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The history, biology, and taxonomy of the Cytophaga group.

The first section of this review covers the important characteristics of the genera Cytophaga and Sporocytophaga. The topics discussed include vegetative cell structure, the spreading habit, and degradation of macromolecules. A historical account of these two genera follows, together with a discussion on the definition of, and species differentiation with the genus Cytophaga, and on the taxonomy of Sporocytophaga. The third section deals with the relationships of the cytophagas with the flavobacteria and includes a brief history of Flavobacterium, reassignation of some species to Cytophaga, differentiation from Cytophaga, and a discussion on the definition of the genus Flavobacterium. This is followed by a section dealing with the relationship of Cytophaga with the flexibacteria, starting with an introduction to the diversity of flexing organisms and taxonomic developments, and proceeding with the differentiation within the family Cytophagaceae, and species differentiation in Flexibacter. The concluding section includes a proposed redefinition of Cytophaga, a proposal regarding species conservation in this genus, and discussions on the relationship between the cytophagas and the myxobacteria and on the significance of cytophagas in the environment. The characteristics of all described species of Cytophaga, Flexibacter, and relevant flavobacteria are tabulated and a bibliography is presented.

Bacteriology↗

Phylogenetic diversity of the genus Cytophaga revealed by 16S rRNA sequencing and menaquinone analysis.

To clarify the intra- and intergeneric relationships of the genus Cytophaga, 16S rRNA sequences and respiratory isoprenoid quinones were determined for the type strains of the 21 validly published species and one isolate in the genus Cytophaga. The sequence analysis revealed extreme heterogeneity of this genus, which diverged into nine distinct lines of descent. Each lineage of Cytophaga was characterized by possessing either menaquinone-6 (MK-6) or MK-7. The MK-6-possessing species were located in the two lineages that were remote from MK-7 species. One of the MK-6 lineages was composed only of terrestrial species and the other only of marine species. Flavobacterium aquatile, the type species of the genus Flavobacterium, was located in the MK-6 terrestrial lineage. The terrestrial Cytophaga species with MK-6 should be transferred to the genus Flavobacterium. The marine facultative anaerobes with MK-7 were located in the bacteroides branch, and possessed signature sequences with features intermediate between the bacteroides and the flavobacteria subdivisions. Cytophaga hutchinsonii, the type species of the genus Cytophaga, had a close relationship only with Cytophaga aurantiaca. The genus Cytophaga should be restricted to these two cellulose-degrading species. The genus Cytophaga is so heterogeneous that it should be divided into several genera and higher taxa in accordance with the phylogenetic relationships.

Base Sequence↗

Coaggregation of human oral Cytophaga species and Actinomyces israelii.

A total of 19 strains of oral Cytophaga sp. obtained from subgingival plaque deposits were tested for their ability to coaggregate with strains of Actinomyces israelii, A. viscosus, A. naeslundii, Streptococcus sanguis, S. mutans, S. salivarius, and S. mitis. Coaggregation was observed only with A. israelii. Based on their coaggregation patterns with eight A. israelii strains, the Cytophaga strains were distributed among three distinct groups: those that coaggregated with A. israelii PK16 but not with A. israelii W1011 (ATCC 29322), those that coaggregated with A. israelii ATCC 29322 but not with A. israelii PK16, and those that coaggregated with none of the eight A. israelii strains. In each of the coaggregations, prior heat treatment (85 degrees C, 30 min) of the Cytophaga cells prevented coaggregation, whereas identical treatment of the A. israelii cells had no effect. The ability of A. israelii PK16 to form adherent plaque on a tooth surface previously coated with Cytophaga plaque was tested with one of the coaggregating Cytophaga strains. White patches of A. israelii plaque were found covering both the amber-colored Cytophaga plaque on the cementum surface as well as the enamel surface to which Cytophaga strains do not adhere. Electron micrographs of thin-sectioned mixed-plaque material revealed both cell types in close proximity. In addition, electron micrographs of negatively stained coaggregated cells showed interbacterial adherence between surface fimbrae on A. israelii and outer membrane blebs on the gram-negative Cytophaga sp. The kinetics of binding of A. israelii to spheroidal hydroxyapatite and to root powder were indicative of a high-affinity binding system with comparatively large numbers of available binding sites on both substrata. These results indicate the highly specific nature of Cytophaga sp.--A. israelii recognition. The contribution of such recognition toward the mechanisms that are responsible for the indigenous nature of these oral bacteria is discussed.

