ivanova
  • , Konstantinos Mavromatis
  • , Galina Ovchinikova
  • , Amrita Pati
  • , Amy Chen
  • , Krishna Palaniappan
  • , Miriam Land,
  • , Loren Hauser,
  • , Yun-Juan Chang,
  • , Cynthia D. Jeffries,
  • , John C. Detter,
  • , Evelyne Brambilla
  • , Manfred Rohde
  • , Brian J. Tindall
  • , Markus Göker
  • , Tanja Woyke
  • , James Bristow
  • , Jonathan A. Eisen,
  • , Victor Markowitz
  • , Philip Hugenholtz,
  • , Nikos C. Kyrpides
  • , Hans-Peter Klenk
  • and Alla Lapidus
Corresponding author

DOI: 10.4056/sigs.1543845

Received: 22 February 2011

Published: 04 March 2011


Cellulophaga algicola Bowman 2000 belongs to the family Flavobacteriaceae within the phylum 'Bacteroidetes' and was isolated from Melosira collected from the Eastern Antarctic coastal zone. The species is of interest because its members produce a wide range of extracellular enzymes capable of degrading proteins and polysaccharides with temperature optima of 20-30°C. This is the first completed genome sequence of a member of the genus Cellulophaga. The 4,888,353 bp long genome with its 4,285 protein-coding and 62 RNA genes consists of one circular chromosome and is a part of the Genomic Encyclopedia of Bacteria and Archaea project.


aerobicmotile by glidingGram-negativeagarolyticchemoorganotrophiccold adapted enzymesFlavobacteriaceaeGEBA


Strain IC166T (= DSM 14237 = CIP 107446 = LMG 21425) is the type strain of C. algicola, which belongs to the family Flavobacteriaceae within the phylum 'Bacteroidetes'. The strain was isolated from the surface of the chain-forming sea-ice diatom Melosira collected from the Eastern Antarctic coastal zone, and was described by Bowman in 2000 [1]. Currently, there are six species placed in the genus Cellulophaga, namely C. algicola [1], C. baltica, C. fucicola, C. lytica [2], C. pacifica [3] and C. tyrosinoxydans [4]. C. lytica is the type species of the genus Cellulophaga [2]. The generic name of the genus derives from the Neo Latin word 'cellulosum' meaning 'cellulose' and the Greek word 'phagein' meaning 'to eat', referring to an eater of cellulose. Here we present a summary classification and a set of features for C. algicola IC166T, together with the description of the complete genomic sequencing and annotation.

Classification and features

A representative genomic 16S rRNA sequence of C. algicola was compared using NCBI BLAST under default settings (e.g., considering only the high-scoring segment pairs (HSPs) from the best 250 hits) with the most recent release of the Greengenes database [5] and the relative frequencies, weighted by BLAST scores, of taxa and keywords (reduced to their stem [6]) were determined. The five most frequent genera were Cellulophaga (39.5%), Maribacter (7.8%), Flavobacterium (5.6%), Cytophaga (5.4%) and Formosa (4.7%) (135 hits in total). Regarding the 21 hits to sequences from members of the species, the average identity within HSPs was 95.8%, whereas the average coverage by HSPs was 94.9%. Regarding the 16 hits to sequences from other members of the genus, the average identity within HSPs was 94.7%, whereas the average coverage by HSPs was 94.7%. Among all other species, the one yielding the highest score was C. baltica, which corresponded to an identity of 98.1% and a HSP coverage of 97.8%. The highest-scoring environmental sequence was GU452686 ('sediments coast oil polluted Black Sea coastal sediment clone 70SZ2'), which showed an identity of 96.5% and a HSP coverage of 98.1%. The five most frequent keywords within the labels of environmental samples which yielded hits were 'marin' (4.7%), 'water' (4.3%), 'sediment' (4.3%), 'sea' (3.5%) and 'coastal' (2.6%) (115 hits in total). Environmental samples which yielded hits of a higher score than the highest scoring species were not found.

