Brevibacterium Lengkap

Genus
Actinobacteria/Actinobacteria/Micrococcales/Brevibacteriaceae/
Brevibacterium
Breed 1953a, 13AL emend. Collins, Jones, Keddie and Sneath 1980, 6
..........................................................................................................................................................................................
Martha E. Trujillo, Universidad de Salamanca, Departamento de Microbiologı̂a y Genêtica, Edificio Departamental,
Lab. 205, Campus Unamuno, Salamanca 37007, Spain
Michael Goodfellow, University of Newcastle, School of Biology, Ridley Building, Newcastle upon Tyne NE1 7RU, UK
Bre.v.i.bac.te’ri.um. L. adj. brevis short; L. neut. n. bac- DNA G+C content (mol%): 55–70.
terium a small rod; N.L. neut. n. Brevibacterium a short Type species: Brevibacterium linens (Wolff 1910)
rodlet. Breed 1953a, 13AL emend. Collins, Jones, Keddie and
Aerobic, catalase-positive, non-acid-fast, asporoge- Sneath 1980, 7 (“Bacterium linens” Wolff 1910).
..................................................................................
nous actinomycetes which have a rod–coccus cell cycle
when grown on complex media. Usually nonmotile, Aerobic, catalase-positive, non-acid-fast, asporogenous acti-
but some species show motility. Both rod and coccoid nomycetes which have a rod–coccus cell cycle when grown on
complex media. Usually nonmotile, but some species show
forms are Gram-stain-positive, but some strains and
motility. Both rod and coccoid forms are Gram-stain-positive,
older cultures decolorize readily. Cells are variable in
but some strains and older cultures decolorize readily. Cells
length but generally are 0.6–1.0 μm in diameter; those
are variable in length but generally are 0.6–1.0 μm in diame-
from older cultures (3–7 d) are composed mainly or ter; those from older cultures (3–7 d) are composed mainly
entirely of coccoid cells or coccobacilli. On transfer to or entirely of coccoid cells or coccobacilli. On transfer to
a suitable fresh medium, these forms grow out to give a suitable fresh medium, these forms grow out to give the
the irregular, slender rods characteristic of exponential irregular, slender rods characteristic of exponential phase
phase cultures. Many cells are arranged at an angle to cultures. Many cells are arranged at an angle to give V-forms.
give V-forms. Primary branching may occur but not Primary branching may occur but not a true mycelium. Respi-
a true mycelium. Respiratory mode of metabolism. ratory mode of metabolism. Optimum growth temperature is
Optimum growth temperature is 20–37∘ C depending 20–37∘ C depending on the species and strain. Grows well on
on the species and strain. Grows well on peptone-yeast peptone-yeast extract agar at neutral pH. Little or no acid is
extract agar at neutral pH. Little or no acid is pro- produced from glucose or other carbohydrates in a peptone
medium. Proteinases produced. The cell-wall peptidoglycan
duced from glucose or other carbohydrates in a
contains meso-diaminopimelic acid as the diamino acid and is
peptone medium. Proteinases produced. The cell-wall
of the A1γ type. Contains dihydrogenated menaquinones with
peptidoglycan contains meso-diaminopimelic acid as
eight isoprene units as the predominant isoprenologue and
the diamino acid and is of the A1γ type. Contains dihy- major proportions of branched cellular fatty acids, notably
drogenated menaquinones with eight isoprene units as C15:0 anteiso and C17:0 anteiso. Mycolic acids are absent.
the predominant isoprenologue and major proportions Isolated from clinical material, dairy products, poultry,
of branched cellular fatty acids, notably C15:0 anteiso and marine and terrestrial habitats.
and C17:0 anteiso. Mycolic acids are absent. DNA G+C content (mol%): 55–70.
......................................................................................................................................................................................................
Bergey’s Manual of Systematics of Archaea and Bacteria, Online © 2015 Bergey’s Manual Trust. This article is © 2012 Bergey’s Manual Trust.
DOI: 10.1002/9781118960608.gbm00062. Published by John Wiley & Sons, Inc., in association with Bergey’s Manual Trust.
2 Bergey’s Manual of Systematics of Archaea and Bacteria
Type species: Brevibacterium linens (Wolff 1910) Breed the type strains of the two latter species which were described
1953a, 13AL emend. Collins, Jones, Keddie and Sneath 1980, around the same time in 2004.
7 (“Bacterium linens” Wolff 1910).
Cell morphology
Number of validated species: 24 ...................................................................................
Brevibacteria grow on a range of media though morphol-
Further descriptive information ogy and staining reactions are best seen on EYGA medium
(Cure and Keddie, 1973). Strains tend to show a distinct
Phylogeny rod–coccus cycle when grown on complex media (Collins,
................................................................................... 2006; Collins et al., 1980). Both young (6–24 h) and older
The genus Brevibacterium forms a distinct line of descent (3–7 d) cultures should be examined as cell shape (Figure 2)
within the high G+C containing actinomycetes (Cai and and staining properties change during the growth cycle
Collins, 1994; Stackebrandt et al., 1997; Zhi et al., 2009). The (Crombach, 1974; Cure and Keddie, 1973; Mulder and
24 species assigned to the genus form a monophyletic group Antheumisse, 1963). Coccoid cells are mainly found in
in the 16S rRNA gene tree (Figure 1) and share sequence older cultures and when transferred to fresh media give
similarities which range from 92.5% between Brevibacterium rise to slender irregular rods. Coccoid and rod-shaped
aurantiacum and Brevibacterium paucivorans, to 99.8% between cells are Gram-stain-positive but generally decolorize with
Brevibacterium sanguinis and Brevibacterium celere. DNA–DNA age (Jones and Keddie, 1986). Members of the genus are
relatedness experiments have not been carried out between typically nonmotile, though motility has been detected in
FIGURE 1. Neighbor-joining tree based on 16S rRNA gene sequences showing the phylogenetic relationships of Brevibacterium
species. Bootstrap values are shown at branching points. Micrococcus luteus ATCC 381T was used as outgroup. Bar = 1 substitu-
tion per 100 nucleotides.
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This article is © 2012 Bergey’s Manual Trust. Published by John Wiley & Sons, Inc., in association with Bergey’s Manual Trust.
Bergey’s Manual of Systematics of Archaea and Bacteria 3
Brevibacterium album (Tang et al., 2008), Brevibacterium iod- Diphosphatidylglycerol (DPG) and phosphatidylglycerol
inum (Colwell et al., 1969), and Brevibacterium samyangense (PG) usually feature as major components in brevibacterial
(Lee, 2006). polar lipid patterns (Bhadra et al., 2008; Collins et al., 1983a;
Kämpfer et al., 2010). Collins et al. (1980) reported both
Chemotaxonomy
................................................................................... of these lipids and a glycolipid, dimannosyldiacylglycerol,
in strains of Brevibacterium iodinum and Brevibacterium linens.
The cell-wall peptidoglycan contains meso-diaminopimelic
Some species also contain phosphatidylinositol (PI) (Collins
acid (Collins et al., 1983a; Fiedler et al., 1981; Kämpfer et
et al., 1980, 1983a; Heyrman et al., 2004; Lee, 2008), though
al., 2010; Keddie and Cure, 1977, 1978; Pascual and Collins,
the presence of this component has been shown to vary with
1999; Tang et al., 2008; Yamada and Komagata, 1972b) and
cultural conditions (Jones and Keddie, 1986). In addition to
is of the A1γ type (Schleifer and Kandler, 1972). A unique
DPG and PG, an unidentified glycolipid has been detected
feature of Brevibacterium paucivorans is the presence of glycine in Brevibacterium picturae (Heyrman et al., 2004) and Brevibac-
and glutamic acid as non-peptidoglycan components of the terium sandarakinum (Kämpfer et al., 2010). Cadavarine and
cell wall (Wauters et al., 2001). Galactose is present in the putrescine are characteristic components of brevibacterial
cell wall of some species such as Brevibacterium album (Tang et polyamine patterns (Altenburgera et al., 1996; Kämpfer et
al., 2008), Brevibacterium avium (Pascual and Collins, 1999), al., 2010).
Brevibacterium casei, and Brevibacterium epidermidis (Sharpe
et al., 1977, 1978). Teichoic acids were initially detected Colony morphology
...................................................................................
in the cell walls of Brevibacterium casei (Fiedler and Bude,
Opaque, white or pale orange, convex colonies (0.5–1.0 mm
1989), Brevibacterium epidermidis (Fiedler and Bude, 1989),
in diameter) with entire edges and smooth, shiny surfaces
Brevibacterium iodinum (Anderton and Wilkinson, 1980) and
are formed within 2 d on nutrient agar. Colonies become
Brevibacterium linens (Fiedler et al., 1981) but have been found
larger (2.0–4.0 mm in diameter) after 4–7 d (Jones and
in additional Brevibacterium species (Gavrish et al., 2005).
Keddie, 1986). Opaque gray-white, convex colonies with
Brevibacterium species have similar fatty acid patterns. All
smooth, shiny surfaces up to 2 mm in diameter are formed
contain major amounts of anteiso- and iso-methyl branched
on blood agar after 24 h; they later increase in size and turn
acids together with small amounts of straight-chain saturated
slightly greenish or yellowish in color (Funke et al., 1997).
acids. The major components are 12-methyltetradecanoic
Collins et al. (1983a) described colonies of Brevibacterium
(C15:0 anteiso) and 14-methylhexadecanoic (C17:0 anteiso)
casei as gray-white in color, but in an earlier study of the same
acids (Bousfield et al., 1983; Bowie et al., 1972; Collins et
strains, Sharpe et al. (1976) recorded the slow production
al., 1980, 1983a). These fatty acids usually account for over
of a brown, water-soluble pigment when the organisms were
75% of the total fatty acid composition (Bhadra et al., 2008;
cultured on milk agar.
Funke and Carlotti, 1994; Gruner et al., 1993; Ivanova et
Brevibacterium linens forms yellow to deep orange-yellow
al., 2004; Kati et al., 2010) though not in species such as colonies when grown on a range of media, with pigment pro-
Brevibacterium mcbrellneri (McBride et al., 1993), Brevibacterium duction in most strains being light-dependent (Crombach,
otitidis (Wauters et al., 2003), and Brevibacterium paucivorans 1974; Mulder et al., 1966). Jones et al. (1973) suggested
(Wauters et al., 2001). In contrast, mycolic acids are not that the pigment of Brevibacterium linens was a carotenoid
found in brevibacteria (Goodfellow et al., 1976; Heyrman et which could be recognized by distinctive color reactions
al., 2004; Keddie and Cure, 1977; Lee, 2006). occurring when pigmented growth was treated with either
Menaquinones are the sole respiratory quinones present glacial acetic acid or solutions of strong bases. The pigments
in the cell membranes of brevibacteria. The major com- of Brevibacterium linens were subsequently shown to consist
ponent is usually dihydrogenated menaquinone with eight of the aromatic carotenoids 3,3′ -dihydroxyisorenieratene,
isoprene units [MK-8(H2 )] (Collins et al., 1980, 1983a, isorenieratene, and 3-hydroxyisorenieratene (Kohl et al.,
Heyrman et al., 2004, Kämpfer et al.,2010; Lee, 2006, 2008), 1983).
though smaller amounts of other components such as MK-8, The characteristic metallic purple to blue color of Brevibac-
MK-7(H2 ), and MK-9(H2 ) may be present (Collins et al., terium iodinum is due to the production of purple, extracellu-
1979, 1980, 1983a). In contrast, Brevibacterium casei and Bre- lar crystals of the phenazine derivative iodinin on and in the
vibacterium linens contain comparable amounts of MK-7(H2 ) colonies and adjacent medium (Collins et al., 1980; Colwell et
and MK-8(H2 ) (Collins et al., 1983a). al., 1969; Davis, 1939; Sneath, 1960). The synthesis of iodinin
......................................................................................................................................................................................................
FIGURE 2. Brevibacterium linens (ATCC 9175) when grown on medium EYGA at 25∘ C; inoculum coccoid cells as in (D). (A)
After 6 h, showing outgrowth of rods from coccoid cells; (B) after 12 h; (C) after 24 h; and (D), after 3 d. Bars = 10 μm.
(Reproduced with permission from Keddie and Jones, 1981. The genus Brochothrix (formerly Microbacterium thermosphactum,
McLean and Sulzbacher). In The Prokaryotes: a Handbook on Habitats, Isolation, and Identification of Bacteria (edited by
Starr, Stolp, Trüper, Balows and Schlegel). Springer, New York, pp. 1866–1869.)
is enhanced on certain media such as Blood Agar Base (Difco) Most strains are proteolytic, metabolizing casein, gelatin, and
plates incubated at 30∘ C for 2 d, but is not affected by light milk (Bousfield, 1972; Colwell et al., 1969; Crombach, 1974;
(Sneath, 1960). Keddie and Jones, 1981; Wauters et al., 2003). In general,
acid is not produced from glucose or other carbohydrates
Nutritional and growth conditions
................................................................................... (Bousfield, 1972; Collins et al., 1983a; Jones, 1975; Kati et al.,
2010), though a range of compounds is used as sole carbon
Brevibacteria grow well on routine laboratory media such
sources (Crombach, 1974; Ivanova et al., 2004; Kämpfer et
as blood, nutrient, tryptone soy, yeast extract–malt extract,
al., 2010; Tang et al., 2008; Yamada and Komagata, 1972a).
and yeast extract–peptone agars. They tolerate and are
Several species including Brevibacterium casei, Brevibacterium
sometimes stimulated by the addition of NaCl to the medium
epidermidis, Brevibacterium linens, and Brevibacterium paucivo-
(Crombach, 1974; Heyrman et al., 2004; Kati et al., 2010;
rans produce methanethiol from L-methionine (Pitcher and
Lee, 2006, 2008; Mulder et al., 1966; Sharpe et al., 1977).
Noble, 1978; Sharpe et al., 1976, 1977, 1978; Wauters et al.,
They grow well at neutral pH and at 30∘ C, though not all
2003).
strains grow optimally at this temperature. Brevibacterium
linens, for instance, grows best between 20–25∘ C (Keddie Genetics
and Jones, 1981), Brevibacterium avium at 37∘ C (Pascual and ...................................................................................
Collins, 1999), Brevibacterium casei and Brevibacterium epider- Little is known about brevibacterial genetics, though plasmids
midis between 30 and 37∘ C (Pitcher and Noble, 1978; Sharpe designated pBL100 and pBL33 have been isolated from Bre-
et al., 1976, 1977), and Brevibacterium iodinum at 28∘ C (Jones, vibacterium linens strains (Kato et al., 1989; Sandoval et al.,
1975). 1985) and purified (Holtz et al., 1992).
Metabolism Antibiotic sensitivity