Actinomyces↗

Sequence and expression analyses of Cytophaga-like hydrolases in a Western arctic metagenomic library and the Sargasso Sea.

Sequence analysis of environmental DNA promises to provide new insights into the ecology and biogeochemistry of uncultured marine microbes. In this study we used the Sargasso Sea Whole Genome Sequence (WGS) data set to search for hydrolases used by Cytophaga-like bacteria to degrade biopolymers such as polysaccharides and proteins. Analysis of the Sargasso WGS data for contigs bearing both the 16S rRNA genes of Cytophaga-like bacteria and hydrolase genes revealed a cellulase gene (celM) most similar to the gene found in Cytophaga hutchinsonii. A BLAST search of the entire Sargasso Sea WGS data set indicated that celM was the most abundant cellulase-like gene in the Sargasso Sea. However, the similarity between CelM-like cellulases and peptidases belonging to metalloprotease family M42 led us to question whether CelM is involved in the degradation of polysaccharides or proteins. PCR primers were designed for the celM genes in the Sargasso Sea WGS data set and used to identify celM in a fosmid library constructed with prokaryotic DNA from the western Arctic Ocean. Expression analysis of the Cytophaga-like Arctic CelM, which is 63% identical and 77% similar to CelM in C. hutchinsonii, indicated that there was peptidase activity, whereas cellulase activity was not detected. Our analysis suggests that the celM gene plays a role in the degradation of protein by Cytophaga-like bacteria. The abundance of peptidase genes in the Cytophaga-like fosmid clone provides further evidence for the importance of Cytophaga-like bacteria in the degradation of protein in high-molecular-weight dissolved organic matter.

Arctic Regions↗

5S rRNA sequences of representatives of the genera Chlorobium, Prosthecochloris, Thermomicrobium, Cytophaga, Flavobacterium, Flexibacter and Saprospira and a discussion of the evolution of eubacteria in general.

5S rRNA sequences were determined for the green sulphur bacteria Chlorobium limicola, Chlorobium phaeobacteroides and Prosthecochloris aestuarii, for Thermomicrobium roseum, which is a relative of the green non-sulphur bacteria, and for Cytophaga aquatilis, Cytophaga heparina, Cytophaga johnsonae, Flavobacterium breve, Flexibacter sp. and Saprospira grandis, organisms allotted to the phylum 'Bacteroides-Cytophaga-Flavobacterium' and relatives as determined by 16S rRNA analyses. By using a clustering algorithm a dendrogram was constructed from these sequences and from all other known eubacterial 5S RNA sequences. The dendrogram showed differences, as well as similarities, with respect to results obtained by 16S RNA analyses. The 5S RNA sequences of green sulphur bacteria were closely related to one another, and to a cluster containing 5S RNA sequences from Bacteroides and its relatives, including Cytophaga aquatilis. 5S RNA sequences of all other representatives of the 'Bacteroides-Cytophaga-Flavobacterium' phylum as distinguished by 16S RNA analysis failed to group with Bacteroides and related clusters. On the basis of 5S RNA sequences, Thermomicrobium roseum clustered with Chloroflexus aurantiacus, as was expected from 16S RNA analysis.

Base Sequence↗

Occurrence of cytophagas in sewage plants.

With the application of plate count methods and of the KOH-flexirubin test, bacteria belonging to the Cytophaga group were proved to occur regularly in samples from biological sewage treatment facilities. Generally, the percentage of Cytophaga colonies of the total heterotrophic colonies was lowest in the inflow sewage water as compared with the values found in activated sludge, trickling filter, and effluent samples. During an observation period of 16 months, the highest percentages of cytophagas were found in winter samples from activated sludge and trickling filters. Furthermore, cytophagas were shown to have high percentages of the bacteria lytic to polymeric substrates such as cellulose, chitin, dextran, pectin, xylan, and gelatin. Thus, it is suggested that cytophagas may contribute to sewage purification, especially at cold temperatures and by polymer breakdown. Cytophaga strains isolated were shown to have gliding motility, flexirubin pigmentation, and a low guanine plus cytosine base ratio in common. The strains were roughly subdivided into a spreading, a nonspreading, and a cellulolytic group.