The environmental samples database (env_nt) contains the marine metagenome clone ctg_1101667042524 (AACY022635173) isolated from Sargasso Sea near Bermuda, sharing 92% identity with IC166T [7] (as of January 2011).

Figure 1 shows the phylogenetic neighborhood of C. algicola IC166T in a 16S rRNA based tree. The sequences of the five 16S rRNA gene copies in the genome differ from each other by up to two nucleotides, and differ by up to 14 nucleotides from the previously published 16S rRNA sequence (AF001366), which contains nine ambiguous base calls.

Figure 1

Phylogenetic tree highlighting the position of C. algicola IC166T relative to the other type strains within the family Flavobacteriaceae. The tree was inferred from 1,458 aligned characters [8,9] of the 16S rRNA gene sequence under the maximum likelihood criterion [10] and rooted in accordance with the current taxonomy. The branches are scaled in terms of the expected number of substitutions per site. Numbers above branches are support values from 350 bootstrap replicates [11] if larger than 60%. Lineages with type strain genome sequencing projects registered in GOLD [12] are shown in blue, published genomes in bold.

The cells of C. algicola are generally rod-shaped with rounded or tapered ends with cell lengths and widths ranging from 1.5 to 4 and 0.4 to 0.5 µm, respectively (Figure 2 and Table 1). C. algicola is motile by gliding [1]. Colonies on marine 2216 agar have yellow-orange pigmentation and a compact center, with a spreading edge possessing lighter pigmentation. Their consistency is slimy and they are slightly sunken into the agar [1]. Flexirubin pigments are not formed. C. algicola grows between 0.5 and 10% NaCl, with the best growth in the presence of about 2% NaCl. The temperature range for growth is between -2°C and 28°C, with an optimum between 15-20°C on solid media and at about 20-25°C in liquid media [1]. The optimal pH for growth is about 7.5 [1].

Figure 2

Scanning electron micrograph of C. algicola IC166T

Table 1

Classification and general features of C. algicola IC166T according to the MIGS recommendations [13].




    Evidence code

    Current classification

   Domain Bacteria

    TAS [14]

   Phylum Bacteroidetes

    TAS [15,16]

   Class Flavobacteria

    TAS [17]

   Order 'Flavobacteriales'

    TAS [15]

   Family Flavobacteriaceae

    TAS [18-21]

   Genus Cellulophaga

    TAS [2]

   Species Cellulophaga algicola

    TAS [1]

   Type strain IC166

    TAS [1]

    Gram stain


    TAS [1]

    Cell shape


    TAS [1]


   motile by gliding

    TAS [1]



    TAS [1]

    Temperature range

   -2 °C – 28°C

    TAS [1]

    Optimum temperature


    TAS [1]


   0.5-10% NaCl

    TAS [1]


    Oxygen requirement


    TAS [1]

    Carbon source


    TAS [1]

    Energy source


    TAS [1]



   sea ice diatoms, macrophyte surfaces

    TAS [1]


    Biotic relationship







    Biosafety level


    TAS [22]


   surfaces of Antarctic algae

    TAS [1]


    Geographic location

   eastern Antarctic coastal zone

    TAS [1]


    Sample collection time





   not reported




   not reported




   not reported




   not reported


Evidence codes - IDA: Inferred from Direct Assay (first time in publication); TAS: Traceable Author Statement (i.e., a direct report exists in the literature); NAS: Non-traceable Author Statement (i.e., not directly observed for the living, isolated sample, but based on a generally accepted property for the species, or anecdotal evidence). These evidence codes are from of the Gene Ontology project [23]. If the evidence code is IDA, then the property was directly observed by one of the authors or an expert mentioned in the acknowledgements.

The organism is strictly aerobic and chemoorganotrophic [1]. C. algicola can hydrolyze agar, starch, gelatine, carboxymethylcellulose (CMC), casein, Tween 80, tributyrin and L-tyrosine, but not urate, xanthine or dextran, when grown in presence of 1% L-tyrosine a reddish-brown diffusible pigment is formed [1]. Nitrate reduction is positive, whereas denitrification, H2S production and indole production are negative [1,18]. Acid is formed oxidatively from D-galactose, D-glucose, D-fructose, sucrose, cellobiose, lactose and mannitol. Strain IC166T is sensitive to ampicillin, streptomycin and carbenicillin and shows resistance to tetracycline [3].


