Genus

Firmicutes/Clostridia/Clostridiales/Eubacteriaceae/

Acetobacterium
Balch, Schoberth, Tanner and Wolfe 1977, 360AL
..........................................................................................................................................................................................
Maria V. Simankova, Russian Academy of Sciences, Winogradsky Institute of Microbiology, 7/2, Prospekt 60-letiya Oktyabrya,
Moscow 117312, Russia
Oleg R. Kotsyurbenko, Helmholtz Centre for Infection Research, Environmental Microbiology Laboratory, Inhoffenstrasse 7,
Braunschweig D-38124, Germany

A.ce.to.bac.teri.um. L. n. acetum vinegar; Gr. neut. n.
bakterion a small rod; N.L. neut. n. Acetobacterium vine-
gar rod.
Oval-shaped, short rods. Gram-stain-positive.
Motile. Endospores not formed. Strictly anaerobic.
Optimal temperature 2730 C for mesophilic species,
2030 C for psychrotolerant species. Optimal pH
7.08.0. Colonies are convex, white, slightly yellow, or
brownish, 0.61.0 mm in diameter. Autotrophic growth
occurs by anaerobic oxidation of H2 and reduction of
CO2 to acetic acid. Chemo-organotrophic, carrying out
homoacetogenic fermentation of reduced substrates,
such as fructose and some other monomeric sugars,
as well as pyruvate, lactate, glycerol, and methanol;
methyl groups of phenyl methyl ethers and betaine are
converted to acetate. The acetyl-CoM pathway serves as
an energy-conserving process and as a mechanism for
autotrophic assimilation of carbon. Cytochromes have
not been detected.
DNA G + C content (mol%): 3945.8.
Type species: Acetobacterium woodii Balch, Scho-
berth, Tanner and Wolfe 1977, 360.
..................................................................................
Oval-shaped, short rods. Gram-stain-positive.
Endospores not formed. Strictly anaerobic. Optimal tem-
perature 2730 C for mesophilic species, 2030 C for
psychrotolerant species. Optimal pH 7.08.0. Colonies are
convex, white, slightly yellow, or brownish, 0.61.0 mm
......................................................................................................................................................................................................
in diameter. Autotrophic growth occurs by anaerobic
oxidation of H2 and reduction of CO2 to acetic acid.
Chemo-organotrophic, carrying out homoacetogenic fer-
mentation of reduced substrates, such as fructose and some
other monomeric sugars, as well as pyruvate, lactate, glycerol,
and methanol; methyl groups of phenyl methyl ethers and
betaine are converted to acetate. The acetyl-CoM pathway
serves as an energy-conserving process and as a mechanism
for autotrophic assimilation of carbon. Cytochromes have
not been detected.
DNA G + C content (mol%): 3945.8.
Type species: Acetobacterium woodii Balch, Schoberth, Tan-
ner and Wolfe 1977, 360.
Number of validated species: 8
Further descriptive information
Oval-shaped, short rods 0.71.0 2.04.0 m, single or in
pairs (Figure 1). Motile by means of one or two subtermi-
nal flagella or peritrichous flagella. Swollen and elongated
cells can appear under nonoptimal growth conditions.
Psychrotolerant species of the genus form swollen cells at
temperatures higher than optimal ones, and usually the
number of the swollen cells reaches 5% of the cell number.
However, at 2730 C swollen cells of the psychrotolerant
Motile. species Acetobacterium tundrae represent 9095% of the cell
number; the size of these cells reaches 23 m in width
and 1015 m in length (Simankova et al., 2000). At tem-
peratures higher than 25 C, the cell size increases, but cell
division does not occur. The breakage of the cell wall of

Bergeys Manual of Systematics of Archaea and Bacteria, Online 2015 Bergeys Manual Trust. This article is 2009 Bergeys Manual Trust.
DOI: 10.1002/9781118960608.gbm00626. Published by John Wiley & Sons, Inc., in association with Bergeys Manual Trust.
 2 Bergeys Manual of Systematics of Archaea and Bacteria
FIGURE 1. Micrograph of Acetobacterium woodii. Bar = 0.5 m.
swollen cells is sometimes observed, occurring when the cell
content reaches a critical amount. If swollen cells grown at
2530 C are transferred to a new fresh medium and are
cultivated at 20 C, normal-sized cells develop. Analysis of
the lipid complex of the cell membrane was carried out with
Acetobacterium bakii, Acetobacterium fimetarium, Acetobacterium
paludosum, and Acetobacterium tundrae. A peculiarity of the
lipid complex structure is the presence of a large amount of
plasmalogens, which are detected as aldehydes released after
sample processing (Kotsyurbenko et al., 1995; Simankova
et al., 2000). The main components of the lipid complex
of the species studied are the saturated fatty acid C16:0 , the
unsaturated fatty acids C16:1 9 and C16:111 , as well as the
saturated aldehyde C16:0 ; however, the latter is absent from
the lipid complex of Acetobacterium fimetarium.
The cell wall has a Gram-stain-positive structure and con-
tains the more rare peptidoglycan of the cross-linkage type
B (Braun and Gottschalk, 1982; Eichler and Schink, 1984),
in which a L-seryl residue replaces the L-alanyl residue in
position 1 of the peptide subunit, and the ornithinyl residues
function as interpeptide bridges (Kandler and Schoberth,
1979).
Colonies are convex, white, 0.61.0 mm in diameter.
Some species produce slightly yellow (Acetobacterium woodii,
Acetobacterium carbinolicum) or brownish (Acetobacterium
wieringae) colonies.
The members of the genus Acetobacterium are metabol-
ically versatile organisms. They are able to grow both
autotrophically and chemo-organotrophically, catalyzing
the formation of acetate from C1 units as the sole or
the major end product of their energy metabolism, pro-
ceeding through the acetyl-CoA (the WoodLjungdahl)
pathway.
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H2 + CO2
...................................................................................
All members of the genus use H2 as the electron donor for
CO2 reduction to acetate. Some species can also similarly
utilize formate and CO, which are intermediates in acetate
formation from CO2 . In the acetyl-CoA pathway, the methyl
group of acetate is derived from CO2 via formate and
tetrahydrofolate-bound C1 intermediates and is then trans-
ferred to corrinoid enzyme prior to incorporation into the
methyl group of acetyl-CoA. The first step of carboxyl group
synthesis is CO2 reduction to a carbonyl group by means
of a carbon monoxide dehydrogenase, the key enzyme
of this metabolic pathway. It also catalyzes the acetyl-CoA
formation from the carbonyl group, the methyl group of
methyl-tetrahydrofolate, and coenzyme A. In the exper-
iments with Acetobacterium woodii, it was found that the
acetyl-CoA pathway is coupled with the generation of a
primary sodium ion potential, which in turn drives ATP
synthesis via Na+ -translocating ATP synthase (Aufurch et al.,
2000; Heise et al., 1992; Mller et al., 2001).
Chemo-organotrophic growth is provided by homoace-
togenic fermentation of different methylated compounds,
alcohols, monomeric sugars, hydroxyacids, and some other
substrates.
Methyl compounds
...................................................................................
Methanol, as well as methyl groups of phenylmethyl ethers,
methyl chlorides, and betaine is converted to acetate.
Methanol is converted to acetate with CO2 as a co-substrate.
The methyl group of methanol is transferred to tetrahy-
drofolate and is further transformed into the methyl group
of the acetate, whereas the carboxyl group is derived from
CO2 . Methoxylated aromatic compounds are demethylated
to the corresponding phenols; the methyl residue is fer-
mented analogously to methanol (Bache and Pfennig, 1981).
o-Demethylase catalyzes the cleavage of the ether bond of
phenyl methyl ethers and the transfer of the methyl group
to tetrahydrofolate (Kaufmann et al., 1997; Messmer et al.,
1996). Acetobacterium woodii dehalogenates chloromethanes
(CH2 Cl2 and CH3 Cl), converting the methyl groups to
acetate in an energy-yielding reaction. It can also catalyze
the reductive dechlorination of CCl4 and its substitutive
transformation to CO2 (Egli et al., 1988; Stromeyer et al.,
1991, 1992). CH3 Cl and CO2 as a co-substrate can also be
converted to acetate by Acetobacterium dehalogenans (Trau-
necker et al., 1991). The methyl transfer reaction is mediated
by a methyl chloride dehalogenase (Messmer et al., 1996;
Messmer et al., 1993; Wohlfarth and Diekert, 1997). Betaine
Bergeys Manual of Systematics of Archaea and Bacteria
is only demethylated to acetate and dimethylglycine (Eichler
and Schink, 1984).
Alcohols
...................................................................................
Acetobacterium carbinolicum is able to utilize aliphatic alcohols
(C2 C5 ), which are fermented to the corresponding fatty
acids and acetate (Eichler and Schink, 1984). Acetobacterium
woodii can also convert these substrates if the bicarbonate
buffer concentration is at least 100 mM; only weak growth is
supported by propanol, butanol, and pentanol (Buschhorn
et al., 1989).
Some species consume diols and acetoin, a component
of the butanediol cycle in microorganisms. 1,2-Propanediol
is fermented to propionate and acetate by Acetobacterium
carbinolicum, Acetobacterium wieringae, and Acetobacterium woodii
(Eichler and Schink, 1984; Schink and Bomar, 1992) or to
propanol, propionate, and acetate by Acetobacterium malicum
(Tanaka and Pfennig, 1988). 2,3-Butane, as well as acetoin,
is converted to acetate by Acetobacterium carbinolicum and
Acetobacterium woodii. A diol dehydratase is considered to
be an enzyme participating in diols cleavage. Acetobacterium
malicum, Acetobacterium bakii, Acetobacterium fimetarium, and
Acetobacterium paludosum can also utilize 2-methoxyethanol,
converting this substrate to methanol and acetaldehyde via
a diol dehydratase-analogous reaction; the latter product is
then oxidized to acetate (Tanaka and Pfennig, 1988).
Sugars and hydroxyacids
...................................................................................
One mol of fructose or any other monomeric sugar is fer-
mented via the EmbdenMeyerhofParnas pathway to 2 mol
of pyruvate, which is further oxidized to 2 mol of acetate and
2 mol of CO2 . The latter is also reduced to acetate. Similarly,
pyruvate and lactate are converted to acetate. Lactate is first
oxidized to pyruvate by a lactic dehydrogenase. Malate is
fermented to acetate by Acetobacterium malicum, Acetobacterium
bakii, Acetobacterium fimetarium, and Acetobacterium paludo-
sum presumably through an NAD-dependent malic enzyme
(Strohhcker and Schink, 1991).
Other substrates
...................................................................................
Acetobacterium woodii saturates the carbon-carbon double
bond of the acrylate side chain of caffeate derivatives and
converts these compounds to the corresponding hydrocaf-
feates (Bache and Pfennig, 1981). The process of caffeate
reduction is coupled to energy conservation (Hansen et al.,
1988; Tschech and Pfennig, 1984). The latter investigation
of hydrogen-dependent caffeate reduction by Acetobacterium
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3
woodii established chemiosmotic mechanism of ATP synthesis
by means of a transmembrane Na+ gradient generation
(Imkamp and Mller, 2002).
Mandelate (phenylglycolate) is oxidized by Acetobacterium
strain LuPhe1 via benzoyl-CoA to benzoate; carboxy carbon of
mandelate serves as one-carbon substrate for acetate forma-
tion (Dorner and Schink, 1991). Acetobacterium strain LuPhe1
also converts 2-phenoxyethanol to phenol and acetate
through acetaldehyde formation (Speranza et al., 2003).
The cytochromes have not been found in Acetobac-
terium woodii (Tschech and Pfennig, 1984), Acetobacterium
carbinolicum (Eichler and Schink, 1984), and Acetobacterium
malicum (Tanaka and Pfennig, 1988). Neither oxidase activity
has ever been reported. Catalase and superoxide dismu-
tase (SOD) activities were investigated in Acetobacterium
woodii, Acetobacterium paludosum, and Acetobacterium wieringae
(Brioukhanov et al., 2002). All studied species were defined
as catalase-positive, but only Acetobacterium wieringae exhibited
a high catalase activity. High specific activities of SOD were
detected in Acetobacterium woodii and Acetobacterium wieringae,
whereas Acetobacterium paludosum demonstrated a low SOD
activity.
All Acetobacterium species are able to fix molecular nitro-
gen, but this process has not been studied in detail (Schink
and Bomar, 1992).
The 16S rRNA gene sequence analysis shows that Aceto-
bacterium species form a tight phylogenetic group exhibiting
a high level (7% and higher) of sequence divergence with
most closely-related species of the genus Eubacterium (Willems
and Collins, 1996). The level of sequence similarity between
species of the genus Acetobacterium ranges between 96.2 and
99.4% (Simankova et al., 2000; Willems and Collins, 1996).
Psychrotolerant species do not exhibit a closer phylogenetic
relationship with each other than with mesophilic species.
DNADNA hybridization experiments between Acetobacterium
species show that hybridization levels are less than 37% for
all species combinations (Simankova et al., 2000). The only
exception is Acetobacterium woodii and Acetobacterium carbino-
licum demonstrating 69% of DNADNA homology. However,
16S rRNA gene sequence analysis shows a sequence similarity
of 98.4% between these species.
Streptomycin, benzylpenicillin, vancomycin, rifampin,
and bacitracin completely inhibit growth of Acetobacterium
bakii, Acetobacterium fimetarium, and Acetobacterium paludosum.
All strains of the genus Acetobacterium were isolated from
strictly anoxic environments. Most of Acetobacterium species
inhabit fresh water ecosystems: anoxic pond sediments,
ditches, wetlands, and anoxic sewage sludge. However, Ace-
tobacterium woodii strain WB1 was isolated from a marine
 4 Bergeys Manual of Systematics of Archaea and Bacteria
estuary, and laboratory experiments with Acetobacterium
strains demonstrated their equally good growth in fresh
water, brackish water, and salt-water medium (Schink, 1994).
In recent years, it was discovered that the acetyl-CoM pathway
is coupled to a chemiosmotic mechanism of ATP synthesis,
namely, to the presence of sodium-proton antiporters in the
cytoplasmic membrane. This finding explains the capacity
of Acetobacterium species to well adapt to environments with
periodical changes in salinity or pH, such as estuaries, soils, or
sewage sludge digestors. However, Acetobacterium species seem
to be more important for fresh water terrestrial ecosystems,
where they can produce acetate from a variety of different
substrates. Representatives of genus Acetobacterium are among
the most typical microorganisms in many cold fresh water
ecosystems (Kotsyurbenko et al., 1993a; Kotsyurbenko et al.,
1993b; Nozhevnikova et al., 1994). All four new psychrotol-
erant species of homoacetogenic bacteria isolated over the
last decade belong to this genus. They play an important eco-
logical role in different cold anoxic environments as strong
competitors of methanogenic archaea for hydrogen (Conrad
et al., 1989; Kotsyurbenko et al., 1996), the key intermediate
in the anaerobic microbial community. Under conditions
of H2 partial pressures higher than 10 Pa, psychrotolerant
homoacetogens are able to outcompete methanogens owing
to higher growth rates at low temperature (Kotsyurbenko
et al., 2001). It can result in essential changes in the trophic
structure of the microbial community and in production of
acetate as one of the end products.
Enrichment and isolation procedures
Enrichment is carried out using a liquid medium and
H2 :CO2 (80:20) as gas phase. Medium contains minerals,
vitamins, yeast extract, sodium sulfide as reducing agent,
sodium bicarbonate (2.5 g/l) as buffer, and resazurin (Kot-
syurbenko et al., 1995). The vitamin and microelement
solutions described by Wolin et al. (1963) and Pfennig and
Lippert (1966) are used, respectively. Yeast extract is not an
obligatory requirement of any species but enhances growth
yield. Methanogens can also be enriched under anoxic
conditions and an atmosphere of H2 /CO2 . However, they
are outgrown by faster-growing homoacetogenic bacteria
when transferred repeatedly on liquid medium using tenfold
serial dilutions. Moreover, methanogens can be inhibited
by the addition of 50 mg/l of sodium dithionite (Balch
et al., 1977) or 1050 mM bromoethanesulfonate (Smith
and Mah, 1981). Growth of Acetobacterium is judged from
acetate production and by observation of characteristic
cell morphology under the microscope. The last positive
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dilution is used to inoculate roll-tubes containing H2 /CO2
as gas phase. Colonies that developed in the roll-tubes
are transferred to liquid medium and cultivated under an
atmosphere of H2 /CO2 . Some species of the genus Aceto-
bacterium can be enriched using selective substrates, such
as methoxylated aromatic compounds (syringate, vanillate,
and trimethoxycinnamate ferulate) for Acetobacterium woodii
isolation (Bache and Pfennig, 1981), ethanol, propanol, or
butanol for Acetobacterium carbinolicum enrichment (Eichler
and Schink, 1984), and 2-methoxyethanol for Acetobacterium
malicum isolation (Tanaka and Pfennig, 1988).
Maintenance procedures
Storage is possible anaerobically in liquid medium at 4 C,
under these conditions, cells remain viable for 6 months. Ace-
tobacterium may be stored after lyophilization according to the
procedure for strict anaerobes (Hippe, 1991).
Differentiation of the genus Acetobacterium from
other genera
The genus Acetobacterium is phylogenetically differentiated
from other genera of homoacetogenic anaerobic members of
the Firmicutes (Gram-positive bacteria with a low DNA G + C
content) on the basis of oligonucleotide composition of the
16S rRNA gene. The phylogenetic relationship with the most
closely related species of the genus Eubacterium is at the level
of not more than 93% sequence similarity (Willems and
Collins, 1996). Acetobacterium is also distinguished from other
homoacetogenic anaerobic Gram-stain-positive bacteria on
the basis of morphological and physiological properties. In
contrast to homoacetogenic bacteria of the genus Clostrid-
ium, the members of the genus Acetobacterium do not form
endospores. Acetobacterium differ in their cell morphology
and peptidoglycan composition from the genera Peptostrep-
tococcus, Syntrophococcus, and Acetitomaculum. The members
of the genus Acetobacterium are distinguished from the only
known species of the genus Acetogenium on the basis of
their substrate spectrum, peptidoglycan composition, and
inability to grow at a high temperature. The members of the
genus Acetobacterium differ from homoacetogenic bacteria
of the genus Eubacterium by their ability to carry out only
homoacetogenic fermentation of reduced substrates.
Taxonomic comments
In the original description of the genus Acetobacterium it was
tentatively placed in the family Propionibacteriaceae (Balch
Bergeys Manual of Systematics of Archaea and Bacteria 5

FIGURE 2. Consensus dendrogram reflecting the phylogenetic relationships of the order Clostridiales (part one) within the class
Clostridia. Analyses were performed as described for Figure 3.

et al., 1977). Later, according to 16S rRNA gene sequence

List of the species of the genus Acetobacterium
data, the genus Acetobacterium was placed in the family
Eubacteriaceae belonging to cluster XV of the Clostridium
Acetobacterium woodii
subphylum of the Gram-positive bacteria (Willems and Balch, Schoberth, Tanner and Wolfe 1977, 360AL
Collins, 1996). According to the 16S rRNA phylogenetic ...................................................................................
analysis presented in the roadmap to this volume (Figure 2),
woodi.i. N.L. gen. n. woodii of Wood; named for H.G. Wood
the genus Acetobacterium is a member of the family Eubac-
teriaceae, order Clostridiales, class Clostridia in the phylum for his pioneering work on the total synthesis of acetate from
Firmicutes. CO2 by bacteria.

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 6 Bergeys Manual of Systematics of Archaea and Bacteria

FIGURE 3. Consensus dendrogram reflecting the phylogenetic relationships of the classes Bacilli and Erysipelotrichia
within the Firmicutes. The tree is based on maximum-likelihood analyses of a dataset comprising about 5000 almost
full-length high-quality 16S rRNA sequences from representatives of the Firmicutes and another 1000 representing the major
lines of decent of the three domains Bacteria, Archaea, and Eucarya. The topology was evaluated by distance matrix and
maximum-parsimony analyses of the dataset. In addition, maximum-parsimony analyses of all currently available almost com-
plete small-subunit rRNA sequences (137,400 of ARB-SILVA release 92, Prsse et al., 2007) were performed. Only alignment
positions invariant in at least 50% of the included primary structures from Firmicutes were included for tree reconstruction.
Multifurcations indicate that a common relative branching order was not significantly supported applying alternative treeing
methods. The (horizontal) branch lengths indicate the significance of the respective node separation.

Cells are oval-shaped, short rods 1.0 2.0 m. Motile by
means of one or two subterminal flagella. Colonies on plates
are circular, convex, and white; 1 mm in diameter. Older
colonies may show a slight yellow pigmentation. Optimal
growth occurs at 30 C. Autotrophic growth on H2 /CO2
and formate; acetate is formed. Methanol, 2,3-butanediol,
acetoin, glycerol, ethylene glycol, lactate, pyruvate, fruc-
tose, as well as methyl groups of methoxylated aromatic
compounds and betaine are fermented to acetate. Ethanol,
propanol, butanol, and pentanol are utilized if medium
contains at least 100 mM bicarbonate buffer. Ethanol is
converted to acetate; propanol, butanol, pentanol, and

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Bergeys Manual of Systematics of Archaea and Bacteria
1,2-propanediol are fermented to corresponding fatty acids
and acetate. Caffeate derivates are reduced to the corre-
sponding hydrocaffeate. Chloromethanes (CH2 Cl2 , CH3 Cl,
and CCl4 ) are dehalogenated; CH2 Cl2 and CH3 Cl are then
fermented to acetate, whereas CCl4 is either reduced to
CH3 Cl or transformed to CO2 .
Habitat: marine and fresh water sediments, and sewage
sludge.
DNA G + C content (mol%): 39 (Bd).
Type strain: WB1, DSM 1030, ATCC 29683.
GenBank accession number (16S rRNA gene): X96954.
Acetobacterium bakii
Kotsyurbenko, Simankova, Nozhevni-
kova, Zhilina, Bolotina, Lysenko and Osipov 1997,
242VP (Effective publication: Kotsyurbenko,
Simankova, Nozhevnikova, Zhilina, Bolotina, Lysenko
and Osipov 1995, 33.)
...................................................................................
baki.i. N.L. gen. n. bakii of Bak, named for F. Bak who isolated
the first psychrotolerant homoacetogenic bacterium.
Short rods, 0.91.5 1.52.7 m. Motile by means of
two subterminal flagella. Colonies on agar are white and
0.61.0 mm in diameter. Psychrotolerant. Temperature
range for growth is 130 C, with an optimum at 20 C.
Growth at pH values 5.58.5, with an optimum at 6.5.
Autotrophic growth on H2 /CO2, CO, and formate; acetate
is formed. Fructose, lactate, malate, as well as methanol
and methyl groups of vanillate and betaine are fermented
to acetate. Weak growth on maltose, glucose, xylose, and
2-methoxyethanol. The main components of the lipid com-
plex of the cell membrane are the saturated fatty acid C16:0,
the saturated aldehyde C16:0 , and the unsaturated fatty acids
C16:19 .
Isolated from anoxic sediments of a pond polluted by
paper-mill wastewater.
DNA G + C content (mol%): 42.1 (Tm ).
Type strain: Z-4391, DSM 8239, ATCC 51794.
GenBank accession number (16S rRNA gene): X96960.
Acetobacterium carbinolicum
Eichler and Schink 1985, 375VP (Effective publication:
Eichler and Schink 1984, 152.)
...................................................................................
car.bi.noli.cum. N.L. adj. carbinolicum metabolizing alcohols.
Rod-shaped cells, 0.81.0 1.52.5 m, with slightly
pointed ends, single or in pairs. Motile. Colonies on agar
are white or slightly yellow. Temperature range for growth is
1540 C, with an optimum at 20 C. Growth at pH range
6.08.0, with an optimum at 7.0. Autotrophic growth
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7
on H2 /CO2 and formate; acetate is formed. Methanol,
ethanol, 2,3-butanediol, acetoin, glycerol, ethylene gly-
col, lactate, pyruvate, glucose, fructose, methyl groups
of methoxylated aromatic compounds, and betaine are
fermented to acetate. Propanol, butanol, pentanol, and
1,2-propanediol are fermented to the corresponding fatty
acids and acetate. Does not require any vitamins. Sulfate,
thiosulfate, elemental sulfur, and nitrate are not reduced. No
cytochromes.
Isolated from anoxic freshwater mud.
DNA G + C content (mol%): 38.5.
Type strain: WoProp1, DSM 2925.
GenBank accession number (16S rRNA gene): X96956.
Acetobacterium fimetarium
Kotsyurbenko, Siman-
kova, Nozhevnikova, Zhilina, Bolotina, Lysenko and
Osipov 1997, 242VP (Effective publication:
Kotsyurbenko, Simankova, Nozhevnikova, Zhilina,
Bolotina, Lysenko and Osipov 1995, 34.)
...................................................................................
fi.me.tari.um. L. n. fimetum a dung-hill; N.L. neut. adj. fimetar-
ium inhabiting manure.
Short rods, 0.81.1 1.52.6 m. Motile by means of per-
itrichous flagella. Colonies on agar are white and 0.61.0 mm
in diameter. Temperature range for growth is 135 C, with
an optimum at 30 C. Growth at pH values 6.08.5, with an
optimum at 7.5. Autotrophic growth on H2 /CO2, CO, and
formate with acetate production. Fructose, lactate, malate
and methyl groups of vanillate and betaine are utilized
for growth and fermented to acetate. 2,3-Butanediol and
2-methoxyethanol are weakly used. The main components
of the lipid complex of the cell membrane are the saturated
aldehyde C16:0 and the unsaturated fatty acids C16:19 and
C16:111 .
Isolated from digested manure.
DNA G + C content (mol%): 45.8 (Tm ).
Type strain: Z-4290, DSM 8238, ATCC 51795.
GenBank accession number (16S rRNA gene): X96959.
cetobacterium malicum
Tanaka and Pfennig 1990, 470VP (Effective
publication: Tanaka and Pfennig 1988, 186.)
...................................................................................
mali.cim. N.L. neut. adj. malicum pertaining to malic acid.
Rod-shaped cells, 1.01.3 1.84.0 m, with slightly
pointed ends, single or in pairs. Motile. Colonies on agar are
white. Optimal temperature is 30 C, no growth at 16 C and
40 C. Optimal pH is 7.58.0, no growth at initial pH 6.0 and
9.5. Autotrophic growth on H2 /CO2 and formate; acetate is
 8 Bergeys Manual of Systematics of Archaea and Bacteria
formed. Fructose, lactate, pyruvate, malate, glycerol, acetoin,
ethylene glycol, 2-methoxyethanol, 2-ethoxyethanol, as well
as methyl groups of methoxylated aromatic compounds
and betaine are fermented to acetate. 1,2-Propanediol is
fermented to propanol, propionate, and acetate. Under
optimal conditions, doubling time on 2-methoxyethanol
and malate is 22 h and 7.5 h, respectively. Sulfate, sulfite,
thiosulfate, elemental sulfur, and nitrate are not reduced. No
cytochromes.
Isolated from anoxic freshwater sediment of a ditch.
DNA G + C content (mol%): 44.1 (Bd).
Type strain: MuME1, DSM 4132, ATCC 51201.
GenBank accession number (16S rRNA gene): X96957.
Acetobacterium paludosum
Kotsyurbenko, Simankova, Nozhevnikova, Zhilina,
Bolotina, Lysenko and Osipov 1997, 242VP (Effective
publication: Kotsyurbenko, Siman-
kova, Nozhevnikova, Zhilina, Bolotina, Lysenko and
Osipov 1995, 34.)
...................................................................................
pa.lu.dosum. L. neut. adj. paludosum inhabiting a fen.
Short rods, 0.81.1 1.32.9 m. Motile by means of
peritrichous flagella. Colonies in roll tube agar are white
and 0.61.0 mm in diameter. Psychrotolerant. Tempera-
ture range for growth is 130 C, with an optimum at 20
C. Growth at pH values 5.08.0, with an optimum at 7.0.
Autotrophic growth on H2 /CO2, CO, and formate; acetate is
formed. Maltose, fructose, glucose, lactate, malate, methanol,
as well as methyl groups of betaine are fermented to acetate.
2-Methoxyethanol, xylose, and cellobiose are weakly used.
The main components of the lipid complex of the cell
membrane are the saturated fatty acid C16:0, the saturated
aldehyde C16:0 , and the unsaturated fatty acids C16:19 and
C16:111. Habitat: anoxic freshwater sediments.
Isolated from anoxic sediments of a fen.
DNA G + C content (mol%): 41.7 (Tm ).
Type strain: Z-4092, DSM 8237, ATCC 51793.
GenBank accession number (16S rRNA gene): X96958.
Acetobacterium tundrae
Simankova, Kotsyurbenko, Stackebrandt, Kostrikina,
Lysenko, Osipov and Nozhevnikova 2001, 793VP
(Effective publication: Simankova, Kotsyurbenko,
Stackebrandt, Kostrikina, Lysenko, Osipov and
Nozhevnikova 2000, 446.)
...................................................................................
tundrae. N.L. fem. gen. n. tundrae from the tundra (a cold
treeless zone located in the north of Eurasia and North
America).
......................................................................................................................................................................................................
This article is 2009 Bergeys Manual Trust. Published by John Wiley & Sons, Inc., in association with Bergeys Manual Trust.
Cells are short rods, 0.71.1 1.14.0 m, with slightly
pointed ends. Motile by means of peritrichous flagella. The
irregular swollen cells 23 m 1015 m appear at tem-
peratures 2530 C as a result of a defect in cell division.
Colonies in agar roll tubes are white and 1.0 mm in diameter.
Psychrotolerant. Temperature range for growth is 130 C,
with an optimum 20 C. Growth at pH values 6.08.0, with an
optimum at 7.0. Autotrophic growth on H2 /CO2, CO, and
formate with acetate production. Maltose, mannose, fruc-
tose, glucose, xylose, pyruvate, lactate, methanol, and methyl
groups of betaine are utilized for growth and fermented
to acetate. The main components of the lipid complex of
the cell membrane are the saturated aldehyde C16:0 and the
unsaturated fatty acid C16:19 .
Isolated from tundra soil.
DNA G + C content (mol%): 39.2 (Tm ).
Type strain: Z-4493, DSM 9173.
GenBank accession number (16S rRNA gene): AJ297449.
Acetobacterium wieringae
Braun and Gottschalk 1983, 438VP (Effective
publication: Braun and Gottschalk 1982, 374.)
...................................................................................
wie.ringae. N.L. gen. n. wieringae of Wieringa; named for K.
T. Wieringa who isolated and described Clostridium aceticum,
the first known acetogenic bacterium.
Cells are oval-shaped, short rods 1.0 2.0 m. Motile by
means of one or two subterminal flagella. Colonies are circu-
lar, smooth, and brownish; 3 mm in diameter. Optimal growth
occurs at temperature 30 C and pH 7.27.4. Autotrophic
growth on H2 /CO2 and formate; acetate is formed. Fructose,
lactate, ethylene glycol, acetoin, glycerol, and ethanol are
fermented to acetate. 1,2-Propanediol is fermented to acetate
and propionate. Under optimal conditions, doubling time
on H2 /CO2 mixture corresponds to 10 h.
Isolated from sewage sludge.
DNA G + C content (mol%): 43 (Tm ).
Type strain: C, DSM 1911, JCM 2380, ATCC 43740.
GenBank accession number (16S rDNA gene): X96955.
Other organisms
Acetobacterium dehalogenans
Traunecker, Preu and Diekert 1991, 417
...................................................................................
de.ha.loge.nans. N.L. adj. part. dehalogenans dehalogenating.
The description of this species is based on the phenotypic
features and G + C content of the DNA. The data of phylo-
genetic analysis of the 16S rRNA gene is lacking. Cells are
elongated cocci 1.1 1.5 m arranged in chains. Nonmotile.
Bergeys Manual of Systematics of Archaea and Bacteria
Gram-stain-positive. Endospores not formed. Strictly anaero-
bic. Colonies are smooth, convex, and yellow, about 1 mm in
diameter. Optimal growth occurs at temperature 25 C and
pH 7.37.7. Autotrophic growth on H2 /CO2 and CO with
acetate production. Fructose, glucose, ribose, pyruvate, lac-
tate, glycerol, as well as methyl groups of phenyl methyl ethers
and methyl chloride are converted to acetate.
Isolated from a sewage digestor.
DNA G + C content (mol%): 47.5 (HPLC).
Type strain: MC.
Acetobacterium psammolithicum
Krumholz, Harris, Tay and Suflita 1999, 2302
...................................................................................
psam.mo.lithi.cum. Gr. fem. n. psammos sand; Gr. masc. n.
lithos stone. N.L. neut. adj. lithicum of stone; N.L. neut. adj.
psammolithicum of sandstone.
The taxonomic status of this organism is uncertain
because its detailed description is incomplete. Cells are
elongated rods 1.1 1.73.3 m. