Antonie van Leeuwenhoek

, Volume 101, Issue 4, pp 803–810 | Cite as

Salinimonas lutimaris sp. nov., a polysaccharide-degrading bacterium isolated from a tidal flat

Original Paper

Abstract

A Gram-negative, non-motile, non-endospore-forming bacterial strain, designated DPSR-4T, was isolated from a tidal flat sediment on the southern coast of Korea. Strain DPSR-4T grew optimally at 25–30°C, at pH 7.0–7.5 and in the presence of 2% (w/v) NaCl. A Neighbour-Joining phylogenetic tree based on 16S rRNA gene sequences revealed that strain DPSR-4T clustered with Salinimonaschungwhensis BH030046T by a high bootstrap resampling value of 99.7%. Strain DPSR-4T exhibited 96.2% 16S rRNA gene sequence similarity to that of S. chungwhensis BH030046T and 93.7–96.6% sequence similarity to the sequences of type strains of Alteromonas species. Strain DPSR-4T contained Q-8 as the predominant ubiquinone and iso-C15:0 2-OH and/or C16:1ω7c, C16:0 and C18:1ω7c as the major fatty acids. The major polar lipids detected in strain DPSR-4T and S. chungwhensis KCTC 12239T were phosphatidylglycerol, phosphatidylethanolamine and an unidentified phospholipid. The DNA G+C content was 53.4 mol%. Differential phenotypic properties and phylogenetic distinctiveness of strain DPSR-4T demonstrated that this strain is distinguishable from the sole recognized species of the genus Salinimonas, S. chungwhensis. On the basis of phenotypic, chemotaxonomic and phylogenetic data, strain DPSR-4T is considered to represent a novel species of the genus Salinimonas, for which the name Salinimonas lutimaris sp. nov. is proposed. The type strain is DPSR-4T (KCTC 23464T, CCUG 60743T).

Keywords

Salinimonas lutimaris Novel species Polysaccharides Tidal flat 

Introduction

In the course of screening of novel polysaccharide-degrading microorganisms from a tidal flat sediment on the southern coast in Korea, many novel bacterial strains that belong to the Proteobacteria have been isolated. One of these isolates, designated DPSR-4T, which shows degradative activities against several polysaccharides, is the subject of this work. Comparative 16S rRNA gene sequence analysis showed that this novel strain was most closely related phylogenetically to members of the genera Alteromonas and Salinimonas of the Gammaproteobacteria. The genus Alteromonas was first proposed by Baumann et al. (1972) and, at the time of writing, comprises 9 species with validly published names (Baumann et al. 1972; Euzéby 1997, http://www.bacterio.cict.fr; Yoon et al. 2003, 2004; Van Trappen et al. 2004; Ivanova et al. 2005; Martínez-Checa et al. 2005; Chiu et al. 2007; Vandecandelaere et al. 2008). The genus Salinimonas was first created by Jeon et al. (2005) with the description of Salinimonaschungwhensis, which is the sole recognized species of the genus to date (Euzéby 1997). The aim of the present work was to determine the exact taxonomic position of strain DPSR-4T by using a polyphasic characterization that included the determination of chemotaxonomic and phenotypic properties and a detailed phylogenetic investigation based on 16S rRNA gene sequences.

Materials and methods

Bacterial strains and culture conditions

Tidal flat sediments were collected from the coast of Geojedo and used as the source for the isolation of bacterial strains. Strain DPSR-4T was isolated by means of the standard dilution plating technique at 25°C on marine agar 2216 (MA; Becton–Dickinson). Strain DPSR-4T was maintained on MA at 4°C for short-term preservation and as a glycerol suspension (20%, w/v in distilled water) at −80°C for long-term preservation. Strain DPSR-4T has been deposited in the Korean Collection for Type Cultures (KCTC; South Korea) and the Culture Collection, University of Göteborg (CCUG; Sweden) as KCTC 23464T and CCUG 60743T, respectively. S. chungwhensis KCTC 12239T, which was used as a reference strain for fatty acid analysis, DNA G + C content analysis and phenotypic characterization, was obtained from the Korean Collection for Type Cultures (KCTC), Daejeon, South Korea.