The fatty acid profile of seven Antarctic strains, including strain IC166T, was analyzed by Bowman in 2000 [1]. The hypothetical median representative of the Antarctic isolates was published. The predominant cellular fatty acids of these seven strains were branched-chain saturated and unsaturated fatty acids and straight-chain saturated and mono-unsaturated fatty acids, namely iso-C15:0 (7.5%), iso-C15:1ω10c (7.5%), iso -C17:1ω7c (6.1%), C15:0 (14.3%), C16:1ω7c (19.2%), iso -C15:0 3-OH (8.6%), iso-C16:0 3-OH (6.5%) and iso -C17:0 3-OH (4.5%) [1]. The isoprenoid quinones of C. algicola were not determined, but for C. pacifica the presence of MK-6 as the major lipoquinone was described [3]. Polar lipids not have been studied.

Genome sequencing and annotation

Genome project history

This organism was selected for sequencing on the basis of its phylogenetic position [24], and is part of the Genomic Encyclopedia of Bacteria and Archaea project [25]. The genome project is deposited in the Genomes OnLine Database [12] and the complete genome sequence is deposited in GenBank. Sequencing, finishing and annotation were performed by the DOE Joint Genome Institute (JGI). A summary of the project information is shown in Table 2.

Table 2

Genome sequencing project information





     Finishing quality



     Libraries used

      Three genomic libraries: one 454 pyrosequence standard library,       one 454 PE library (12 kb insert size), one Illumina library


     Sequencing platforms

      Illumina GAii, 454 GS FLX Titanium


     Sequencing coverage

      146.0 × Illumina; 53.5 × pyrosequence



      Newbler version,       Velvet version 0.7.63, phrap version SPS D 4.24


     Gene calling method

      Prodigal 1.4, GenePRIMP



     Genbank Date of Release

      January 18, 2011

     GOLD ID


     NCBI project ID


     Database: IMG-GEBA



     Source material identifier

      DSM 14237

     Project relevance

      Tree of Life, GEBA

Growth conditions and DNA isolation

C. algicola IC166T, DSM 14237, was grown in DSMZ medium 514 (BACTO marine broth) [26] at 15°C. DNA was isolated from 0.5-1 g of cell paste using MasterPure Gram-positive DNA purification kit (Epicentre MGP04100) following the standard protocol as recommended by the manufacturer with modification st/DL for cell lysis as described in Wu et al. [25]. DNA is available through the DNA Bank Network [27].

Genome sequencing and assembly

The genome was sequenced using a combination of Illumina and 454 sequencing platforms. All general aspects of library construction and sequencing can be found at the JGI website [28]. Pyrosequencing reads were assembled using the Newbler assembler version 2.3-PreRelease-09-14-2009-bin (Roche). The initial Newbler assembly consisting of 128 contigs in two scaffolds was converted into a phrap assembly by [29] making fake reads from the consensus, to collect the read pairs in the 454 paired end library. Illumina GAii sequencing data (710 Mb) was assembled with Velvet [30] and the consensus sequences were shredded into 1.5 kb overlapped fake reads and assembled together with the 454 data. The 454 draft assembly was based on 263.4Mb 454 draft data and all of the 454 paired end data. Newbler parameters are -consed -a 50 -l 350 -g -m -ml 20. The Phred/Phrap/Consed software package [29] was used for sequence assembly and quality assessment in the subsequent finishing process. After the shotgun stage, reads were assembled with parallel phrap (High Performance Software, LLC). Possible mis-assemblies were corrected with gapResolution [28], Dupfinisher [31], or sequencing cloned bridging PCR fragments with subcloning or transposon bombing (Epicentre Biotechnologies, Madison, WI). Gaps between contigs were closed by editing in Consed, by PCR and by Bubble PCR primer walks (J.-F.Chang, unpublished). A total of 1,054 additional reactions and three shatter libraries were necessary to close gaps and to raise the quality of the finished sequence. Illumina reads were also used to correct potential base errors and increase consensus quality using a software Polisher developed at JGI [32]. The error rate of the completed genome sequence is less than 1 in 100,000. Together, the combination of the Illumina and 454 sequencing platforms provided 199.5 × coverage of the genome. The final assembly contained 697,305 pyrosequence and 20,331,123 Illumina reads