Journal Article↗

Polyamine distribution profiles in newly validated genera and species within the Flavobacterium-Flexibacter-Cytophaga-Sphingobacterium complex.

Cellular polyamines of 58 strains belonging to the Flavobacterium-Flexibacter-Cytophaga-Sphingobacterium complex were analysed by HPLC. Homospermidine was found in all species of Flavobacterium, Chryseobacterium, Empedobacter, Myroides, Cellulophaga, Salegentibacter, Psychroserpens and Gelidibacter of the family Flavobacteriaceae. Flavobacterium ferrugineum located outside of this family also contained homospermidine. Cytophaga fermentans and C. xylanolytica belonging to the family Bacteroidaceae contained spermidine. Cytophaga marinoflava and C. latercula belonging to Flavobacteriaceae contained homospermidine. The Cytophaga hutchinsonii/C. aurantiaca group contained homospermidine which was the major polyamine in Flexibacter maritimus/ F. ovolyticus of the family Flavobacteriaceae. The Flexibacter sancti/F filiformis/ Cytophaga arvensicola group, F. elegans, F. ruber, F. canadensis, F. flexilis and F. tractuosus, were located separately in different six clusters, and contained homospermidine. The Flexibacter litoralis/F. polymorphus/F. aggregans group contained spermidine, which was detected in Flexibacter roseolus belonging to a divergent cluster. Sphingobacterium and Pedobacter species of the family Sphingobacteriaceae contained homospermidine. Polyamine profiles serve, as a phenotypic chemotaxonomic marker, for the classification of this complex.

Bacteria↗

Deoxyribonucleic acid relatedness of some menaquinone-producing Flavobacterium and Cytophaga strains.

Nine menaquinone-forming strains of the Flavobacterium--Cytophaga complex with DNA base compositions between 35 and 45 moles percent guanine-plus-cytosine were investigated for genome sizes and DNA relatedness by DNA:DNA hybridization in vitro, using the optically recorded initial reassociation kinetics. Two strains representing C. hutchinsonii and C. marinoflava proved to be related on the 50 percent binding level, i.e. on a level of DNA relatedness commonly found within well-classified conventional genera of bacteria. Strains of C. johnsonae, F. heparinum, F. meningosepticum, F. odoratum, F. pectinovorum, and an unnamed Flavobacterium--Cytophaga strain were found to be interrelated, and linked to the genus Cytophaga, on the 30, or 20 percent binding levels, respectively. These findings indicate that the organisms in question are related to Cytophaga. They therefore should be transferred into the family Cytophagaceae.

Base Composition↗

Influence of physiological factors on the lysis effect of Cytophaga on the red microalga Rhodella reticulata.

The influence of different factors on the lysis of the red microalga, Rhodella reticulata, by Cytophaga sp. LR2 was studied. The pathogenic bacterial strain was more resistant than the alga to the physiological parameters studied, which assured long-term survival of bacteria in algal cultures. Cytophaga sp. LR2 infected R. reticulata at temperatures between 15 and 30 degrees C, in the illuminated as well as the non-illuminated cultures, at pH values between 5.0 and 9.0, and in the presence of NaCl and CaCl2 in the culture medium. SEM showed a different morphology of the bacteria in algal cultures from those of axenic cultures of Cytophaga. Observations of specific associations between algal and bacterial cells revealed that the role of the slime extrusions on the bacterial surface was attachment of Cytophaga to algal cells, and that their clumping leads to rapid lysis.

Culture Media↗

Extra- and intra-cellular lytic effects of Cytophaga sp. LR2 on the red microalgae Rhodella reticulata.