Cell biomass of strain DPSR-4T for DNA extraction and for the analyses of isoprenoid quinones and polar lipids was obtained from cultures grown for 2 days in marine broth 2216 (MB; Becton–Dickinson) at 30°C. Cell biomass of S. chungwhensis KCTC 12239T for DNA extraction and for polar lipid analysis was obtained from cultures grown for 2 days in MB at 30°C. For fatty acid methyl ester analysis, cell mass of strain DPSR-4T and S. chungwhensis KCTC 12239T was harvested from MA plates after incubation for 2 days at 30°C.

Morphological, physiological and biochemical characterization

The morphological, physiological and biochemical characteristics of strain DPSR-4T were investigated using routine cultivation on MA at 25°C. The cell morphology was examined by light microscopy (Olympus BX51) and transmission electron microscopy. Flagellation was determined by using a Philips CM-20 transmission electron microscope with cells from exponentially growing cultures. For this purpose, the cells were negatively stained with 1% (w/v) phosphotungstic acid and the grids were examined after being air-dried. The Gram reaction was determined by using the bioMérieux Gram stain kit according to the manufacturer’s instructions. Growth at various temperatures (4, 10, 20, 25, 28, 30, 35, 37, 40 and 45°C) was measured on MA. The pH range for growth was determined in MB that was adjusted to various pH values (pH 4.5–9.5, using increments of 0.5 pH units) by the addition of HCl or Na2CO3. Growth in the presence of 0–22.0% (w/v) NaCl was investigated by using marine broth prepared according to the formula of the Difco medium except that NaCl was excluded. Growth under anaerobic conditions was determined after incubation in a Forma anaerobic chamber on MA and on MA supplemented with potassium nitrate (0.1%, w/v), both of which had been prepared anaerobically under nitrogen atmosphere. Catalase and oxidase activities were determined as described by Cowan and Steel (1965). Hydrolysis of casein, starch, hypoxanthine, tyrosine and xanthine was tested on MA by using the substrate concentrations described by Cowan and Steel (1965). Hydrolysis of aesculin, gelatin, Tweens 20, 40, 60 and 80 and urea and nitrate reduction were investigated as described previously (Lányí 1987) with the modification that artificial seawater was used for preparation of media. The artificial seawater contained (l−1 distilled water): 23.6 g NaCl, 0.64 g KCl, 4.53 g MgCl2·6H2O, 5.94 g MgSO4·7H2O and 1.3 g CaCl2·2H2O (Bruns et al. 2001). Degradation of various polysaccharides was tested on basal medium agar [12 g gellan gum, 1 g yeast extract, 0.5 g NH4Cl, 50 ml 1 M Tris/HCl (pH 7.4) and 5 g low melting agarose l−1 artificial seawater] containing alginate, carboxymethylcellulose (CMC), ι-carrageenan, colloidal chitin, curdlan, pectin from apple peels and birch wood xylan, respectively. All polysaccharides were purchased from Sigma except curdlan and low viscous alginate (Wako Chemicals). Colloidal chitin was prepared using shrimp shell according to a modification of the method of Rodriguez-Kabana et al. (1983). 5 g of shrimp shell was dissolved in 50 ml of concentrated HCl and stirred at 4°C overnight. The chitin solution was slowly poured into 50% chilled ethanol with vigorous stirring. The pH of the mixture was neutralized by NaOH and after centrifugation for 30 min at 6,000 rpm, the supernatant was decanted and the colloid was dried in a desiccator. Alginate lyase activity was revealed by flooding the agar with a 10% (w/v) cetylpyridinium chloride solution. Degradation of CMC and chitin was revealed by flooding the agar with Congo red aqueous solution. Degradation of carrageenan, pectin and xylan was revealed by flooding the agar with 10× diluted Lugol’s iodine solution. For the detection of curdlanase activity, 0.005% (w/v) of aniline blue was added to the medium before autoclaving and the activity was revealed as halos formed around colonies (Mahasneh and Stewart 1980). Utilization of substrates as sole carbon and energy sources was tested as described by Baumann and Baumann (1981) using supplementation with 2% (v/v) Hutner’s mineral base (Cohen-Bazire et al. 1957) and 1% (v/v) vitamin solution (Staley 1968). Acid production from carbohydrates was determined as described by Leifson (1963). Susceptibility to antibiotics was tested on MA plates using antibiotic discs containing the following concentrations: ampicillin (10 μg), carbenicillin (100 μg), cephalothin (30 μg), chloramphenicol (100 μg), gentamicin (30 μg), kanamycin (30 μg), lincomycin (15 μg), neomycin (30 μg), novobiocin (5 μg), oleandomycin (15 μg), penicillin G (20 U), polymyxin B (100 U), streptomycin (50 μg) and tetracycline (30 μg). Other biochemical tests were performed with the API ZYM system (bioMérieux).