Genome annotation

Genes were identified using Prodigal [33] as part of the Oak Ridge National Laboratory genome annotation pipeline, followed by a round of manual curation using the JGI GenePRIMP pipeline [34]. The predicted CDSs were translated and used to search the National Center for Biotechnology Information (NCBI) nonredundant database, UniProt, TIGR-Fam, Pfam, PRIAM, KEGG, COG, and InterPro databases. Additional gene prediction analysis and functional annotation was performed within the Integrated Microbial Genomes - Expert Review (IMG-ER) platform [35].

Genome properties

The genome consists of a 4,888,353 bp long chromosome with a GC content of 33.8% (Table 3 and Figure 3). Of the 4,347 genes predicted, 4,285 were protein-coding genes, and 62 RNAs; 122 pseudogenes were also identified. The majority of the protein-coding genes (59.5%) were assigned with a putative function while the remaining ones were annotated as hypothetical proteins. The distribution of genes into COGs functional categories is presented in Table 4.

Table 3

Genome Statistics



     % of Total

Genome size (bp)



DNA coding region (bp)



DNA G+C content (bp)



Number of replicons


Extrachromosomal elements


Total genes



RNA genes



rRNA operons


Protein-coding genes



Pseudo genes



Genes with function prediction



Genes in paralog clusters



Genes assigned to COGs



Genes assigned Pfam domains



Genes with signal peptides



Genes with transmembrane helices



CRISPR repeats


Figure 3

Graphical circular map of the chromosome. From outside to the center: Genes on forward strand (color by COG categories), Genes on reverse strand (color by COG categories), RNA genes (tRNAs green, rRNAs red, other RNAs black), GC content, GC skew.

Table 4

Number of genes associated with the general COG functional categories








      Translation, ribosomal structure and biogenesis




      RNA processing and modification








      Replication, recombination and repair




      Chromatin structure and dynamics




      Cell cycle control, cell division, chromosome partitioning




      Nuclear structure




      Defense mechanisms




      Signal transduction mechanisms




      Cell wall/membrane/envelope biogenesis




      Cell motility








      Extracellular structures




      Intracellular trafficking, secretion, and vesicular transport




      Posttranslational modification, protein turnover, chaperones




      Energy production and conversion




      Carbohydrate transport and metabolism




      Amino acid transport and metabolism




      Nucleotide transport and metabolism




      Coenzyme transport and metabolism




      Lipid transport and metabolism




      Inorganic ion transport and metabolism




      Secondary metabolites biosynthesis, transport and catabolism




      General function prediction only




      Function unknown




      Not in COGs

Insights from genome sequence

A closer look on the genome sequence of strain IC166T revealed a set of genes which might be responsible for the yellow-orange color of C. algicola cells by encoding enzymes that are involved in the synthesis of carotenoids. Carotenoids are produced by the action of geranylgeranyl pyrophosphate synthase (Celal_1770), phytoene synthase (Celal_2446), phytoene desaturase (Celal_2447), lycopene cyclase (Celal_1771) and carotene hydroxylase (Celal_2445). Geranylgeranyl pyrophosphate synthases start the biosynthesis of carotenoids by combining farnesyl pyrophosphate with C5 isoprenoid units to C20-molecules, geranylgeranyl pyrophosphate. The phytoene synthase catalyzes the condensation of two geranylgeranyl pyrophosphate molecules followed by the removal of diphosphate and a proton shift leading to the formation of phytoene. Sequential desaturation steps are conducted by the phytoene desaturase followed by cyclisation of the ends of the molecules catalyzed by the lycopene cyclase [36].

