Class II. Betaproteobacteria Class. Nov. GEORGE M
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CLASS II. BETAPROTEOBACTERIA 575 Class II. Betaproteobacteria class. nov. GEORGE M. GARRITY, JULIA A. BELL AND TIMOTHY LILBURN Be.ta.pro.te.o.bac.teЈri.a. Gr. n. beta name of second letter of Greek alphabet; Gr. n. Proteus ocean god able to change shape; Gr. n. bakterion a small rod; M.L. fem. pl. n. Betaproteobacteria class of bacteria having 16S rRNA gene sequences related to those of the members of the order Spirillales. The class Betaproteobacteria was circumscribed for this volume on ylophilales, Neisseriales, Nitrosomonadales, “Procabacteriales”, and Rho- the basis of phylogenetic analysis of 16S rRNA sequences; the docyclales. class contains the orders Burkholderiales, Hydrogenophilales, Meth- Type order: Burkholderiales ord. nov. Order I. Burkholderiales ord. nov. GEORGE M. GARRITY, JULIA A. BELL AND TIMOTHY LILBURN Burk.hol.de.ri.aЈles. M.L. fem. n. Burkholderia type genus of the order; -ales ending to denote order; M.L. fem. n. Burkholderiales the Burkholderia order. The order Burkholderiales was circumscribed for this volume on trogen-fixing organisms; and plant, animal, and human patho- the basis of phylogenetic analysis of 16S rRNA sequences; the gens. order contains the families Burkholderiaceae, Oxalobacteraceae, Al- Type genus: Burkholderia Yabuuchi, Kosako, Oyaizu, Yano, caligenaceae, and Comamonadaceae. Hotta, Hashimoto, Ezaki and Arakawa 1993, 398 (Effective pub- Order is phenotypically, metabolically, and ecologically di- lication: Yabuuchi, Kosako, Oyaizu, Yano, Hotta, Hashimoto, verse. Includes strictly aerobic and facultatively anaerobic che- Ezaki and Arakawa 1992, 1268) emend. Gillis, Van, Bardin, Goor, moorganotrophs; obligate and facultative chemolithotrophs; ni- Hebbar, Willems, Segers, Kersters, Heulin and Fernandez 1995, 286. Family I. Burkholderiaceae fam. nov. GEORGE M. GARRITY, JULIA A. BELL AND TIMOTHY LILBURN Burk.hol.de.ri.aЈce.ae. M.L. fem. n. Burkholderia type genus of the family; -aceae ending to denote family; M.L. fem. pl. n. Burkholderiaceae the Burkholderia family. The family Burkholderiaceae was circumscribed for this volume on chemoorganotrophs and obligate and facultative chemolitho- the basis of phylogenetic analysis of 16S rRNA sequences; the trophs. family contains the genera Burkholderia (type genus), Cupriavidus, Type genus: Burkholderia Yabuuchi, Kosako, Oyaizu, Yano, Lautropia, Limnobacter, Pandoraea, Paucimonas, Polynucleobacter,Ral- Hotta, Hashimoto, Ezaki and Arakawa 1993, 398 (Effective pub- stonia, and Thermothrix. Limnobacter was proposed after the cut- lication: Yabuuchi, Kosako, Oyaizu, Yano, Hotta, Hashimoto, off date for inclusion in this volume ( June 30, 2001) and is not Ezaki and Arakawa 1992, 1268) emend. Gillis, Van, Bardin, Goor, described here (see Spring et al. (2001). Hebbar, Willems, Segers, Kersters, Heulin and Fernandez 1995, Family is phenotypically, metabolically, and ecologically di- 286. verse. Includes both strictly aerobic and facultatively anaerobic Genus I. Burkholderia Yabuuchi, Kosako, Oyaizu, Yano, Hotta, Hashimoto, Ezaki and Arakawa 1993, 398VP (Effective publication: Yabuuchi, Kosako, Oyaizu, Yano, Hotta, Hashimoto, Ezaki and Arakawa 1992, 1268) emend. Gillis, Van, Bardin, Goor, Hebbar, Willems, Segers, Kersters, Heulin and Fernandez 1995, 286* NORBERTO J. PALLERONI Burk.hol.deЈri.a. M.L. fem. n. Burkholderia named after W.H. Burkholder, American bacteriologist who discovered the etiological agent of onion rot. Cells single or in pairs, straight or curved rods, but not helical. lm. Motile by means of 4–1.5 ן Dimensions, generally 0.5–1 *Editorial Note: The literature search for the chapter on Burkholderia was completed one or, more commonly, several polar flagella. One species (Burk- in January, 2000. During the course of unavoidable publication delays, a number of new species were described or reclassified after the chapter was completed. It holderia mallei) lacks flagella and is nonmotile. Do not produce was not possible to include these species in the text or to include their characteristics sheaths or prosthecae. No resting stages are known. Gram neg- in the comparative tables. The reader is encouraged to consult the studies listed ative. Most species accumulate poly-b-hydroxybutyrate (PHB) as in the Further Reading section. 576 FAMILY I. BURKHOLDERIACEAE carbon reserve material. Chemoorganotrophs. Have a strictly res- rRNA similarity group II of Palleroni et al. (1973), which includes piratory type of metabolism with oxygen as the terminal electron the genus Burkholderia, have a fatty acid composition containing acceptor. Some species can exhibit anaerobic respiration with C14:0 3OH and C16:0, and C18:1 2OH. Most strains also contained nitrate. Strains of some of the species (B. cepacia, B. vietnamiensis) C16:0 2OH and C16:1. Even though hydroxylated fatty acids are pres- are able to fix N2. Catalase positive. A wide variety of organic ent in the lipids of members of other groups, group II including compounds can be used as sources of carbon and energy for Burkholderia is the only one having hydroxylated fatty acids of 14, growth. Although hydroxylated fatty acids are present in the lip- 16, and 18 carbon atoms (Table BXII.b.1). A recent evaluation ids of members of other genera of aerobic pseudomonads, spe- of the taxonomic significance of fatty acid composition empha- cies of Burkholderia are characterized by the presence of hydroxy sizes the diagnostic value of the above results (Vancanneyt et al., fatty acids of 14, 16, and 18 carbon atoms (C14:0 3OH and C16:0, 1996), confirming earlier findings on this approach for the char- and C16:0 2OH, C16:1, and C18:1). The most characteristic of these acterization of major phylogenetic groups within the pseudo- acids is the C16:0 3OH. Two different ornithine lipids are present monads (Wollenweber and Rietschel, 1990). in strains of some of the species. Over one-half of the species Ornithine-containing lipids in Pseudomonas aeruginosa, P. pu- are pathogenic for plants or animals (including humans). The tida, and B. cepacia represent from 2–15% of the total of extract- genus belongs to the ribosomal RNA similarity group II, which able lipids. The amino acid was not found in the phospholipids can be differentiated from other groups of aerobic pseudomo- that amount to more than 80% of all the extractable lipids (Kawai nads by rRNA/DNA hybridization experiments or by rDNA se- et al., 1988). An analysis of the polar lipids and fatty acids of B. quencing. cepacia has shown that the only significant phospholipids in this -(C of the DNA is: 59–69.6. species are phosphatidyl-ethanolamine and bis(phosphatidyl ם The mol% G Type species: Burkholderia cepacia (Palleroni and Holmes glycerol. These characteristics, taken together with the unusual 1981) Yabuuchi, Kosako, Oyaizu, Yano, Hotta, Hashimoto, Ezaki lipid profiles of B. cepacia, can be used as markers of chemotax- and Arakawa 1993, 398 (Effective publication: Yabuuchi, Kosako, onomic importance (Cox and Wilkinson, 1989a). Oyaizu, Yano, Hotta, Hashimoto, Ezaki and Arakawa 1992, 1271) A striking feature of the cellular composition of B. cepacia is (Pseudomonas cepacia Palleroni and Holmes 1981, 479.) the range of polar lipids, which include two forms (with and without 2-OH fatty acids) of phosphatidyl-ethanolamine and or- FURTHER DESCRIPTIVE INFORMATION nithine amide lipids. Variations in the lipid composition, as well Cell morphology The cells of the genus Burkholderia corre- as in pigmentation and flagellation, were observed as the con- spond in their general characteristics to those of other aerobic sequence of changes in growth temperature and limiting oxygen, pseudomonads: Gram-negative rods, straight or slightly curved, carbon, phosphorus, and magnesium supplies in the medium. with rounded ends, usually motile when suspended in liquid. Phosphorus limitation appears to be the only nutritional factor Motility is due to several polar flagella, but a single flagellum that results in a composition with polar lipids represented only per cell has been reported for B. andropogonis, B. glathei, and B. by ornithine amide lipids (Taylor et al., 1998). norimbergensis.* The single flagellum of B. andropogonis is sheathed Interestingly, the 3-hydroxylated fatty acid of 10 carbon atoms (Fuerst and Hayward, 1969a). One species, B. mallei, is nonmotile is a component of the lipids of B. gladioli but not of those of B. and lacks flagella (Redfearn et al., 1966). cepacia, as indicated by lipid analysis performed on B. gladioli strains isolated from respiratory tract infections in cystic fibrosis Intracellular granules The cells of species of the genus ac- patients (Christenson et al., 1989). cumulate granules of carbon reserve material (poly-b-hydroxy- Differentiation of the plant pathogenic species of Burkholderia b butyrate, PHB, which may be part of a copolymer with poly- - can be done by a direct colony thin-layer chromatographic hydroxyvalerate, PHA). Proteins (“phasins”) have been found to be associated with the granules (Wieczorek et al., 1996). Only B. pseudomallei and some strains of B. mallei use extracellular PHB for growth (Ramsay et al., 1990). All species able to accumulate TABLE BXII.b.1. Fatty acid and ubiquinone composition of the genus PHB can use the intracellular polymer when needed; however, Burkholderia (rRNA group II) and of aerobic pseudomonads of other a,b in the description of some species, the statement “can use PHB” rRNA groups has been included, without indicating whether the PHB was Ribosomal RNA groups endo- or extracellular. Fatty acids I II III IV V The colonies of most species of the genus are smooth, but 3-OH ם ם ם those of the human pathogen B. pseudomallei often have a rough C10:0 ם surface. C11:0 ם C11:0 iso ם ם ם Lipids The first detailed analysis of the fatty acids of aerobic C12:0 ם pseudomonads, using both saprophytic and phytopathogenic C12:0 iso ם strains, demonstrated the possibility of establishing a correlation C13:0 iso ם ם with the phylogenetic subdivision of the genus Pseudomonas, as C14:0 ם C16:1 classically defined, on the basis of rRNA–DNA hybridization 2-OH (ם) Oyaizu and Komagata, 1983). Years later, the results of this sur- C12:0) (ם) vey were confirmed and extended (Stead, 1992).