16S rRNA gene sequencing and phylogenetic analysis

Chromosomal DNA was isolated and purified according to the method described previously (Yoon et al. 1996), with the exception that RNase T1 was used in combination with RNase A to minimize contamination with RNA. The 16S rRNA gene amplification was performed using two universal primers (5′-GAGTTTGATCCTGGCTCAG-3′ and 5′-AGAAAGGAGGTGATCCAGCC-3′) as described previously (Yoon et al. 1998) and the PCR products were purified by using a QIAquick PCR purification kit (Qiagen). Sequencing of the amplified 16S rRNA gene was performed as described by Yoon et al. (2003). Alignment of sequences was carried out with CLUSTAL W software (Thompson et al. 1994). Gaps at the 5′ and 3′ ends of the alignment were omitted from further analysis. Phylogenetic trees were inferred by using three tree-making algorithms, the neighbour-joining (Saitou and Nei 1987), maximum-likelihood (Felsenstein 1981) and maximum-parsimony (Kluge and Farris 1969) methods implemented within the PHYLIP package (Felsenstein 1993). Evolutionary distance matrices for the neighbour-joining method were calculated by the algorithm of Jukes and Cantor (1969) using the program DNADIST. The stability of relationships was assessed by a bootstrap analysis based on 1,000 resampling of the neighbour-joining dataset by using the programs SEQBOOT, DNADIST, NEIGHBOR and CONSENSE of the PHYLIP package.

Chemotaxonomic characterization

Isoprenoid quinones were extracted according to the method of Komagata and Suzuki (1987) and analyzed using reversed-phase HPLC and a YMC ODS-A (250 × 4.6 mm) column. The isoprenoid quinones were eluted by a mixture of methanol/isopropanol (2:1, v/v), using a flow rate of 1 ml min−1 at room temperature and detected by UV absorbance at 270 nm. Fatty acids were saponified, methylated and extracted using the standard protocol of the MIDI (Sherlock Microbial Identification System, version 4.0). The fatty acids were analysed by GC (Hewlett Packard 6890) and identified by using TSBA40 database of the Microbial Identification System (Sasser 1990). Polar lipids were extracted according to the procedures described by Minnikin et al. (1984), and separated by two-dimensional TLC using chloroform/methanol/water (65:25:3.8, v/v) for the first dimension and chloroform/methanol/acetic acid/water (40:7.5:6:1.8, v/v) for the second dimension as described by Minnikin et al. (1977). Individual polar lipids were identified by spraying with the ethanolic molybdophosphoric acid, molybdenum blue, ninhydrin and α-naphthol reagents (Minnikin et al. 1984; Komagata and Suzuki 1987) and the Dragendorff’s reagent (Sigma). The DNA G+C content was determined by the method of Tamaoka and Komagata (1984) with the modification that DNA was hydrolysed and the resultant nucleotides were analysed by reversed-phase HPLC equipped with a YMC ODS-A (250 × 4.6 mm) column. The nucleotides were eluted by a mixture of 0.55 M NH4H2PO4 (pH 4.0) and acetonitrile (40:1, v/v), using flow rate of 1 ml min−1 at room temperature and detected by UV absorbance at 270 nm.

Results and discussion

Morphological, cultural, physiological, and biochemical characteristics

Strain DPSR-4T was found to be aerobic, Gram-negative, non-spore-forming short rods or rods. It was non-motile whereas S. chungwhensis BH030046T was motile by means of a polar flagellum. Strain DPSR-4T grew optimally at 25–30°C and pH 7.0–7.5. Strain DPSR-4T is a moderately halophile as it grew optimally in the presence of 2% (w/v) NaCl; it grew also without NaCl, whereas S. chungwhensis BH030046T did not grow without NaCl (Jeon et al. 2005). Strain DPSR-4T showed catalase and oxidase activities but no urease activity. It did not reduce nitrate to nitrite. Strain DPSR-4T degraded agarose, alginate, ι-carrageenan, curdlan, pectin, starch and xylan but did not degrade chitin and CMC. Morphological, cultural, physiological and biochemical characteristics of strain DPSR-4T are given in the species description (see below) or in Table 1.
Table 1