Strain IC166T produces a wide range of extracellular enzymes degrading proteins and polysaccharides. These enzymes are cold adapted, they have temperature optima between 15-30°C and can tolerate temperatures below 0°C [37]. For that reason they are of special interest for industrial and biotechnical applications. C. algicola like the other members of the genus Cellulophaga, cannot hydrolyze filter paper or cellulose in its crystalline form, though they can hydrolyze the soluble cellulose derivative carboxymethylcellulose (CMC). The genome sequence of strain IC166T revealed the presence of three cellulases (Celal_0025, Celal_2753, Celal_3912), probably responsible for the hydrolysis of CMC. In addition two β-glucosidases (Celal_0470, Celal_1802) were identified in the genome, catalyzing the break down of the glycosidic β-1,4 bond between two glucose molecules in cellobiose.

The IC166T genome contains 22 genes coding for sulfatases, which are located in close proximity to glycoside hydrolase genes suggesting that sulfated polysaccharides may be used as substrates. α-L-fucoidan could be a substrate, as five α-L-fucosidases (Celal_2459, Celal_2466, Celal_2469, Celal_2470, Celal_2473) are located in close proximity to three sulfatases (Celal_2464, Celal_2468, Celal_2472). Sakai and colleagues report the existence of intracellular α-L-fucosidases and sulfatases, which enable 'Fucophilus fucoidanolyticus' to degrade fucoidan [38]. This fucoidan degrading ability could be also shared by Coraliomargarita akajimensis, as the annotation of the genome sequence revealed the existence of 49 sulfatases and twelve α-L-fucosidases [39]. In addition, three β-agarases (Celal_2463, Celal_2494, Celal_3979) were identified, with two of them located in the above mentioned region, which is rich in genes encoding glycoside hydrolases and sulfatases.



We would like to gratefully acknowledge the help of Regine Fähnrich (DSMZ) for growing C. algicola cultures. This work was performed under the auspices of the US Department of Energy Office of Science, Biological and Environmental Research Program, and by the University of California, Lawrence Berkeley National Laboratory under contract No. DE-AC02-05CH11231, Lawrence Livermore National Laboratory under Contract No. DE-AC52-07NA27344, and Los Alamos National Laboratory under contract No. DE-AC02-06NA25396, UT-Battelle and Oak Ridge National Laboratory under contract DE-AC05-00OR22725, as well as German Research Foundation (DFG) INST 599/1-2.