AIMS: To evaluate the lytic activities of crude enzymes from Cytophaga sp. LR2 on Rhodella reticulata cells and isolated algal polysaccharide. METHODS AND RESULTS: The Cytophaga compartment was separated after centrifugation in a cell suspension for 30 min at 18,000 g. The extracellular enzyme was obtained from the supernatant and the intracellular from the pelleted cells after sonication and removal of debris. Algal cells were incubated with extra- or intracellular preparations and sowed onto agar medium. The suppressive effect of the extracellular enzyme on colony-forming units was found to be almost twice as high. The result was still more pronounced when treated cells had been shocked osmotically before seeding. Saccharolytic activity was evaluated by changes in the reducing sugars in the media. Concerning isolated algal polysaccharide, the reducing power of the two bacterial preparates was relatively low. A combined fraction showed the highest lytic activity. Using native and SDS electrophoresis some relation between the prevalence of the extra and intracellular protein patterns was registered. Two of the common components' molecular weight masses of 50 and 21 kDa were found to be reproducible in native- and SDS-containing gel. CONCLUSIONS: Cytophaga sp. LR2 produce extra- and intracellular enzymes active in destroying Rhodella cultures. The agents excreted in the medium are more effective.We suppose that two or three different classes of enzymes are involved in the lysis process. The comparative electrophoresis in this case shows the protein components with predictable functions. SIGNIFICANCE AND IMPACT OF THE STUDY: Combining different simple and reproducible approaches to identify the lytic capability of Cytophaga sp. LR2 on R. reticulata.

Cell Count↗

Occupational lung disease related to cytophaga endotoxin exposure in a nylon plant.

Workers at a nylon plant developed pulmonary disease with systemic symptoms. Differentiating between humidifier fever and hypersensitivity pneumonitis (HP) is challenging. Cytophaga, an endotoxin-producing bacteria, was isolated from the plant air-conditioning system. A number of workers had systemic and pulmonary symptoms. Precipitins to Cytophaga endotoxin were identified. Several workers underwent lung biopsies demonstrating HP. Inhalation challenges with purified Cytophaga endotoxin were performed on three pairs of subjects: group 1, or employees with clinical features and biopsy consistent with HP; group 2, asymptomatic exposed workers with precipitins; and group 3, nonexposed healthy individuals. All subjects had fever and leukocytosis after inhalation challenge. Acute and/or late pulmonary function changes occurred in groups 1 and 2. Group 3 only had acute and transient pulmonary function changes. Cytophaga bacterial endotoxin is capable of inducing HP as well as humidifier fever.

Alveolitis, Extrinsic Allergic↗

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.

Phenotypic data indicate that gliding, yellow/orange-pigmented, agar-digesting bacterial strains were members of the Cytophaga-Flavobacterium-Bacteroides (CFB) group. The strains were isolated from the surface of the marine benthic macroalga Fucus serratus L. and the surrounding seawater at three localities in Danish waters. The bacteria were Gram-negative, flexirubin-negative, aerobic, catalase-positive and oxidase-negative and were psychrophilic and halophilic. All strains utilized D-fructose, L-fucose and alpha-ketobutyric acid and degraded alginic acid, carrageenan, starch and autoclaved yeast cells. Amplification with primers specific for repetitive extragenic palindromic elements by PCR divided the strains of this study into two groups. Both groups showed unique PCR amplification patterns compared to reference strains of the CFB group. Phylogenetic analysis of 16S rDNA sequences showed association of these organisms and [Cytophaga] lytica at the genus level. Hybridization of total chromosomal DNA revealed that the new strains and [Cytophaga] lytica ATCC 23178T were clearly distinct from each other and other previously described species of the CFB group. A new genus is described, Cellulophaga gen. nov. comprising two new species, Cellulophaga baltica gen. nov., sp. nov. (NN015840T = LMG 18535T) and Cellulophaga fucicola gen. nov., sp. nov. (NN015860T = LMG 18536T), as well as the emendation of [Cytophaga] lytica to Cellulophaga lytica gen. nov., comb. nov.

Bacteria↗

Zobellia galactanovorans gen. nov., sp. nov., a marine species of Flavobacteriaceae isolated from a red alga, and classification of [Cytophaga] uliginosa (ZoBell and Upham 1944) Reichenbach 1989 as Zobellia uliginosa gen. nov., comb. nov.