Phenotypic characteristics of Salinimonas lutimaris DPSR-4T and Salinimonas chungwhensis KCTC 12239T

Characteristic

1

2

Motility

+

Optimal growth temperature (°C)

25–30

30–35

Growth at

  

 40°C

+

 0% NaCl

+

 20% NaCl

+

Hydrolysis of

  

 Hypoxanthine

+

 Urea

+

Utilization of

  

  l-Arabinose

+

  d-Fructose

+

  d-Galactose

+

  Salicin

+

  d-Xylose

+

Acid production from

  

  l-Arabinose

+

  d-Glucose

+

 Maltose

+

  d-Ribose

+

  d-Trehalose

+

  d-Xylose

+

Susceptibility to antibiotics

  

 Carbenicillin

+

 Neomycin

+

 Polymyxin B

+

 Streptomycin

+

 Tetracycline

+

Enzyme activity (API ZYM)

  

 β-Galactosidase

+

DNA G + C mol%

53.4

50.6

Strains: 1, S. lutimaris DPSR-4T; 2, S. chungwhensis KCTC 12239T. All data from this study except for optimal growth temperature, growth at 4 and 40°C, Gram-staining, morphology, catalase, oxidase, nitrate reduction and hydrolysis of substrates; these data are from Jeon et al. (2005). Both species are Gram-staining-negative and rod-shaped. Both strains are positive for catalase, oxidase, hydrolysis of aesculin, casein, gelatin, starch, Tween 80 and tyrosine, utilization of d-cellobiose, d-glucose, maltose, sucrose, d-trehalose, acetate and pyruvate, susceptibility to chloramphenicol, gentamicin, kanamycin and oleandomycin, and activity of alkaline phosphatase, esterase (C 4), esterase lipase (C 8), leucine arylamidase, acid phosphatase and naphthol-AS-BI-phosphohydrolase. Both strains are negative for growth at 4°C, nitrate reduction, hydrolysis of xanthine, utilization of d-mannose, citrate, benzoate, formate, l-malate, succinate and l-glutamate, acid production from d-fructose, lactose, d-mannose, melibiose, d-raffinose, l-rhamnose and d-mannitol, susceptibility to ampicillin, cephalothin, penicillin G, lincomycin, and novobiocin, activity of lipase (C 14), valine arylamidase, cystine arylamidase, trypsin, α-chymotrypsin, α-galactosidase, β-glucuronidase, α-glucosidase, β-glucosidase, N-acetyl-β-glucosaminidase, α-mannosidase and α-fucosidase

Symbol: +, positive reaction; –, negative reaction

Phylogenetic analysis

The almost complete 16S rRNA gene sequence of strain DPSR-4T determined in this study comprised 1,457 nucleotides (approximately 96% of the Escherichiacoli 16S rRNA sequence). In the neighbour-joining phylogenetic tree based on 16S rRNA gene sequences, strain DPSR-4T joined the type strain of S. chungwhensis with a bootstrap resampling value of 99.7% and this cluster joined the clade comprising Alteromonas species by a bootstrap resampling value of 94.2% (Fig. 1). The relationship between strain DPSR-4T and S. chungwhensis BH030046T was also maintained in the trees constructed using the maximum-likelihood and maximum-parsimony algorithms (Supplementary Figures 1 and 2). Strain DPSR-4T exhibited 16S rRNA gene sequence similarity values of 96.2% to S. chungwhensis BH030046T. It exhibited slightly higher 16S rRNA gene sequence similarity values (96.6 and 96.5%) to the type strains of Alteromonashispanica and Alteromonasgenovensis than to S. chungwhensis BH030046T, and 93.7–96.1% sequence similarity to the type strains of the other Alteromonas species. It has been shown that the 16S rRNA gene sequence similarity may not be necessarily consistent with the topology of phylogenetic trees (Nedashkovskaya et al. 2006; Xue et al. 2006; Yoon et al. 2010), as it the case here. S. chungwhensis BH030046T exhibited 92.2–94.7% 16S rRNA gene sequence similarity to the type strains of Alteromonas species.
Fig. 1

Neighbour-joining phylogenetic tree based on 16S rRNA gene sequences showing the positions of Salinimonas lutimaris DPSR-4T, Salinimonas chungwhensis BH030046T and some other related taxa. Bootstrap values (expressed as percentages of 1,000 replications) of >70% are shown at branching points. Filled circles indicate that the corresponding nodes were also recovered in the trees generated with the maximum-likelihood and maximum parsimony algorithms. Vibrio cholerae CECT 514T (GenBank accession number, X76337) was used as an outgroup. Scale bar, 0.01 substitutions per nucleotide position

Chemotaxonomic characteristics

The predominant isoprenoid quinone detected in strain DPSR-4T was ubiquinone-8 (Q-8) which is the same as that found in S. chungwhensis BH030046T (Jeon et al. 2005). The cellular fatty acid profiles of strain DPSR-4T and S. chungwhensis KCTC 12239T, which were analyzed in this study, are shown in Table 2. The major fatty acids (>10% of total fatty acids) found in strain DPSR-4T were iso-C15:0 2-OH and/or C16:1ω7c, C16:0 and C18:1ω7c (Table 2). The fatty acid profiles were similar in both strains, even though there were differences in the proportions of some fatty acids (Table 2). The fatty acids (C16:0 N alcohol, C16:1ω7c and 10-methyl-C17:0), which were present as traces or not detected in S. chungwhensis BH030046T but were detected as significant amounts in Alteromonas species as described by Jeon et al. (2005), were not detected in strain DPSR-4T. The major polar lipids detected in strain DPSR-4T were phosphatidylglycerol, phosphatidylethanolamine and an unidentified phospholipid, and the major polar lipids of S. chungwhensis KCTC 12239T analysed also in this study were the same with those of strain DPSR-4T (Supplementary Figure 3). The DNA G+C content of strain DPSR-4T was 53.4 mol%, a little higher than that of S. chungwhensis KCTC 12239T (50.6 mol%), which was also determined in this study (Table 1).
Table 2

Cellular fatty acid compositions (%) of Salinimonas lutimaris DPSR-4T and Salinimonas chungwhensis KCTC 12239T

Fatty acid

1

2

Straight-chain fatty acid

  

 C12:0

5.2

3.4

 C13:0

0.7

0.4

 C14:0

2.3

2.6

 C15:0

1.7

2.1

 C16:0

22.3

23.2

 C17:0

4.5

4.1

 C18:0

1.4

1.0

Branched fatty acid

  

 Iso-C16:0

1.1

0.7

 Anteiso-C17:0

0.7

0.7

Unsaturated fatty acid

  

 C15:1ω8c

0.5

1.1

 C17:1ω8c

3.3

4.4

 C18:1ω7c

11.3

11.0

Hydroxy fatty acid

  

 C10:0 3-OH

1.4

0.9

 C11:0 3-OH

2.3

1.4

 C12:0 3-OH

4.7

2.4

 C12:1 3-OH

0.6

0.7

Cyclo-C17:0

3.5

11-methyl-C18:1ω7c

0.8

Summed featuresa

  

 1

0.9

0.6

 2

5.6

4.6

 3

24.8

27.3

 7

0.3

1.2

Strains: 1, S. lutimaris DPSR-4T; 2, S. chungwhensis KCTC 12239T. All data from this study; cells of both strains were grown for 2 days at 30°C on MA. Fatty acids that represented <0·5% in both strains were omitted. −, Not detected

aSummed feature represent groups of two or three fatty acids which could not be separated by GLC with the MIDI system. Summed feature 1 contained C13:0 3-OH and/or iso-C15:1. Summed feature 2 contained C14:0 3-OH and/or iso-C16:1. Summed feature 3 contained iso-C15:0 2-OH and/or C16:1ω7c. Summed feature 7 contained unknown fatty acid (ECL 18.846), C19:1ω6c and/or cyclo-C19:0ω10c

Conclusion

It is appropriate to place strain DPSR-4T within the genus Salinimonas on the basis of the absence of distinct chemotaxonomic properties between strain DPSR-4T and S. chungwhensis as well as the phylogenetic data. Strain DPSR-4T was distinguishable from S. chungwhensis by differences in several phenotypic characteristics including motility, optimal growth temperature, hydrolysis of hypoxanthine and urea, utilization of some substrates, acid production from some substrates, susceptibility to some antibiotics and activity of some enzymes, all of which were determined under the same conditions and methods (Jeon et al. 2005; Table 1). The phylogenetic distinctiveness of strain DPSR-4T, together with the differential phenotypic properties, is sufficient to show that this strain is separate from S. chungwhensis (Stackebrandt and Goebel 1994). Therefore, on the basis of the phenotypic, chemotaxonomic and phylogenetic data, strain DPSR-4T is considered to represent a novel species of the genus Salinimonas, for which the name Salinimonas lutimaris sp. nov. is proposed.

Description of Salinimonas lutimaris sp. nov

Salinimonas lutimaris (lu.ti.ma’ris. L. n. lutum mud; L. gen. n. maris of the sea, marine; N.L. gen. n. lutimaris of a marine mud).

Cells are Gram-negative, non-spore-forming, non-flagellated and short rods or rods (0.4–0.7 × 0.9–3.5 μm). Colonies on MA are circular to slightly irregular, flat, smooth, glistening, light grayish yellow in colour and 1.0–2.0 mm in diameter after incubation for 2 days at 25°C; Colonies on MA at 30°C are wrinkled. Optimal growth occurs at 25–30°C; growth occurs at 10 (weak) and 37°C, but not at 4 and 40°C. Optimal pH for growth is between 7.0 and 7.5; growth occurs at pH 6.0, but not at pH 5.5. Optimal growth occurs in the presence of 2% (w/v) NaCl; growth occurs at 0–20.0% (w/v) NaCl. Growth does not occur under anaerobic conditions on MA and on MA supplemented with nitrate. Catalase- and oxidase-positive. Nitrate reduction is negative. Aesculin, casein, gelatin, starch, Tweens 20, 40, 60 and 80 and tyrosine are hydrolysed, but hypoxanthine, xanthine and urea are not. d-Cellobiose, d-fructose, d-galactose, d-glucose, maltose, sucrose, salicin, d-trehalose, acetate and pyruvate are utilized, but l-arabinose, d-mannose, d-xylose, citrate, benzoate, formate, l-malate, succinate and l-glutamate are not. Acid is not production from l-arabinose, d-fructose, d-glucose, lactose, maltose, d-mannose, melibiose, d-raffinose, l-rhamnose, d-ribose, d-trehalose, d-xylose and d-mannitol. Susceptible to chloramphenicol, gentamicin, kanamycin, neomycin, oleandomycin, streptomycin and tetracycline, but not to ampicillin, carbenicillin, cephalothin, penicillin G, polymyxin B, lincomycin and novobiocin. In assays with API ZYM system, alkaline phosphatase, esterase (C 4), esterase lipase (C 8), leucine arylamidase, acid phosphatase, naphthol-AS-BI-phosphohydrolase and β-galactosidase are present, but lipase (C 14), valine arylamidase, cystine arylamidase, trypsin, α-chymotrypsin, α-galactosidase, β-glucuronidase, α-glucosidase, β-glucosidase, N-acetyl-β-glucosaminidase, α-mannosidase and α-fucosidase are not absent. The predominant ubiquinone is Q-8. The major fatty acids (>10% of total fatty acids) are iso-C15:0 2-OH and/or C16:1ω7c, C16:0 and C18:1ω7c. The major polar lipids are phosphatidylglycerol, phosphatidylethanolamine and an unidentified phospholipid. The DNA G+C content is 53.4 mol % (HPLC). Other phenotypic properties are shown in Table 1. The type strain, DPSR-4T (KCTC 23464T, CCUG 60743T), was isolated from a tidal flat sediment of Geojedo on the South Sea, Korea.

Notes

Acknowledgments

This work was supported by the Program for Collection, Management and Utilization of Biological Resources (grant M10867010003) and BK21 program from the Ministry of Education, Science and Technology (MEST) of the Republic of Korea.

Supplementary material

10482_2011_9695_MOESM1_ESM.ppt (112 kb)
Supplementary material 1 (PPT 112 kb)
10482_2011_9695_MOESM2_ESM.ppt (112 kb)
Supplementary material 2 (PPT 112 kb)
10482_2011_9695_MOESM3_ESM.ppt (126 kb)
Supplementary material 3 (PPT 126 kb)

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Copyright information

© Springer Science+Business Media B.V. 2011

Authors and Affiliations

  1. 1.Department of Food Science and BiotechnologySungkyunkwan UniversityJangan-guSouth Korea
  2. 2.Korea Research Institute of Bioscience and Biotechnology (KRIBB)YuseongSouth Korea

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