................................................................................... ...................................................................................
Brevibacteria are aerobic, catalase-positive, chemoorgan- Few studies have been directed toward establishing the antibi-
otrophic actinomycetes with an oxidative type of metabolism. otic sensitivity patterns of Brevibacterium species (Bhadra et
......................................................................................................................................................................................................
al., 2008; Colwell et al., 1969; Ivanova et al., 2004; Jones, otitidis from an ear infection (Pascual et al., 1996a), and
1975; McBride et al., 1993; Sneath, 1960), though this is Brevibacterium paucivorans from clinical samples (Wauters et
likely to change given the clinical significance of species al., 2001).
such as Brevibacterium casei (Brazzola et al., 2000; Funke et
Ecology
al., 1997). Members of this taxon tend to be resistant to ...................................................................................
β-lactams, ciprofloxacin, clindamycin, and erythromycin and
Brevibacteria have been isolated from numerous habitats,
sensitive to gentamicin, rifampin, and tetracycline (Funke
notably ones with a high salt concentration. Most species have
et al., 1996). Pitcher and Noble (1978) noted that nearly
been derived either from dairy milk products, human skin,
90% of methanethiol-producing skin brevibacteria isolated
or clinical material. The usual habitat of Brevibacterium linens
from the feet of patients infected with Trichophyton species
is the exterior of surface-ripened cheeses such as Limburger
were resistant to penicillin. The antibiotic sensitivity profiles
(Brennan et al., 2002), but it also occurs on cheeses such as
of individual Brevibacterium species are cited in the species
Brick, Camembert, and Roquefort (El-Erian, 1969; Mulder et
descriptions.
al., 1966). This organism is considered to be responsible for
Bacteriocins and antimicrobials the surface color of red-smear cheeses and also, in part, by its
................................................................................... proteolytic activity, to the ripening of such cheeses (Brennan
Several bacteriocins and antimicrobials are produced et al., 2002). It is for this latter reason that Brevibacterium
by Brevibacterium linens (Kato et al., 1984, 1991). These linens is often inoculated onto the surface of smear cheeses
include Linocin A which shows inhibition to some other during the early stages of ripening, either as a commercial
members of this species but not to other Brevibacterium preparation or as the so-called “old-young” smearing method
species or to Corynebacterium, Listeria, and Micrococcus species whereby young cheeses are washed with a brine suspension
(Valdes-Stauber and Scherer, 1994). Further, Linenscin OC2 of microorganisms from the surface of mature cheese.
from another Brevibacterium linens strain inhibits food borne Strains now classified as Brevibacterium casei (Collins et
pathogens including Listeria monocytogenes and Staphylococcus al., 1983a) have been isolated from cheese curd, milk, and
aureus. In addition, Brevibacterium linens strains isolated from cheddar cheese, though the primary habitat of this organism
brine used to salt red-smear cheeses produce an antimi- is considered to be the human skin (Sharpe et al., 1976).
crobial agent active against Listeria species (Martin et al., Brevibacterium epidermidis is also thought to be part of the
1995). resident flora of human skin (Collins et al., 1983a; Pitcher
Clinical significance and Noble, 1978). Other Brevibacterium species isolated
................................................................................... from human material include Brevibacterium luteolum (corrig.
Members of the genus Brevibacterium, notably Brevibacterium Wauters et al., 2003), Brevibacterium mcbrellneri (McBride et
casei, have quite recently been associated with human infec- al., 1993), Brevibacterium otitidis (Pascual et al., 1996a), and
tions (Collins, 2006; Funke et al., 1997; McCaughey and Brevibacterium paucivorans (Wauters et al., 2004b).
Damani, 1991; Pitcher and Malnick, 1984). They have been Brevibacteria have been recovered from marine habi-
implicated as agents of continuous ambulatory peritoneal tats, as seen by the isolation of Brevibacterium celere from
dialysis (Antoniou et al., 1997; Gruner et al., 1993; Hummel the degraded thallus of the brown alga Fucus evanescens
et al., 1996; Wauters, Van Bosterhaut, Avesani, Cuvelier, Char- (Ivanova, Christen, Alexeeva, Zhukova, Gorshkova, Lysenko,
lier, Janssens and Delmmée, 2000), corneal ulcers (Ghosheh Mikhailov and Nicolau, 2004), Brevibacterium marinum from
et al., 2007), endocarditis (Dass et al., 2002), osteomyelitis sea water (Lee, 2008), Brevibacterium oceani from deep-sea
(Neumeister et al., 1993), peritonitis (Cannon et al., 2005), sediment, and Brevibacterium samyangense from beach sand
septicemia (Carlotti et al., 1993; Kaukoranta-Tolvanen et al., (Lee, 2006). Additional species have been isolated from
1995; Lina et al., 1994; Lipsky et al., 1982; Reinert et al., 1995; more unusual sources such as Brevibacterium album from
Ulrich et al., 2006), and sepsis in AIDS patients (Brazzola saline soil (Tang et al., 2008), Brevibacterium salitolerans
et al., 2000; Janda et al., 2003). In addition, Brevibacterium from a hypersaline lake sediment sample (Guan et al.,
luteolum has been isolated from an ear discharge (Wauters et 2010), Brevibacterium picturae from a damaged mural paint-
al., 2003), Brevibacterium mcbrellneri from infected genital hair ing (Heyrman et al., 2004), Brevibacterium pityocampae from
of patients with white piedra in association with Trichosporon healthy larvae of the moth, Thaumelopoea pityocampae, a
beigelli (McBride et al., 1993), Brevibacterium massiliense from pest of Pinus species in Mediterranean countries, including
an ankle discharge (Roux and Raoult, 2009), Brevibacterium Turkey (Kati et al., 2010), and Brevibacterium sandarakinum
......................................................................................................................................................................................................
from an indoor wall colonized by fungi (Kämpfer et al., A medium which has been reported to be particularly suit-
2010). able for the isolation of Brevibacterium linens from cheese is
that of Albert et al. (1944). On this medium, which contains
Enrichment and isolation procedures ripened cheese, Brevibacterium linens is reputed to give good
pigment production in 5–7 d at 21∘ C or at room temperature,
Brevibacteria can be isolated on nutrient media based on especially when plates are incubated in an oxygen-enriched
glucose, peptone, and yeast extract, as exemplified by try- atmosphere (Keddie and Jones, 1981).
pone soy agar supplemented with 4% (w/v) NaCl (TSAS;
[Oxoid] 17 g; peptone, 3 g; NaCl, 9 g; K2 HPO4 , 2.5 g; glu- Isolation from skin
...................................................................................
cose, 2.5 g; agar, 12 g; pH is adjusted to 7.0. The medium is
sterilized at 121∘ C for 15 min). Isolation of the organisms Swabs moistened in physiological saline are rubbed firmly
is usually achieved by the use of such media supplemented over the appropriate skin area and then streaked onto TSAS
with cheese, milk, and an elevated concentration of NaCl or agar or Blood Agar Base no. 2 (Difco). After incubation at
30∘ C for 5–7 d colonies are randomly selected and examined
sea water, as appropriate, depending on the environmental
for coryneform morphology. Further identification tests are
sample. Inoculated plates should be incubated at 20–25∘ C
then carried out.
for 3–7 d, though temperatures of 30–37∘ C are better for the
isolation of skin brevibacteria. It is important when isolating Isolation from other sources
Brevibacterium linens that plates are exposed to light during ...................................................................................
some period of active growth to enhance pigment formation Brevibacterium marinum and Brevibacterium samyangense were
(Crombach, 1974; Mulder et al., 1966). A convenient practice isolated from sea water and sediment samples, respec-
is to incubate plates at 25∘ C until small colonies are evident tively, using starch casein agar (Küster and Williams, 1964)
and then to expose plates to light on the laboratory bench prepared using 60% natural sea water (Lee, 2006, 2008).
for the rest of the incubation period (Jones and Keddie, Similarly, Brevibacterium oceani was isolated by plating out
1986). a sediment sample suspended in 2% (w/v) NaCl onto
Since isolation media are non-selective for brevibac- yeast extract–peptone plates (Bhadra et al., 2008) and Bre-
teria, it is necessary to randomly select pigmented and vibacterium album was isolated by inoculating a saline soil
nonpigmented colony types to establish whether cells have suspension onto glycerol-asparagine agar (Tang et al., 2008).
a coryneform morphology. When identification is limited Similarly, Brevibacterium picturae strains were isolated on R2A
to a consideration of isolates showing a rod–coccus growth agar (Difco) supplemented with 1% (w/v) NaCl and on
cycle care must be taken to distinguish brevibacteria from tryptic soy agar supplemented with 10% (w/v) NaCl, Brevibac-
organisms such as arthrobacters and rhodococci. Tests for terium pityocampae by inoculating suspensions of the contents
this purpose are available (see section on differentiating the of surface-sterilized insect larvae onto nutrient agar (Kati et
genus Brevibacterium from other genera). al., 2010), and Brevibacterium sandarakinum by enriching a
sample taken from a wall colonized by fungi in a 10 ml 0.9
Isolation from cheese (w/v) NaCl solution containing 0.01% (w/v) Tween 80 and
...................................................................................
plating dilutions onto M79 agar (Kämpfer et al., 2010).
Samples of cheese or cheese curd are homogenized in 2%
(w/v) trisodium citrate supplemented with NaCl (El-Erian, Maintenance procedures
1969). This medium is based on Tryptone Soy Broth (Oxoid)
though similar products are available from other sources such Brevibacteria may be preserved for some months by stab inoc-
as BBL and Difco. Colonies growing on the isolation plates are ulation in nutrient agar (Oxoid) or similar media in screw-
examined as described above. capped containers. After overnight incubation with the caps
Sharpe et al. (1976) isolated brevibacteria from various loose at 25 or 30∘ C, the caps should be screwed tightly to pre-
dairy products by direct plating onto a medium containing vent evaporation. The containers may be stored at room tem-
30% (w/v) skim milk and 2% (w/v) agar. All isolates were perature (∼20∘ C), or better at ∼4∘ C in the dark. Cultures may
tested for the production of methanethiol from L-methionine be preserved for longer periods (at least 7 years) by freezing
(see Procedures for testing for special characteristics, below) and in glass beads at −70∘ C (Feltham et al., 1978) or by lyophiliza-
then examined for a coryneform morphology. tion.
......................................................................................................................................................................................................
and contain meso-A2 pm in the wall peptidoglycan. Mor-