This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


  1. Bowman JP. Description of Cellulophaga algicola sp. nov., isolated from the surfaces of Antarctic algae, and reclassification of Cytophaga uliginosa (ZoBell and Upham 1944) Reichenbach 1989 as Cellulophaga uliginosa comb. nov. Int J Syst Evol Microbiol. 2000; 50:1861-1868PubMed
  2. Johansen JE, Nielsen P and Sjøholm C. Description of Cellulophaga baltica gen. nov., sp. nov. and Cellulophaga fucicola gen. nov., sp. nov. and reclassification of [Cytophaga] lytica to Cellulophaga lytica gen. nov., comb. nov. Int J Syst Evol Microbiol. 1999; 49:1231-1240PubMed
  3. Nedashkovskaya OI, Suzuki M, Lysenko AM, Snauwaert C, Vancanneyt M, Swings J, Vysotskii MV and Mikhailov VV. Cellulophaga pacifica sp. nov. Int J Syst Evol Microbiol. 2004; 54:609-613 View ArticlePubMed
  4. Kahng HY, Chung BS, Lee DH, Jung JS, Park JH and Joen CO. Cellulophaga tyrosinoxydans sp. nov., a tyrosinase producing bacterium isolated from seawater. Int J Syst Evol Microbiol. 2009; 59:654-657 View ArticlePubMed
  5. DeSantis TZ, Hugenholtz P, Larsen N, Rojas M, Brodie EL, Keller K, Huber T, Dalevi D, Hu P and Andersen GL. Greengenes, a chimera-checked 16S rRNA gene database and workbench compatible with ARB. Appl Environ Microbiol. 2006; 72:5069-5072 View ArticlePubMed
  6. Porter MF. An algorithm for suffix stripping. Program: electronic library and information systems. 1980; 14:130-137 View Article
  7. Venter JC, Remington K, Heidelberg JF, Halpern AL, Rusch D, Eisen JA, Wu D, Paulsen I, Nelson KE and Nelson W. Environmental genome shotgun sequencing of the Sargasso Sea. Science. 2004; 304:66-74 View ArticlePubMed
  8. Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol Biol Evol. 2000; 17:540-552PubMed
  9. Lee C, Grasso C and Sharlow MF. Multiple sequence alignment using partial order graphs. Bioinformatics. 2002; 18:452-464 View ArticlePubMed
  10. Stamatakis A, Hoover P and Rougemont J. A rapid bootstrap algorithm for the RAxML Web servers. Syst Biol. 2008; 57:758-771 View ArticlePubMed
  11. Pattengale ND, Alipour M, Bininda-Emonds ORP, Moret BME and Stamatakis A. How many bootstrap replicates are necessary? Lect Notes Comput Sci. 2009; 5541:184-200 View Article
  12. Liolios K, Chen IM, Mavromatis K, Tavernarakis N, Hugenholtz P, Markowitz VM and Kyrpides NC. The Genomes On Line Database (GOLD) in 2009: status of genomic and metagenomic projects and their associated metadata. Nucleic Acids Res. 2010; 38:D346-D354 View ArticlePubMed
  13. Field D, Garrity G, Gray T, Morrison N, Selengut J, Sterk P, Tatusova T, Thomson N, Allen MJ and Angiuoli SV. The minimum information about a genome sequence (MIGS) specification. Nat Biotechnol. 2008; 26:541-547 View ArticlePubMed
  14. Woese CR, Kandler O and Wheelis ML. Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya. Proc Natl Acad Sci USA. 1990; 87:4576-4579 View ArticlePubMed
  15. Garrity GM, Holt J. Taxonomic outline of the Archaea and Bacteria In: Bergey's Manual of Systematic Bacteriology, 2nd ed. vol. 1. The Archaea, Deeply Branchingand Phototrophic Bacteria Garrity GM, Boone DR and Castenholz RW (eds). 2001; 155-166.
  16. Garrity GM, Holt JG. The Road Map to the Manual. In: Garrity GM, Boone DR, Castenholz RW (eds), Bergey's Manual of Systematic Bacteriology, Second Edition, Volume 1, Springer, New York, 2001, p. 119-169.
  17. Ludwig W, Euzeby J, Whitman WG. Draft taxonomic outline of the Bacteroidetes, Planctomycetes, Chlamydiae, Spirochaetes, Fibrobacteres, Fusobacteria, Acidobacteria, Verrucomicrobia, Dictyoglomi, and Gemmatimonadetes Taxonomic Outline 2008.Web Site
  18. Bernardet JF, Nakagawa Y and Holmes B. Proposed minimal standards for describing new taxa of the family Flavobacteriaceae and emended description of the family. Int J Syst Evol Microbiol. 2002; 52:1049-1070 View ArticlePubMed
  19. . Validation of the publication of new names and new combinations previously effectively published outside the IJSB. List No. 41. Int J Syst Bacteriol. 1992; 42:327-328 View Article
  20. Reichenbach H. Order 1. Cytophagales Leadbetter 1974, 99AL. In: Holt JG (ed), Bergey's Manual of Systematic Bacteriology, First Edition, Volume 3, The Williams and Wilkins Co., Baltimore, 1989, p. 2011-2013.
  21. Bernardet JF, Segers P, Vancanneyt M, Berthe F, Kersters K and Vandamme P. Cutting a Gordian knot: emended classification and description of the genus Flavobacterium, emended description of the family Flavobacteriaceae, and proposal of Flavobacterium hydatis nom. nov. (Basonym, Cytophaga aquatilis Strohl and Tait 1978). Int J Syst Bacteriol. 1996; 46:128-148 View Article
  22. Classification of. Bacteria and Archaea in risk groups. Web Site
  23. Ashburner M, Ball CA, Blake JA, Botstein D, Butler H, Cherry JM, Davis AP, Dolinski K, Dwight SS and Eppig JT. Gene Ontology: tool for the unification of biology. Nat Genet. 2000; 25:25-29 View ArticlePubMed
  24. Klenk HP and Goeker M. En route to a genome-based classification of Archaea and Bacteria? Syst Appl Microbiol. 2010; 33:175-182 View ArticlePubMed
  25. Wu D, Hugenholtz P, Mavromatis K, Pukall R, Dalin E, Ivanova NN, Kunin V, Goodwin L, Wu M and Tindall BJ. A phylogeny-driven genomic encyclopaedia of Bacteria and Archaea. Nature. 2009; 462:1056-1060 View ArticlePubMed
  26. List of growth media used at DSMZ: Web Site
  27. Gemeinholzer B, Dröge G, Zetzsche H, Haszprunar G, Klenk HP, Güntsch A, Berendsohn WG and Wägele JW. The DNA Bank Network: the start from a German initiative. Biopreservation and Biobanking. (In press).
  28. . Web Site
  29. Phrap and Phred for Windows. MacOS, Linux, and Unix. Web Site
  30. Zerbino DR and Birney E. Velvet: algorithms for de novo short read assembly using de Bruijn graphs. Genome Res. 2008; 18:821-829 View ArticlePubMed
  31. Han C, Chain P. 2006. Finishing repeat regions automatically with Dupfinisher. in Proceeding of the 2006 international conference on bioinformatics & computational biology. Edited by Hamid R. Arabnia & Homayoun Valafar, CSREA Press. June 26-29, 2006: 141-146.
  32. Lapidus A, LaButti K, Foster B, Lowry S, Trong S, Goltsman E. POLISHER: An effective tool for using ultra short reads in microbial genome assembly and finishing. AGBT, Marco Island, FL, 2008.
  33. Hyatt D, Chen GL, LoCascio PF, Land ML, Larimer FW and Hauser LJ. Prodigal: prokaryotic gene recognition and translation initiation site identification. BMC Bioinformatics. 2010; 11:119 View ArticlePubMed
  34. Pati A, Ivanova NN, Mikhailova N, Ovchinnikova G, Hooper SD, Lykidis A and Kyrpides NC. GenePRIMP: a gene prediction improvement pipeline for prokaryotic genomes. Nat Methods. 2010; 7:455-457 View ArticlePubMed
  35. Markowitz VM, Ivanova NN, Chen IMA, Chu K and Kyrpides NC. IMG ER: a system for microbial genome annotation expert review and curation. Bioinformatics. 2009; 25:2271-2278 View ArticlePubMed
  36. Sandmann G. Carotenoid biosynthesis and biotechnological application. Arch Biochem Biophys. 2001; 385:4-12 View ArticlePubMed
  37. Nichols D, Bowman J, Sanderson K, Nichols CM, Lewis T, McMeekin T and Nichols PD. Developments with Antarctic microorganisms: culture collections, bioactivity screening, taxonomy, PUFA production and cold-adapted enzymes. Curr Opin Biotechnol. 1999; 10:240-246 View ArticlePubMed
  38. Sakai T, Ishizuka K and Kato I. Isolation and characterization of fucoidan-degrading marine bacterium. Mar Biotechnol. 2003; 5:409-416 View ArticlePubMed
  39. Mavromatis K, Abt B, Brambilla E, Lapidus A, Copeland A, Desphande S, Nolan M, Lucas S, Tice H and Cheng JF. Complete genome sequence of Coraliomargarita akajimensis type strain (04OKA010-24 T). Stand Genomic Sci. 2010; 2:290-299 View ArticlePubMed