A mesophilic, aerobic, non-flagellated, gliding bacterium, forming yellow colonies and designated DsijT, was isolated from a red alga on the sea-shore of Roscoff, Brittany, France. DsijT was selected for its ability to actively degrade both agars and carrageenans. The Gram-negative cells occurred singly or in pairs as long rods. The temperature range for growth was 13-45 degrees C, with an optimum at 35 degrees C. The pH range for growth at 35 degrees C was from 6.0 to 8.5, with an optimum around pH 7.0. The NaCl concentrations required for growth at 35 degrees C and pH 7.0 ranged from 5 to 60 g l(-1), with an optimum around 25 g l(-1). The G+C content of the genomic DNA was 42-43 mol%. Phylogenetic analysis of 16S rRNA gene sequences indicated that strain DsijT is closely related to [Cytophaga] uliginosa DSM 2061T. Phenotypic features, however, allowed DsijT and [Cytophaga] uliginosa strains to be distinguished on the basis of ten traits (spreading behaviour, assimilation of eight compounds and amylase production). Their total protein profiles were also different and DNA-DNA hybridization experiments confirmed that DsijT constitutes a new species, distinct from [Cytophaga] uliginosa. Based on the phenotypic features and the phylogenetic relationships of the Flavobacteriaceae, a new genus designated Zobellia gen. nov. is proposed to include Zobellia galactanovorans gen. nov., sp. nov., while [Cytophaga] uliginosa becomes Zobellia uliginosa comb. nov. The type strain of Zobellia galactanovorans is DsijT (= DSM 12802T = CIP 106680T).

Anti-Bacterial Agents↗

Development of techniques to genetically manipulate members of the genera Cytophaga, Flavobacterium, Flexibacter, and Sporocytophaga.

The Bacteroides-Cytophaga-Flavobacterium branch of the eubacterial phylogenetic tree contains a diverse group of bacterial species. Techniques for the genetic manipulation of Bacteroides spp. are well developed (A. A. Salyers, N. B. Shoemaker, and E. P. Guthrie, Crit. Rev. Microbiol. 14:49-71, 1987). Recently we developed techniques to genetically manipulate the gliding bacterium Cytophaga johnsonae (M. J. McBride and M. J. Kempf, J. Bacteriol. 178:583-590, 1996). We now demonstrate that some of these techniques allow genetic manipulation of a number of environmentally or medically significant bacteria in this group. The Bacteroides transposon Tn4351 was introduced into Cytophaga hutchinsonii, Cytophaga succinicans, Flavobacterium meningosepticum, Flexibacter canadensis, Flexibacter sp. strain FS1, and Sporocytophaga myxococcoides by conjugation. Tn4351 integrated itself into the host chromosomes and conferred erythromycin resistance. We isolated several auxotrophic mutants of Flavobacterium meningosepticum following Tn4351 mutagenesis. The C. johnsonae-Escherichia coli shuttle vector pCP11 functioned in C. succinicans but not in the other bacteria. pLYL03 did not replicate in any of these bacteria and should function as a convenient suicide vector. The identification of a system of gene transfer, a selectable marker, a suicide vector, and a transposon that functions in these diverse bacteria allows genetic manipulations to be performed.

Blotting, Southern↗

A Cytophaga species endotoxin as a putative agent of occupation-related lung disease.

A previous study suggested that a biologically active bacterial endotoxin was a putative agent of lung disease in a textile-producing facility. The endotoxin was isolated from the biomass growing in a chilled-water spray air humidification system. The bacterial flora of the air humidification system were isolated and taxonomically identified to the genus level. By using indirect immunofluorescence assays, a serologically reactive Cytophaga species was identified. A serologically reactive, biologically active (Limulus assay) endotoxin was purified from phenol extracts of the Cytophaga species. The endotoxin contained sugars, hexosamines, and lipids identical to those found in the humidifier biomass endotoxin. All subjects with biopsy-proven and suspected lung disease had antibodies directed toward the purified Cytophaga endotoxin. The data suggest that the Cytophaga endotoxin is the putative agent of lung disease in the textile facility.

Bacterial Infections↗

Novel psychrophilic and thermolabile L-threonine dehydrogenase from psychrophilic Cytophaga sp. strain KUC-1.

A psychrophilic bacterium, Cytophaga sp. strain KUC-1, that abundantly produces a NAD(+)-dependent L-threonine dehydrogenase was isolated from Antarctic seawater, and the enzyme was purified. The molecular weight of the enzyme was estimated to be 139,000, and that of the subunit was determined to be 35,000. The enzyme is a homotetramer. Atomic absorption analysis showed that the enzyme contains no metals. In these respects, the Cytophaga enzyme is distinct from other L-threonine dehydrogenases that have thus far been studied. L-Threonine and DL-threo-3-hydroxynorvaline were the substrates, and NAD(+) and some of its analogs served as coenzymes. The enzyme showed maximum activity at pH 9.5 and at 45 degrees C. The kinetic parameters of the enzyme are highly influenced by temperatures. The K(m) for L-threonine was lowest at 20 degrees C. Dead-end inhibition studies with pyruvate and adenosine-5'-diphosphoribose showed that the enzyme reaction proceeds via the ordered Bi Bi mechanism in which NAD(+) binds to an enzyme prior to L-threonine and 2-amino-3-oxobutyrate is released from the enzyme prior to NADH. The enzyme gene was cloned into Escherichia coli, and its nucleotides were sequenced. The enzyme gene contains an open reading frame of 939 bp encoding a protein of 312 amino acid residues. The amino acid sequence of the enzyme showed a significant similarity to that of UDP-glucose 4-epimerase from Staphylococcus aureus and belongs to the short-chain dehydrogenase-reductase superfamily. In contrast, L-threonine dehydrogenase from E. coli belongs to the medium-chain alcohol dehydrogenase family, and its amino acid sequence is not at all similar to that of the Cytophaga enzyme. L-Threonine dehydrogenase is significantly similar to an epimerase, which was shown for the first time. The amino acid residues playing an important role in the catalysis of the E. coli and human UDP-glucose 4-epimerases are highly conserved in the Cytophaga enzyme, except for the residues participating in the substrate binding.

Alcohol Oxidoreductases↗

Psychrophilic valine dehydrogenase of the antarctic psychrophile, Cytophaga sp. KUC-1: purification, molecular characterization and expression.

We found the occurrence of valine dehydrogenase in the cell extract of a psychrophilic bacterium, Cytophaga sp. KUC-1, isolated from Antarctic seawater and purified the enzyme to homogeneity. The molecular mass of the enzyme was determined to be approximately 154 kDa by gel filtration and that of the subunit was 43 kDa by SDS/PAGE: the enzyme was a homotetramer. The enzyme required NAD+ as a coenzyme, and catalyzed the oxidative deamination of L-valine, L-isoleucine, L-leucine and the reductive amination of alpha-ketoisovalerate, alpha-ketovalerate, alpha-ketoisocaproate, and alpha-ketocaproate. The reaction proceeds through an iso-ordered bi-bi mechanism. The enzyme was highly susceptible to heat treatment and the half-life at 45 degrees C was estimated to be 2.4 min. The kcat/Km (micro(-1).s(-1)) values for L-valine and NAD+ at 20 degrees C were 27.48 and 421.6, respectively. The enzyme showed pro-S stereospecificity for hydrogen transfer at the C4 position of the nicotinamide moiety of coenzyme. The gene encoding valine dehydrogenase was cloned into Escherichia coli (Novablue), and the primary structure of the enzyme was deduced on the basis of the nucleotide sequence of the gene encoding the enzyme. The enzyme contains 370 amino-acid residues, and is highly homologous with S. coelicolor ValDH (identity, 46.7%) and S. fradiae ValDH (43.1%). Cytophaga sp. KUC-1 ValDH contains much lower numbers of proline and arginine residues than those of other ValDHs. The changes probably lead to an increase in conformational flexibility of the Cytophaga enzyme molecule to enhance the catalytic activity at low temperatures.

Amino Acid Oxidoreductases↗

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.

A group of strains with potent extracellular enzymic activity were isolated from the surfaces of the chain-forming sea-ice diatom Melosira and from an unidentified macrophyte collected from the Eastern Antarctic coastal zone. 16S rDNA sequence analysis indicated that the strains belonged to the genus Cellulophaga and showed greatest similarity to the species Cellulophaga baltica (sequence similarity 97%). Phenotypic characteristics, DNA base composition and DNA-DNA hybridization values clearly separate the Antarctic strains from Cellulophaga baltica and other Cellulophaga species. Thus, the strains form a distinct and novel species and have the proposed name Cellulophaga algicola sp. nov. (type strain IC166T = ACAM 630T). In addition, it was recognized that the species Cytophaga uliginosa (ZoBell and Upham 1944) Reichenbach 1989, a species phylogenetically remote from the type species of the genus Cytophaga, possessed 16S rDNA sequences and phenotypic and chemotaxonomic traits similar to those of other Cellulophaga species. Thus, it was proposed that the species Cytophaga uliginosa be renamed as Cellulophaga uliginosa comb. nov.

Antarctic Regions↗