Procedures for testing for special characters
phologically, members of the genus can be confused with
Corynebacterium, Dietzia, Gordonia, Rhodococcus, and Williamsia
Presence of rod–coccus cycle strains but can be readily distinguished from the latter as
...................................................................................
they have fatty acids rich in iso- and anteiso-methyl branched
This may be demonstrated by using the methods described by components but lack mycolic acids and phosphatidylinos-
Cure and Keddie (1973). itol dimannosides (Collins et al., 1980, 1983a; Goodfellow
and Maldonado, 2006). Similarly, brevibacteria can be dis-
Cell-wall composition
................................................................................... tinguished from arthrobacters and cellulomonads as they
contain meso-A2 pm and not lysine or ornithine in the wall
Satisfactory results may be obtained with one of the methods
peptidoglycan (Collins, 2006). Bousfield et al. (1983) sug-
described by Bousfield et al. (1985).
gested that computer-assisted interpretation of fatty acid
Detection of mycolic acids data combined with morphological examination may be
................................................................................... sufficient to identify strains as members of the genera Bre-
This may be achieved by whole-organism methanolysis as vibacterium, Arthrobacter, Cellulomonas, Corynebacterium, Kurthia,
described by Minnikin et al. (1984). or Oerskovia.
Polar lipid patterns Taxonomic comments

...................................................................................
These may be determined by two dimensional TLC analysis The genus Brevibacterium was proposed by Breed (1953a)
of free lipid extracts (Collins et al., 1983a). to accommodate several Gram-stain-positive, asporogenous,
nonbranching rod-shaped bacteria which had previously
Test for the production of methanethiol
................................................................................... been classified in the genus Bacterium; Brevibacterium linens
was designated the type species. The genus was recognized
Sharpe et al. (1978) recommended the DTNB method of
in the 7th edition of the Manual where it was classified
Laakso (1976) because it is more rapid and, in their expe-
with the genus Kurthia in the family Brevibacteriaceae. At that
rience, much more sensitive than the Conway diffusion time, the genus Brevibacterium contained 23 species which
method (Sharpe et al., 1978). were described as typically short, unbranched rods that were
Organisms grown on nutrient agar slopes are suspended usually nonmotile; no mention was made of a coryneform
(to give an E580 of 10) in 0.05 M Tris/HCl buffer at pH 8.0; morphology.
1 ml samples are then incubated in rubber-stoppered test It was subsequently shown that Brevibacterium linens had a
tubes with 12.5 mM L-methionine and 0.25 mM 5.5-dithiols coryneform morphology and that it exhibited a rod–coccus
(2-nitrobenzoic acid) (DTNB) in a total volume of 5 ml growth cycle similar to that seen in Arthrobacter globiformis
for 2 h at 30∘ C. Methanethiol production is indicated by (Mulder and Antheumisse, 1963; Schefferle, 1966). In some
the development of a yellow color in the test incubations. earlier numerical taxonomic studies, it was even proposed
Positive tests are discernable by eye, but weak methanethiol that Brevibacterium linens should be transferred to the genus
producers may be confirmed colorimetrically at E420 after Arthrobacter as Arthrobacter linens (Bousfield, 1972; da Silva
centrifugation of the bacterial growth. Controls (a) without and Holt, 1965; Davis and Newton, 1969). However, Fiedler
cell suspensions and (b) without L-methionine should be et al. (1970) showed not only that the peptidoglycan in Bre-
incubated. vibacterium linens was quite different from that in Arthrobacter
In addition, Pitcher and Malnick (1984) described a rapid globiformis and closely related species, but also that the 28
method for the detection of methanethiol. species named Brevibacterium which they examined were
heterogeneous in peptidoglycan type. It was mainly for these
Differentiation of the genus Brevibacterium from reasons that the genus Brevibacterium was listed as incertae sedis
other genera in the 8th edition of the Manual (Rogosa and Keddie, 1974).
The heterogeneity of the genus was underpinned by
16S rRNA gene sequencing is currently the most effective way chemotaxonomic (Keddie and Cure, 1978; Minnikin et al.,
of distinguishing brevibacteria from other actinomycetes, 1978; Schleifer and Kandler, 1972) and numerical taxonomic
notably from those which show a rod–coccus growth cycle (Jones, 1978; Seiler et al., 1977) studies and Brevibacterium
......................................................................................................................................................................................................
linens was seen as a distinct taxon which could form the basis iodinum is diagnostic for this organism (Clemo and Daglish,
of a redefined genus (Jones, 1975; Keddie and Cure, 1977; 1950; Davis, 1939; Sneath, 1960). In practice, closely related
Sharpe et al., 1978; Yamada and Komagata, 1972a). These and Brevibacterium species are distinguished using a combination
further studies led Collins et al. (1980) to redefine the genus of 16S rRNA gene sequence and phenotypic data (Kämpfer
Brevibacterium and at the same time to reclassify the organism et al., 2010; Kati et al., 2010). Preliminary evidence suggests
previously designated “Chromobacterium iodinum” as Brevibac- that Brevibacterium species can be separated on the basis of
terium iodinum. Later, two groups of methanethiol-producing their MALDI-TOF mass spectra (Roux and Raoult, 2009).
coryneform bacteria, one isolated from cheese and milk and
the other from human skin, were classified as Brevibacterium List of species of the genus Brevibacterium
casei and Brevibacterium epidermidis by Collins et al. (1983a).
The methanethiol-producing organisms from cheese and
Brevibacterium linens
milk were first isolated and well described by Sharpe et al.
(Wolff 1910) Breed 1953a, 13AL emend. Collins et al.,
(1976, 1977, 1978) and those from skin by Pitcher and Noble 1980, 7 (“Bacterium linens” Wolff 1910)
(1978) and Sharpe et al. (1978). All of these investigators ...................................................................................
emphasized the similarity between these isolates and Brevibac-
li’nens. L. part. adj. linens spreading over, smearing.
terium linens, though at that time the genus Brevibacterium was
Aerobic, Gram-stain-positive, nonmotile actinomycetes
incertae sedis.
which have a coryneform morphology and show a rod–coccus
The four Brevibacterium species mentioned above were
growth cycle when grown on complex media. Surface
recognized in the last edition of Bergey’s Manual of System-
colonies on nutrient agar are small (0.1–0.2 mm in diam-
atic Bacteriology by Jones and Keddie (1986) who also listed
eter) after 1–2 d, becoming larger (2.5 mm) on extended
additional species that required further study so that they
incubation. Mature colonies are shiny, convex, and have
might be reclassified with confidence. In the meantime, new
entire margins. On peptone–yeast extract media, an orange
species have been assigned to the genus Brevibacterium, which
now encompasses 24 validly published species forming a to orange-red pigment is produced. Pigment production by
monophyletic branch in the 16S rRNA gene tree (Figure 1). most strains, including the type strain, is light dependent.
In contrast, species previously associated with the genus Pigmented growth gives characteristic color reactions with
have been transferred to other genera, as exemplified by various acids and bases. Optimum growth temperature is
the reclassification of Brevibacterium ammoniagenes Cooke and 20–25∘ C; growth is generally poor or absent at 5∘ C and
Keith 1927 and Brevibacterium stationis (ZoBell and Upham 37∘ C. Grows in the presence of 10% (w/v) NaCl. Acid is not
1944) Breed 1953a as Corynebacterium ammoniagenes comb. produced from glucose or other sugars in peptone media.
nov. Collins 1987 and Corynebacterium stationis comb. nov. Gelatin is liquefied, hippurate hydrolyzed, and extracellular
Bernard et al. 2010. DNase produced. Weakly oxidase-positive. Methanethiol is
produced from L-methionine. Additional phenotypic charac-
Differentiation of the species of the genus teristics are shown in Table 1. The major fatty acids are C17:0
Brevibacterium anteiso and C15:0 anteiso.
Common on surface-ripened smear cheeses.
Brevibacterium species can be distinguished using a combina-
Source: harzer cheese.
tion of phenotypic tests (Table 1) but not always with confi-
DNA G+C content (mol%): 60–64 (Tm ).
dence, as a common set of tests are not yet available for the
Type strain: ATCC 9172, CIP 101125, DSM 20425, HAMBI
delineation of all species. Brevibacterium linens is most easily
recognized by its ability to produce yellow to deep orange 2038, JCM 1327, NBRC 12142, NRRL B-4210, VKM Ac-2112.
colonies (Jones et al., 1973; Mulder et al., 1966), though it Sequence accession no. (16S rRNA gene): X77451.
needs to be remembered that DNA relatedness studies show Additional comments: Brevibacterium linens has been shown
that this taxon encompasses at least two species (Fiedler et to be heterogeneous on the basis of nutritional proper-
al., 1981). The color reaction of Brevibacterium linens illicited ties (Mulder et al., 1966), electrophoretic protein patterns
by various chemicals has been used to distinguish members of (Foissy, 1974) and DNA–DNA relatedness studies (Fiedler
this taxon (Grecz and Dack, 1961), but this test is not exclu- et al., 1981). Fiedler and his colleagues showed that strains
sively selective (Jones et al., 1973). The production of dis- designated Brevibacterium linens in public and private culture
tinct purple extracellular crystals of iodinin by Brevibacterium collections fell into at least two distinct species.
......................................................................................................................................................................................................
TABLE 1. Characteristics differentiating species of the genus Brevibacteriuma
B. ravenspurgense
B. sandarakinum
B. aurantiacum
B. samyangense
B. paucivorans
B. pityocampae
B. epidermidis
B. salitolerans
B. massiliense
B. mcbrellneri
B. antiquum
B. sanguinis
B. marinum
B. permense
B. luteolum
B. iodinum
B. picturae
B. otitidis
B. avium
B. album
B. oceani
B. linens
B. celere
B. casei
Characteristic
Colony colorb yo wh wo o gw gw wy wy g y wb y wb po wy g o wh y gw wy bry lr gw
Colony texturec s s nd nd s s s s s s s s d st sc s–st nd nd nd st s nd nd st–s
Growth at:
10°C − − + + − − − − − − − + − + − − + − − nd − + + nd
37°C w + − − + + + + + + + − + − + + + v + + + + − +
42°C − + − − nd nd nd − + nd + − nd − nd nd nd − + nd + + − nd
NaCl tolerance (%, w/v) 10 5 18 15 nd 15 15 15 12 10 10 10 nd 12 nd nd 18 15 10 nd 18 9 10 10
Caseinase + nd + + + + − − + + nd + + + + − + nd nd nd nd − nd +
Lipase − − nd nd − nd nd nd − + w − nd − − − nd − nd + nd − nd nd
Oxidase w − + + + − + − + nd − − nd − nd nd + + − nd − − nd −
Pyrazinamidase + + + + nd + − + + + w − − + + − + + nd + + + nd +
-Glucosidase − − − − − v + − − − − + − − − − − − nd − − + nd +
Nitrate reduction + − − + + v − v + − − + − − − − − v + − − − nd v
Acid from phenylacetate + nd nd nd nd + nd nd + nd nd nd + + − − nd nd nd nd nd nd nd +
Utilization of:
D -Arabinose − nd nd − − + − + − − − nd − − − − − nd nd − − nd nd +
Gluconate − nd + − − + − + − − nd nd − + − − + nd nd − nd nd nd +
Mannitol − + + + + − − + − − − + − − − − + nd − − − + − −
a Symbols: +, positive; −, negative; w, weak; v, variable; nd, not determined.

b bry; bright yellow; g, grayish; gw, grayish-white; light red; o, orange; po, pale-orange; y, wb, whitish-beige; wh, white, wo, white to orange;
wy, whitish-yellow; yellow; yo, yellow to orange.

c s,
smooth; st, sticky; d, dry; sc, smooth-creamy.
Brevibacterium album arylamidase, esterase (C4), esterase lipase (C8), leucine

Tang, Wang, Schumann, Stackebrandt, Lou, Jiang, Xu arylamidase, naphthol-AS-BI-phosphohydrolase, trypsin, and
and Li 2008, 576VP valine arylamidase (API ZYM tests). Additional phenotypic
................................................................................... characteristics are shown in Table 1.
al’bum. L. neut. adj. album white. Cell wall contains galactose. Major fatty acids are anteiso
Aerobic, Gram-stain-positive, motile actinomycete which C15:0 , C17:0 anteiso, C15:0 iso, and C16:0 iso. The predominant
forms rods (1.8 × 4.0–5.0 μm). Colonies are white, smooth, isoprenologue is MK-8(H2 ); also contains MK-6, MK-6(H2 ),
circular, opaque, and 1–2 mm in diameter after incubation MK-7(H2 ), MK-8, and MK-9(H2 ). The major phospholipids
for 2 d on gelatin-asparagine agar (ISP medium 5) supple- are diphosphatidylgylcerol and phosphatidylglycerol.
mented with 5% (w/v) NaCl. Growth occurs at 28–45∘ C with Source: a saline soil collected in northwest China.
an optimum of 37∘ C; pH range 6–8 with an optimum of 7.5. DNA G+C content (mol%): 70.0 (HPLC).
Grows in the presence of NaCl, 0–10% (w/v) with an opti- Type strain: YIM 90718, CCTCC AB 20612, DSM 18261,
mum of 0–5%. JCM 15617, KCTC 19173.
The following carbon sources are used for growth Sequence accession no. (16S rRNA gene): EF158852.
in Biolog GP2 microplates: D-alanine, β-cyclodextrin,
Brevibacterium antiquum
N-acetyl-D-glucosamine, fructose 6-phosphate, L-fucose,
Gavrish, Krauzova, Potekhina, Karasev, Plotnikova,
D-gluconic acid, myo-inositol, lactulose, D-mannitol, L-mannose,
Altyntseva, Korosteleva and Evtushenko 2005, 1VP
methyl β-D-galactoside, methyl α-D-glucoside, D-malic acid, (Effective publication: Gavrish, Krauzova, Potekhina,
palatinose, putrescine, salicin, succinic acid, trehalose, tura- Karasev, Plotnikova, Altyntseva, Korosteleva and
nose, L-alanyl glycine, Tween 80, and uridine 5′ -monopho- Evtushenko 2004, 181.)
sphate. ...................................................................................
Positive for alkaline phosphatase, gelatin hydrolysis, pyraz- an.ti’qu.um. L.neut. adj. antiquum ancient, old.
inamidase, pyrrolidonyl arylamidase, and acid production Aerobic, Gram-stain-positive, nonmotile actinomycete
from D-ribose (API CORYNE tests). In addition, it is positive which is composed mainly of coccoid cells in old cultures.
for acid and alkaline phosphatase, α-chemotrypsin, cystine Colonies are white to orange, circular, and slightly convex.
......................................................................................................................................................................................................
Orange pigment is produced in the light. Grows at 7∘ C, but extract, but adonitol, L- and D-arabinose, D-arabitol, dulci-
not at 37∘ C; optimal growth is between 24–26∘ C. Grows in tol, meso-erythritol, L-fucose, inositol, lactose, D-melibiose,
the presence of 18% (w/v) NaCl. raffinose, salicin, sorbitol, and trehalose are not. Acid is pro-
Gelatin is hydrolyzed, but esculin and starch are not. duced from galactose, D-glucose, mannose, and salicin, but
Oxidase-positive. The ability to hydrolyze urea and to pro- not from inositol, inulin, D-mannitol, melezitose, raffinose,
duce H2 S is strain-specific. L-Arabinose (slowly), D-fructose, sorbose, sorbitol, or trehalose. Alkaline phosphatase-positive,
D-galactose (slowly), D-glucose, glycerol, D-mannose, and but negative for pyrrolidonyl arylamidase, β-glucuronidase,
D-xylose are used as carbon sources in a mineral medium sup- β-galactosidase, and N-acetyl-β-glucosaminidase (API Coryne
plemented with 0.05% (w/v) yeast extract, but D-melibiose, system). Additional phenotypic characteristics are shown in
melezitose, raffinose and sorbitol are not. Acid is produced Table 1.
from fructose, D-galactose, D-glucose, mannose, and sor- Major fatty acids are C15:0 anteiso and C17:0 anteiso, and
bitol, but not from inositol, inulin, D-mannitol, melezitose, the predominant menaquinone MK-8(H2 ). Contains glycerol
raffinose, sorbose, rhamnose, or trehalose. Pyrrolidonyl teichoic acids.
arylamidase-positive, but negative for β-galactosidase, Source: cheese.
β-glucuronidase, and N-acetyl-β-glucosaminidase (API Coryne DNA G+C content (mol%): 60.1–64.3 (Tm ).
tests). Methyl red and Voges–Proskauer tests are negative. Type strain: ATCC 9175, JCM 2590, LMG 22546, NCIMB
Additional phenotypic characteristics are shown in Table 1. 8546), VKM Ac-2111.
The major fatty acids are C15:0 anteiso and C17:0 anteiso, and Sequence accession no. (16S rRNA gene): X76566.
predominant menaquinone is MK-8(H2 ). Cell-wall teichoic Additional comments: the culture collection accession num-
acids contain glucose, glycerol, and mannitol. ber LMG 22546 was provided on request for validation (List
Source: 1.8–3-million-year-old ancient permafrost sedi- Editor, 2005).
ments located on the Kolyma Lowland, East Siberia, Russia.
DNA G+C content (mol%): 60.1–64.3 (Tm ). Brevibacterium avium
Type strain: JCM 13317, LMG 22206, UCM Ac-411, VKM Pascual and Collins 1999, 1529VP
...................................................................................
Ac-2118.
Sequence accession no. (16S rRNA gene): AY243344. a’vi.um. L. n. avis a bird; L. gen. pl. n. avium of birds.
Additional comments: the specific epithet antiquum is a Aerobic, Gram-stain-positive, non-acid-fast, nonmotile
“Latin adjective in the neuter gender” not a “modern noun actinomycetes which have a coryneform morphology.
adjective” (sic) as cited in the publication by Gavrish et al. Colonies are convex, with entire margins and are round,
(2005). The culture collection accession number LMG 22206 smooth, grayish-white and butyrous. Grows well at 22 and
was provided on request for validation (List Editor, 2005). 30∘ C and optimally at 37∘ C.
Tyrosine is decomposed and extracellular DNase pro-
Brevibacterium aurantiacum duced. Gelatin and xanthine are degraded, but not esculin,
Gavrish, Krauzova, Potekhina, Karasev, Plotnikova,
starch or urea. L-Arabinose, D-arabitol, D-fructose, galac-
Altyntseva, Korosteleva and Evtushenko 2005, 1VP
tose, D-glucose, glycerol, D-mannose, and mannitol are
(Effective publication: Gavrish, Krauzova, Potekhina,
Karasev, Plotnikova, Altyntseva, Korosteleva and used as sole carbon sources, but not adonitol, amygdalin,
Evtushenko 2004, 181.) L-arabitol, arbutin, cellobiose, dulcitol, erythritol, D-fucose,
................................................................................... L-fucose, β-gentiobiose, 2-ketogluconate, 5-ketogluconate,
au.ran.ti’a.cum. N.L. neut. adj. aurantiacum orange-colored. N-acetylglucosamine, methyl-α-D-glucoside, glycogen, inosi-
Aerobic, Gram-stain-positive, nonmotile actinomycetes tol, inulin, lactose, D-lyxose, maltose, methyl α-D-mannoside,
which are mainly present as coccoid cells in older cultures. melezitose, melibiose, D-raffinose, rhamnose, ribose, salicin,
Colonies are orange, circular, slightly convex, and glistening. sorbitol, L-sorbose, starch, sucrose, D-tagatose, trehalose,
Grows at 7∘ C and optimally between 24–26∘ C. Grows in the D-turanose, xylitol, D-xylose, or methyl β-xyloside. Does not
presence of 15% (w/v) NaCl. produce acid from glucose, maltose, mannitol, sucrose,
H2 S is produced. Does not hydrolyze esculin, starch, or or xylose. Positive for alkaline and acid phosphatase,
urea. Production of acid from fructose and gelatin hydrolysis esterase (C4), ester lipase (C8), leucine arylamidase,
are strain-specific. Cellobiose, fructose, D-galactose, D-glucose, and phosphoamidase, but negative for α-chymotrypsin,
glycerol, D-mannose, and D-xylose are used as carbon sources α-fucosidase, β-fucosidase, α-galactosidase, β-glucuronidase,
in a mineral medium supplemented with 0.05% (w/v) yeast α-mannosidase, N-acetyl-β-glucosaminidase, trypsin, and
......................................................................................................................................................................................................
valine arylamidase (API ZYM tests). Additional phenotypic Brevibacterium celere

characteristics are shown in Table 1. Ivanova, Christen, Alexeeva, Zhukova, Gorshkova,
Galactose, but not arabinose, is present in the cell wall. Lysenko, Mikhailov and Nicolau 2004, 2110VP
...................................................................................
Source: bumble-foot-like manifestations of poultry.
ce’le.re. L. neut. adj. celere rapid, indicating the rapid growth
DNA G+C content (mol%): not determined.
on nutrient media.
Type strain: CCUG 43455, CIP 106532, JCM 11680, NCIMB
Aerobic, Gram-stain-positive, non-acid-fast, nonmotile
703055. actinomycetes which have a coryneform morphology.
Sequence accession no. (16S rRNA gene): Y17962. Colonies are circular, convex with entire margins, whitish
Brevibacterium casei yellow with a butyrous consistency. Growth occurs between
Collins, Farrow, Goodfellow and Minnikin 1983b, 12–42∘ C, and from pH 5–10 with an optimum at 8.5–9.0.
896VP (Effective publication: Collins, Farrow, Tolerant to 15% (w/v) NaCl.
Goodfellow and Minnikin 1983a, 393.) Alginate, gelatin, and laminarin are degraded but
................................................................................... starch is not. Negative for urease production. The fol-
ca’se.i. L. n. caseus cheese; L. gen. n. casei of cheese. lowing carbon sources are used for growth in Biolog GP2
Aerobic, Gram-stain-positive, nonmotile actinomycetes microplates: acetic acid, cis-aconitic acid, alaninamide,
γ-aminobutyric acid, bromosuccinic acid, D-and L-alamine,
which have a coryneform morphology. Gray-white colonies
D-and L-carnitine, D-gluconic acid, α-D-glucose, D- and L-lactic
are produced on nutrient agar. A brown, water soluble
acid, D-trehalose, dextrin, formic acid, glycerol, glycyl
pigment is formed on milk agar. Grows well at 40∘ C, and opti-
L-glutamic acid, α-hydroxybutyric acid, β-hydroxybutyric acid,
mally between 30–37∘ C. Survives heating in nutrient broth γ-hydroxybutyric acid, p-hydroxyphenylacetic acid, hydrox-
(Oxoid) at 60∘ C for 30 min. Tolerant to 15% (w/v) NaCl. yproline, α-ketobutyric acid, L-alanyl glycine, L-asparagine,
Gelatin is hydrolyzed and extracellular DNase produced. L-aspartic acid, L-fucose, L-glutamic acid, L-histidine, L-leucine,
Reduction of nitrate is variable. Weak acid production from L-ornithine, L-phenylalanine, L-proline, L-pyroglutamic
glucose has been reported using the method of Hugh and acid, L-serine, maltose, methylpyruvate, monomethyl suc-
Leifson (1953). Glucose, sucrose, acetate, and lactate are used cinate, N-acetyl-D-galactosamine, N-acetyl-D-glucosamine,
as sole carbon sources. Additional phenotypic characteristics phenylethylamine, propionic acid, putrescine, quinic acid,
sebacic acid, succinic acid, sucrose, turanose, Tween 40,
are shown in Table 1.
Tween 80, and urocanic acid. Does not form acid from
DNA–DNA relatedness values between the type strain and
galactose, glucose, lactose, maltose, sorbose, or xylose.
representatives of related species are as follows: Brevibacterium
Susceptible to (μg/ml): carbenicillin (10) and olean-
epidermidis (22–23%), Brevibacterium iodinum (14% with type domycin (30), but resistant to: ampicillin (10), benzylpeni-
strain), and Brevibacterium linens (19% with type strain). cillin (10), gentamicin (10), kanamycin (30), lincomycin
Source: cheddar cheese, cheese curd, and raw milk. (15), neomycin (30), streptomycin (30), and polymyxin B
DNA G+C content (mol%): 66–67 (Tm ). (25). Additional phenotypic characteristics are shown in
Type strain: ATCC 35515, CCM 4100, BCRC 12214, CCUG Table 1.
29001, CIP 102111, DSM 20657, IMET 10997, JCM 2594, Major fatty acids are C15:0 anteiso and C17:0 anteiso.
Source: the brown alga Fucus evanescens.
KCTC 3082, KCTC 3085, NBRC 14812, NCIMB 702048, VKM
DNA G+C content (mol%): 61.4 (Tm ).
Ac-2114.
Type strain: ATCC BAA-809, DSM 15453, JCM 13521, KMM
Sequence accession no. (16S rRNA gene): AJ251418, X76564.
3637.
Additional comments: Brevibacterium casei Collins et al., Sequence accession no. (16S rRNA gene): AY228463.
(1983a) was previously known as CDC coryneform groups
B-1 and B-3. The name Brevibacterium casei (effective publi- Brevibacterium epidermidis
Collins, Farrow, Goodfellow and Minnikin 1983b,
cation: Collins et al., 1983a), type strain CMD1 = NCIMB
896VP (Effective publication: Collins, Farrow,
702048 (formerly NCDO 2048) appears on validation lists 12 Goodfellow and Minnikin 1983a, 393.)
and 13. The date of valid publication of the name is the first ...................................................................................
citation, namely the one in validation list 12 (Euzéby and e.pi.der’mi.dis. Gr. n. epidermida the outer skin; N.L. gen. n.
Kudo, 2001). epidermidis of the epidermis.
......................................................................................................................................................................................................
Aerobic, Gram-stain-positive, nonmotile actinomycetes Major fatty acids are C15:0 anteiso and C17:0 anteiso.
which have a coryneform morphology. Gray-white colonies Source: milk.
are formed on nutrient agar. Grows optimally between DNA G+C content (mol%): 60–63 (Tm ).
30–37∘ C; some strains grow at 40∘ C. Grows in presence of Type strain: ATCC 49514, BCRC 12216, DSM 20626, IMET
15% (w/v) NaCl. 10995, JCM 2591, KCTC 3083, NBRC 15230, NCIMB 700613,
Gelatin is hydrolyzed and extracellular DNase is pro- NCTC 12955, NRRL B-1717, VKM Ac-2106.
duced. Nitrate reduction is strain-dependent. Acetate and
Sequence accession no. (16S rRNA gene): X76567, X83813.
lactate are used as sole carbon sources. Acid is not produced
from glucose in peptone media. Additional phenotypic Brevibacterium luteolum corrig.
characteristics are shown in Table 1. Wauters, Avesani, Laffineur, Charlier, Janssens, Van
DNA–DNA relatedness values between the type strain and Bosterhaut and Delmée 2003, 1324VP
...................................................................................
representatives of related species are as follows: Brevibacterium
casei (31–44%), Brevibacterium iodinum (33% with type strain), lu.te’o.lum. L. neut. adj. luteolum yellowish, because colonies
and Brevibacterium linens (14% with type strain). exhibit a yellow pigment.
Source: human skin. Aerobic, Gram-stain-positive rods (2–3 μm long) which
DNA G+C content (mol%): 63.5 (Tm ). show a diphtheroid morphology, but not a rod–coccus
Type strain: ATCC 35514, BCRC 2215, CCM 4098, CCUG growth cycle. Colonies are smooth, yellowish, and approxi-
30540, CIP 102110, DSM 20660, IMET 10998, JCM 2593, mately 1–2 mm in diameter after incubation for 2 d at 37∘ C.
KCTC 3090, NBRC 14811, NCIMB 702286, NCTC 12997, Grows optimally between 30 and 37∘ C; some strains grow at
VKM Ac-2108. 20∘ C. Grows in the presence of 10% (w/v) NaCl.
Sequence accession no. (16S rRNA gene): X76565.
Gelatin is degraded, but esculin, tyrosine, and xanthine
Additional comments: the species name Brevibacterium epider-
are not. Ethylene glycol is acidified. γ-Aminobutyric acid is uti-
midis [effective publication: Collins et al., 1983a; type strain:
lized and alkalinized on Simmon’s agar base. Phenylacetate
strain D731 = NCIMB 702286 formerly NCDO 2286] appears
is acidified and methanethiol produced from L-methionine.
on Validation Lists 12 and 13. The date of the valid publica-
Pyrrolidonyl peptidase-positive. N-Acetyl-β-D-glucosaminidase
tion of this name is the first citation, namely the one cited in
Validation List 12 (Euzéby and Kudo, 2001). is positive when the nitrophenyl compound is used as sub-
strate (Rosco). Positive for lipase esterase (C8), leucine
Brevibacterium iodinum arylamidase, and phosphoamidase, and acid and alkaline
(ex Davis 1939) Collins, Jones, Keddie and Sneath
phosphatase variable (negative for the type strain) (API ZYM
1981, 216VP (Effective publication Collins, Jones,
tests). Additional phenotypic characteristics are shown in
Keddie and Sneath 1980, 7.)
................................................................................... Table 1.
i.o.di’num. N.L. neut. n. iodinum iodine. Main fatty acids are C15:0 anteiso and C17:0 anteiso. Isolated
Aerobic, nonmotile actinomycetes which have a rod– from clinical and environmental samples.
coccus growth cycle. Cells frequently appear Gram-stain- Source: peritoneal fluid of a patient undergoing dialysis.
negative. In very young cultures (approx. 8 h), cells frequently DNA G+C content (mol%): 68.8 (Tm ).
stain one-half Gram-positive, the other half Gram-negative. Type strain: CF87, CCUG 46604, DSM 15022, JCM 12367.
Colonies on nutrient agar are whitish-gray and smooth, Sequence accession no. (16S rRNA gene): AJ488509.
convex with entire margin, and shiny. Blue to purple extra- Additional comments: Wauters et al. (2003) proposed the
cellular crystals of iodinin are produced on a variety of media. specific epithet lutescens. However, lutescens (L. part. adj. from
Iodinin production is not influenced by light. Grows between L. v. lutesco) means becoming muddy, not yellowish as cited in
20–37∘ C, optimally between 25–30∘ C, but does not grow the paper by Wauters et al. Consequently, this specific epithet
below 12∘ C.
was corrected on notification with the author’s agreement
Catalase-positive. Gelatin is hydrolyzed. DNase is pro-
(Euzéby, 2004).
duced. Hippurate is not hydrolyzed. H2 S is produced from
cysteine but not from sodium thiosulfate. Does not produce Brevibacterium marinum
acid from glucose or other sugars in peptone based media. Lee 2008, 503VP
Citrate is utilized. Additional phenotypic characteristics are ...................................................................................
shown in Table 1. ma.ri’num. L. neut. adj. marinum of the sea, marine.
......................................................................................................................................................................................................
Aerobic, Gram-stain-positive, nonmotile actinomycete between 30–37∘ C. Good growth in the presence of 1–10%
which forms long rods (1.5–2.4 × 0.6–0.9 μm) at the expo- (w/v) NaCl.
nential growth phase; as the cultures age these elements Does not hydrolyze gelatin. Positive for esterase (C4) and
fragment into short rods (0.5–0.6 × 0.7–1.0 μm) which occa- leucine arylamidase, but negative for alkaline phosphatase,
sionally occur singly but more often, in pairs or in chains. α-fucosidase, β-glucosidase, β-glucuronidase, α-mannosidase,
V-forms are observed. Colonies are translucent, convex, N-acetyl-β-glucosaminidase, and naphthol-
smooth, and circular with entire margins. Pigment produc- AS-BI-phosphohydrolase (API ZYM tests). Positive for ala-
tion is variable depending on the culture conditions. In the nine arylamidase, arginine arylamidase, glycine arylamidase,
dark colonies are white; in the light, they are bright yellow. leucine arylamidase, leucyl-glycine arylamidase, phenylala-
Grows from 10–30∘ C, optimally at 30∘ C; does not grow at 4 nine arylamidase, proline arylamidase, serine arylamidase,
or 37∘ C. Growth occurs in the pH range 5–12. Tolerant to and tyrosine arylamidase (API 32A tests).
10% (w/v) NaCl. The following sole carbon sources are used for growth
Hypoxanthine is degraded, but not DNA, gelatin, in Biolog GP2 microplates: acetic acid, 2,3-butanediol,
starch, cellulose, tyrosine, or xanthine. Positive for nitrate D-alanine, glycyl L-glutamic acid, α-hydroxybutyric acid,
reduction, β-galactosidase, and urease, but negative for β-hydroxybutyric acid, γ-hydroxybutyric acid, α-ketovaleric
alkaline phosphatase, β-glucosidase, β-glucuronidase, acid, L-alaninamide, L-alanine, L-alanyl glycine, L-aspara-
N-acetyl-β-glucosaminidase, and pyrrolidonyl arylmidase gine, L-glutamic acid, L-lactic acid, L-serine, N-acetyl-L--
(API Coryne tests). Esterase (C4) and esterase lipase glutamic acid, propionic acid, pyruvic acid, pyruvic acid
(C8) are weakly positive, but acid phosphatase, alkaline methyl ester, succinic acid monomethyl ester, Tween 40, and
phosphatase, α-chemotrypsin, cysteine arylamidase, galac- Tween 80. Additional phenotypic characteristics are shown in
Table 1. The major fatty acids are C15:0 anteiso, C17:0 anteiso,
tosidase, β-glucosidase, leucine arylamidase, lipase (C14),
C15:0 iso, and C16:0 iso.
α-mannosidase, naphthol-AS-BI-phosphohydrolase, trypsin,
Source: a human ankle discharge sample.
and valine arylamidase are negative (API ZYM tests). Does not
DNA G+C content (mol%): not determined.
produce acid from D-glucose, D-lactose, maltose, D-mannitol,
Type strain: 5401308, CCUG 53855, CIP 109422, CSUR P26.
D-ribose, sucrose, or D-xylose. Additional phenotypic char-
Sequence accession no. (16S rRNA gene): EU868814.
acteristics are shown in Table 1. The major fatty acids are
C15:0 anteiso, C17:0 anteiso, and C18:0 . The predominant Brevibacterium mcbrellneri
menaquinone is MK-8(H2 ), and the major polar lipids McBride, Ellner, Black, Clarridge and Wolf 1994,
are diphosphatidylglycerol, phosphatidylglycerol, and phos- 852VP (Effective publication: McBride, Ellner, Black,
phatidyl methyl inositol; an unknown phospholipid is present Clarridge and Wolf 1993, 260.)
...................................................................................
as a minor component.
Source: seawater from Hwasun Beach in Jeju, Republic of mc.brell’ne.ri. N.L. masc. gen. n. mcbrellneri of McBrEllner,
named for two of the authors (M.E. McBride and K.M. Ellner)
Korea.
of the publication describing its isolation and characteristics.
DNA G+C content (mol%): 71.4 (HPLC).
Type strain: HFW-26, DSM 18964, JBRI 2001, JCM 15618,
which show a rod–coccus growth cycle. Colonies are
KCTC 19221.
whitish-beige, dry, and friable. Grows at 37∘ C, but not at 20∘ C.
Sequence accession no. (16S rRNA gene): AM421807.
Tolerates 6.5% (w/v) NaCl but not higher concentrations
Brevibacterium massiliense of salt.
Roux and Raoult 2009, 1962VP DNA and gelatin are hydrolyzed. Xanthine degrada-
................................................................................... tion is variable. Pyrrolidone peptidase is negative. Produces
mas.si.li.en’se. L. neut. adj. massiliense of Massilia, the old methanethiol from L-methionine. Ribose is used as a sole
Roman name for Marseille, where the type strain was isolated. carbon source, but mannose is not. Acid is produced from
Aerobic, Gram-stain-positive, nonmotile actinomycetes 2,3-butylene glycol.
which form short, irregular straight rods (0.4–1.4 × 0.3–- Sensitive (μg/ml) to ampicillin (10), cephalothin, metho-
0.5 μm). Colonies are beige, shiny, smooth, circular, and prim sulfate (24), penicillin (2), and vancomycin (15), but
0.2 mm in diameter after growth for 24 h in 5% CO2 resistant to erythromycin (15), oxacillin (10), and tetracy-
on sheep blood agar. Grows between 25–45∘ C, optimally cline (30). Additional phenotypic characteristics are shown
......................................................................................................................................................................................................
in Table 1. The major fatty acids are C15:0 antieso and C17:0 Brevibacterium otitidis
anteiso. Cell walls contain galactose, glucose, and xylose. Pascual, Collins, Funke and Pitcher 1996b, 1189VP
Source: infected genital hair of patients with white piedra. (Effective publication: Pascual, Collins, Funke and
Pitcher 1996a, 121.)
DNA G+C content (mol%): 63.05 (Tm ). ...................................................................................
Type strain: E2cr, ATCC 49030, CCUG 37313, DSM 9583,
o.ti’ti.dis. Gr. n. ous, otos ear; N.L. suff. -itis, idis suffix used in
JCM 11682. names of inflammations; N.L. gen. n. otitidis of inflammation
Sequence accession no. (16S rRNA gene): X93594. of the ear.
Brevibacterium oceani
which have a coryneform morphology. Colonies are
Bhadra, Raghukumar, Pindi and Shivaji 2008, 59VP
................................................................................... whitish-yellowish and friable with a dry texture. Growth
is observed at 37∘ C but not at 20∘ C.
o.ce.a’ni. L. gen n. oceani of the ocean.
Gelatin and xanthine are degraded. Positive for produc-
Aerobic, Gram-stain-positive, nonmotile, rod-shaped
tion of pyrrolidone peptidase. Additional phenotypic charac-
(2.0–3.0 × 1.0–1.2 μm) actinomycetes. Sticky, pale orange teristics are shown in Table 1.
colonies (∼1.5–2.0 mm in diameter) with entire margins are Source: a human ear infection.
formed at 28∘ C on nutrient agar. Growth occurs between DNA G+C content (mol%): not determined.
10–35∘ C, but not at 5 or 37∘ C. pH for growth is between Type strain: A37/73, ATCC 700348, BBH7, CCUG 37257,
5.2–9.5 with an optimum of 6.8. CIP 10544, DSM 10718, JCM 11683, NCIMB 703053.
Sequence accession no. (16S rRNA gene): X93593.
L-Arginine, L-asparagine, L-lysine, malonate, L-proline,
L-serine, and L-tyrosine are used as carbon sources, but Brevibacterium paucivorans
adonitol, L-cystine, erythritol, D-glucuronate, and 2-oxogluta- Wauters, Charlier and Janssens Delmée 2001, 1706VP
...................................................................................
rate are not. Acid is produced from phenylacetate, but not
from adonitol, D- and L-arabinose, D-cellobiose, dulcitol, pau.ci.vo’rans. L. adj. paucus little; L. v. vorare to eat; N.L. part.
adj. paucivorans eating little, because few substrates are assim-
fructose, D-galactose, glucose, inositol, lactose, maltose,
ilated.
D-mannitol, mannose, D-melibiose, D-raffinose, rham-
nose, D-ribose, sucrose, trehalose, or D-xylose. Positive which form short, plump rods with a coryneform morphol-
for lysine decarboxylase and ornithine decarboxylase, ogy. Colonies are grayish, glistening and smooth or sticky
but negative for citrate utilization, esculin hydrolysis, and approximately 2 mm in diameter after incubation for 2
β-galactosidase, H2 S production, indole production, methyl d at 37∘ C on blood agar. Grows at 37∘ C, but not 20∘ C.
red and Voges–Proskauer tests, oxidase, and phenylalanine Carbohydrates are neither acidified nor assimilated.
deamination. Gelatin is liquefied weakly and slowly. Does not hydrolyze
esculin, or degrade tyrosine or xanthine. Ethylene glycol
Resistant to (μg/ml): chloramphenicol (25), nalidixic
is acidified but 2,3-butylene glycol is not. Methanethiol is
acid (15), and polymixin B (15), but sensitive to ampicillin
produced from L-methionine. Urea is hydrolyzed. Pyrroli-
(25), kanamycin (30), roxithromycin (30), streptomycin donyl peptidase is negative. Positive for leucine arylamidase
(25), and tetracycline (25). Additional phenotypic charac- and phosphoamidase; weakly positive for esterase lipase
teristics are shown in Table 1. The major fatty acids are C15:0 (C8), but negative for acid phosphatase, alkaline phos-
anteiso and C17:0 anteiso. The predominant menaquinone phatase, α-chymotrypsin, cystine arylamidase, esterase
is MK-8(H2 ), and the major polar lipids are diphosphatidyl- (C4), α-fucosidase, α- and β-glucosidase, α-mannosidase,
glycerol and phosphatidylglycerol. N-acetyl-β-glucosaminidase, and valine arylamidase; some
strains are positive for trypsin (API ZYM tests). Additional
Source: a 50–70 cm section of a deep-sea sediment core
phenotypic characteristics are shown in Table 1.
of 4.6 m length obtained from the Chagos Trench, Indian
The cell wall contains an excess of glutamic acid and
Ocean, at a water depth of 5904 m.
glycine moieties that are not bound to the peptidoglycan.
DNA G+C content (mol%): 59.8–60.2 (Tm ). The cellular fatty acids are of the branched type with C15:0
Type strain: BBH7, IAM 15353, JCM 21798, LMG 23457. anteiso and C17:0 anteiso as major components. Isolated from
Sequence accession no. (16S rRNA gene): AM158906. clinical material.
......................................................................................................................................................................................................
Source: blood. Brevibacterium picturae

DNA G+C content (mol%): 55.8 (Tm ). Heyrman, Verbeeren, Schumann, Devos, Swings and
Type strain: CF62, CCUG 45688, DSM 13657, JCM 11567, De Vos 2004, 1540VP
...................................................................................
LMG 19814.
pic.tu’ra.e. L. gen. n. picturae of a painting.
Sequence accession no. (16S rRNA gene): AJ251463.
Brevibacterium permense which do not form a clear rod–coccus cycle on nutrient agar.
Gavrish, Krauzova, Potekhina, Karasev, Plotnikova, Single coccoid cells (0.8 × 1.0 μm) are formed after growth
Altyntseva, Korosteleva and Evtushenko 2005, 1VP for 6 and 16 h, single coccoid to oval cells after 24 h; after 72
(Effective publication: Gavrish, Krauzova, Potekhina, h growth, cells are oval or short rods which occur singly or in
Karasev, Plotnikova, Altyntseva, Korosteleva and
pairs (1.0–1.5 × 4.0 μm). Colonies on nutrient agar are white,
Evtush enko 2004, 182.)
................................................................................... low-convex, slightly transparent to opaque, and circular with
entire margins. Growth is variable at 37∘ C, optimal between
per.men’se. N.L. neut. adj. permense pertaining to Perm, a
20–30∘ C, but does not grow at 10∘ C. The pH range for
region of Russia.
growth is 6–9, with an optimum at pH 7.0. Tolerant to 10%
(w/v) NaCl.
which are mainly present as coccoid cells in older cultures. Positive for alkaline phosphatase, urease, and gelatin
Colonies are white to orange, circular, slightly convex, and hydrolysis, and negative for pyrrolidonyl arylamidase,
glistening. Orange pigment is produced in the light. Grows β-glucuronidase, esculin, and fermentation of ribose, xylose,
at 37∘ C; optimal temperature for growth is 24∘ C. Grows in mannitol, and glycogen; variable for nitrate reduction,
the presence of 18% (w/v) NaCl. β-galactosidase, and acid production from glucose, lactose,
Gelatin is hydrolyzed but esculin and urea are not. maltose, and sucrose (API Coryne tests).
H2 S is produced. The methyl red and Voges–Proskauer Weakly positive for esterase (C4), esterase lipase (C8),
tests are negative. Does not degrade starch. L-Arabinose, and naphthol-AS-BI-phosphohydrolase; variable for acid
cellobiose, fructose, D-galactose, D-glucose, gluconate, glyc- phosphatase and leucine arylamidase, and negative for
erol, D-mannose, raffinose, salicin, trehalose, and D-xylose α-chymotrypsin, cystine arylamidase, α-fucosidase, α-galacto-
are used as carbon sources when prepared in a mineral sidase, β-glucosidase, α-mannosidase, trypsin, and valine
medium supplemented with 0.05% (w/v) yeast extract, but arylamidase (API ZYM tests). Additional phenotypic charac-
teristics are shown in Table 1. The major fatty acids are C15:0
D-arabinose, D-arabitol, adonitol, dulcitol, meso-erythritol,
anteiso and C17:0 anteiso. The predominant menaquinone
L-fucose, inositol, lactose, D-melibiose, and sorbitol are not.
is MK-8(H2 ), and the major polar lipids are diphosphatidyl-
Acid is produced from D-galactose, D-glucose, fructose, man-
glycerol, phosphatidylglycerol, phosphatidylinositol, an
nose, salicin, and sorbitol, but not from inositol, inulin,
unknown glycolipid, and an unknown phospholipid.
D-mannitol, melezitose, raffinose, rhamnose, sorbose, or
Source: a mural painting discolored by microbial growth.
trehalose. Positive for alkaline phosphatase, but negative for DNA G+C content (mol%): 63.3 (HPLC).
pyrrolidonyl arylmidase, β-glucuronidase, β-galactosidase, Type strain: DSM 16132, JCM 13319, LMG 22061.
and N-acetyl-β-glucosaminidase (API Coryne tests). Addi- Sequence accession no. (16S rRNA gene): AJ620364.
tional phenotypic characteristics are shown in Table 1. The
major menaquinone is MK-8(H2 ). Cell-wall teichoic acid
Brevibacterium pityocampae
Kati, İnce, Demir and Demirbağ 2010, 315VP
contains glycerol, mannose, and mannitol. ...................................................................................
Source: salt-contaminated soil samples collected in the
pi.ty.o.cam’pae. L. fem. n. pityocampa a pine grub; L. fem. gen.
Perm region of Russia.
n. pityocampae of a pine grub, referring to the isolation of the
DNA G+C content (mol%): 60.1–64.3 (Tm ). type strain from larvae of Thaumetopoea pityocampa.
Type strain: JCM 13318, LMG 22207, UMC Ac-413, VKM Aerobic, Gram-stain-positive, nonmotile actinomycete
Ac-2280. which forms rods (1.0 × 0.5 μm). Colonies are yellow, circu-
Sequence accession no. (16S rRNA gene): AY243343. lar, and convex with entire margins. Grows at 25–40∘ C, from
Additional comments: the culture collection number was pH 6.0–9.0, and optimally at pH 8.0. Grows in 0–10%, but
provided on request for validation (List Editor, 2005). not 15% (w/v) NaCl.
......................................................................................................................................................................................................
Arginine dihydrolase, β-galactosidase, lysine decarboxy- Brevibacterium salitolerans

lase, ornithine decarboxylase, and tryptophan deaminase are Guan, Zhau, Xiao, Liu, Xia, Zhang and Zhang 2010,
2993VP
not formed. Does not produce acetoin, hydrogen sulfide, or
...................................................................................
indole; gelatin is not degraded and citrate is not utilized. Does
sa.li.to.le’rans. L. n. sal salis salt; L. part. adj. tolerans tolerating;
not produce acid from amygdalin, L-arabinose, D-glucose,
N.L. part. adj. salitolerans salt-tolerating, originating from a
inositol, D-mannitol, melibiose, L-rhamnose, D-sorbitol, or saline habitat.
sucrose. Additional phenotypic characteristics are shown Aerobic, Gram-stain-positive, nonmotile actinomycete
in Table 1. The major fatty acids are C15:0 anteiso and C17:0 which forms rods (1.3–1.8 × 0.6–1.0 μm) but does not show
anteiso. a rod–coccus growth cycle. Colonies on glycerol-asparagine
agar (ISP 5 medium) supplemented with 2% (w/v) NaCl, 1%
Source: healthy larvae of Thaumetopoea pityocampa from the
(w/v) KCl, and 0.5% (w/v) MgCl2 are 2.0–3.5 mm in diam-
Middle Black Sea region of Turkey.
eter, smooth, circular, and whitish-yellow after 5 d. Colonial
DNA G+C content (mol%): 69.8 (HPLC). pigmentation is always whitish-yellow in both the dark and
Type strain: Tp12, DSM 21720, NCCB 100255. light. Grows from 15–50∘ C, optimally between 28 and 40∘ C,
Sequence accession no. (16S rRNA gene): EU484189. but does not grow at 10∘ C or 52∘ C. Grows from pH 5.0–9.0,
optimally at pH 7.0–7.5. Tolerant up to 18% (w/v) NaCl, but
Brevibacterium ravenspurgense grows optimally in the presence of 3.8% (w/v) NaCl.
Mages, Frodl, Bernard and Funke 2009, 1VP (Effective
Degrades gelatin, but not cellulose, starch, or Tweens 20,
publication: Mages, Frodl, Bernard and Funke 2008,
40, or 80. Positive for alkaline phosphatase, gelatin hydrolysis,
2985.)
................................................................................... β-glucosidase, pyrazinamidase, pyrrolidonyl arylamidase, and
acid production from D-ribose (API CORYNE tests). Does not
ra.vens.pur.gen’se. N.L. adj. ravenspurgense pertaining to
coagulate or peptonize milk or produce H2 S. Methyl red and
Ravenspurgum, Latin name of the town of Ravensburg, Voges–Proskauer tests are negative.
Germany, where the type strain of this species was isolated. The following sole carbon sources are used for growth
Aerobic, Gram-stain-positive, nonmotile actinomycetes in Biolog GP2 microplates: acetic acid, N-acetyl-L-glutamic
which show a coryneform morphology. Colonies are acid, adenosine, L-alaninamide, D-alanine, L-alanine, L-alanyl
whitish-grayish, slightly convex, have a sticky consistency, glycine, L-asparagine, 2,3-butanediol, α-cyclodextr-
in, 2′ -deoxy adenosine, dextrin, L-fucose, D-gluconic acid,
and are up to 2 mm in diameter after incubation for 24 h at
L-glutamic acid, glycogen, glycyl L-glutamic acid, α-, β- and
35∘ C on Columbia SBA plates.
γ-hydroxybutyric acid, inosine, α-ketovaleric acid, lactamide,
Positive for esterase (C4), esterase lipase (C8), leucine L-lactic acid, D-lactic acid methyl ester, D-malic acid, L-malic
arylamidase, and naphthol-AS-BI-phosphohydrolase, but acid, methyl pyruvate, monomethyl succinate, propionic
negative for acid phosphatase, α-chymotrypsin, α-fucosidase, acid, pyruvic acid, L-serine, succinamic acid, succinic acid,
α-galactosidase, β-galactosidase, gelatinase, N-acetyl-β- thymidine, Tween 40, and Tween 80. Additional phenotypic
glucosaminidase, α-glucosidase, β-glucosidase, β-glucuronidase, characteristics are shown in Table 1.
The DNA–DNA relatedness value between the type strain
lipase (C14), α-mannosidase, nitrate reductase, pyrrolidonyl
and the corresponding strain of Brevibacterium album is 41.3%.
arylamidase, urease, and valine arylamidase; variable for
The major fatty acids are C17:0 anteiso and C15:0 anteiso.
alkaline phosphatase and trypsin (API ZYM tests). Does not The predominant menaquinone is MK-8(H ), but smaller
2
use carbohydrates in the system described by Funke and amounts of MK-6(H2 ), MK-7, MK-7(H2 ), and MK-8 are also
Carlotti (1994). Additional phenotypic characteristics are present. The major polar lipids are diphosphatidylglycerol
shown in Table 1. The major fatty acids are C15:0 anteiso and and phosphatidylglycerol.
Source: a hypersaline sediment sample collected from Lop
C17:0 .
Nur salt lake in Xinjiang Province, north-west China.
Source: the town of Ravensburg in Germany.
DNA G+C content (mol%): 69 (HPLC).
DNA G+C content (mol%): not determined. Type strain: TRM 415, CCTCC AB 208328, JCM 15900,
Type strain: CCUG 56047, DSM 21258. KCTC 19616.
Sequence accession no. (16S rRNA gene): EU086793. Sequence accession no. (16S rRNA gene): GU117109.
......................................................................................................................................................................................................
Brevibacterium samyangense in Table 1. The major fatty acids are C17:0 anteiso, C15:0
Lee 2006, 1891VP anteiso, and C15:0 iso. The predominant menaquinone is
................................................................................... MK8(H2 ). The polar lipid pattern contains phosphatidylglyc-
sam.yang.en’se. N.L. neut. adj. samyangense of Samyang erol as a major component but lacks diphosphatidylglycerol
Beach, Jeju, Republic of Korea, from where the type strain and phosphatidylinositol.
was isolated. Source: sand sediment of Samyang Beach, Jeju Island,
Aerobic, Gram-stain-positive, motile actinomycetes Republic of Korea.
which show an apparent rod–coccus growth cycle on yeast DNA G+C content (mol%): 70.7 (Tm ).
extract-sea water agar; single or paired coccoid cells are seen Type strain: SST-8, JCM 14546, KCCM 42316, NRRL
after 6 h together with rare short rods; after 12–16 h most B-41420.
cells are irregular, slender rods, arranged in V-forms, while Sequence accession no. (16S rRNA gene): DQ344485.
coccoid cells occur singly, in pairs, or in chains after 24–72
h. Colonies are opaque, convex, circular, and bright yellow Brevibacterium sandarakinum
Kämpfer, Schäfer, Lodders and Busse 2010, 911VP
in color with an entire edge. Grows from 10–45∘ C, but not
...................................................................................
at 4 or 55∘ C, from pH 6.1–10.1, and in the presence of 15%
san.da.ra.ki’num. N.L. neut. adj. sandarakinum (from Gr.
(w/v) NaCl.
neut. adj. sandarakinos -e -on) of light-red color.
Degrades D- and L-tyrosine, but not xanthine. The follow-
Aerobic, Gram-stain-positive, nonmotile actinomycete
ing sole carbon sources are used for growth in Biolog
GP2 microplates: acetic acid, N-acetyl-L-glutamic acid, which does not form a clear rod–coccus cycle on nutrient
N-acetyl-D-glucosamine, N-acetyl-β-mannosamine, adeno- agar. After 12 h growth, cells are coccoid (0.8–1.2 μm) and
sine, adenosine 5′ -monophosphate, L-alaninamide, D- and occur singly; after 24 h cells are coccoid to oval. Good growth
L-alanine, L-alanylglycine, amygdalin, L-arabinose, arabitol,
occurs after 3 d of incubation on nutrient agar at 25–30∘ C.
L-asparagine, D-cellobiose, β-cyclodextrin, 2-deoxyadenos-
Growth occurs between 4∘ C (weak) and 36∘ C (but not above
ine, dextrin, D-fructose, L-fucose, D-galactose, D-galacturonic this temperature), from pH 5.5–12.5 (optimally from pH
acid, α-D-glucose, D-glucose 6-phosphate, α-D-glucoside 7.5–9.5), and in the presence of 1–10% NaCl.
1-phosphate, L-glutamic acid, DL-α-glycerol phosphate, glyco- Acetate (weak), cis-aconitate, citrate, fumarate (weak),
D-galactose, D-glucose, glutarate, histidine, DL-3-hydroxybu-
gen, glycyl L-glutamic acid, p-hydroxyphenylacetic acid,
α-, β-, and γ-hydroxybutyric acids, inosine, myo-inositol, tyrate, D-mannose, 2-oxoglutarate, propionate, pyruvate,
inulin, α-ketoglutaric acid, lactamide, L-lactic acid, D-lactic ribose, and D-sorbitol are used as sole carbon sources,
acid methyl ester, α-D-lactose, lactulose, L-malic acid, but N-acetyl-D-galactosamine, N-acetyl-D-glucosamine, trans-
maltose, maltotriose, mannan, D-mannose, D-melezitose, aconitate, D-adonitol, 4-aminobutyrate, L-arabinose, arbutin,
D-melibiose, methyl ethyl α-D-mannoside, methyl α-D-galact- azelate, cellobiose, D-fructose, myo-inositol, itaconate,
oside, 3-methyl D-glucoside, methyl β-D-glucoside, methyl D-maltitol, maltose, D-mannitol, melibiose, mesaconate,
pyruvate, monomethyl succinate, palatinose, propionic 2-oxoglutarate, putrescine, L-rhamnose, salicin, sucrose,
acid, D-psicose, putrescine, L-pyroglutamic acid, D-raffinose, trehalose, and D-xylose are not. Additional phenotypic
L-rhamnose, D-ribose, salicin, sedoheptulosan, L-serine, characteristics are shown in Table 1.
D-sorbitol, stachyose, succinamic acid, succinic acid, sucrose, Major fatty acids are anteiso-branched fatty acids.
D-tagatose, thymidine, thymidine 5′ -monophosphate, Menaquinones include MK-8(H2 ), MK-7(H2 ), and minor
D-trehalose, turanose, Tweens 40 and 80, uridine, uridine amounts of MK-9(H2 ). The polar lipid profile includes
5′ -monophosphate, xylitol, and D-xylose. diphosphatidylglycerol, phosphatidylglycerol, an uniden-
Positive for alkaline phosphatase, β-galactosidase, and tified aminophospholipid, an unidentified glycolipid, an
pyrrolidonyl arylamidase, but negative for N-acetyl-β- unidentified polar lipid, and minor amounts of three
glucosaminidase, gelatin hydrolysis, β-glucosidase, β-glucu- unidentified phospholipids. Cadaverine and putrescine are
ronidase, urease, and fermentation of glycogen, D-lactose, major components in the polyamine profile.
D-mannitol, and D-ribose (API Coryne tests). Positive for Source: a sample from the wall of a house colonized with
esterase (C4), esterase lipase (C8), leucine arylamidase, molds in Jena, Germany.
naphthol-AS-BI-phosphohydrolase, and trypsin, and weakly DNA G+C content: not determined.
positive for cystine arylamidase and valine arylamidase (API Type strain: 01-Je-003, CCM 7649, DSM 22082.
ZYM tests). Additional phenotypic characteristics are shown Sequence accession no. (16S rRNA gene): FN293377.
......................................................................................................................................................................................................
Additional comments: the type strain of Brevibacterium san- Bousfield, I.J. 1972. A taxonomic study of some coryneform
darakinum showed relatively low DNA–DNA relatedness to bacteria. J. Gen. Microbiol. 71: 441–455.
Brevibacterium marinum (31.0%, reciprocal 35.7%) and Bre-
Bousfield, I.J., G.L. Smith, T.R. Dando and G. Hobbs. 1983.
vibacterium picturae (36.3%, reciprocal 37.9%) (Kämpfer et
Numerical analysis of total fatty acid profiles in the iden-
al., 2010).
tification of coryneform, nocardioform and some other
Brevibacterium sanguinis bacteria. J. Gen. Microbiol. 129: 375–394.
Wauters, Haase, Avesani, Charlier, Janssens, Van
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tion of higher ranks of the class Actinobacteria, with the
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Brevibacterium Lengkap

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Genus

Metabolism Antibiotic sensitivity

and contain meso-A2 pm in the wall peptidoglycan. Mor-

Polar lipid patterns Taxonomic comments

TABLE 1. Characteristics differentiating species of the genus Brevibacteriuma

a Symbols: +, positive; −, negative; w, weak; v, variable; nd, not determined.

wy, whitish-yellow; yellow; yo, yellow to orange.

Brevibacterium album arylamidase, esterase (C4), esterase lipase (C8), leucine

................................................................................... L-fucose, β-gentiobiose, 2-ketogluconate, 5-ketogluconate,

valine arylamidase (API ZYM tests). Additional phenotypic Brevibacterium celere

Source: blood. Brevibacterium picturae

Arginine dihydrolase, β-galactosidase, lysine decarboxy- Brevibacterium salitolerans

Cannon, J.P., S.L. Spandoni, S. Pesh-Iman and S. Johnson.

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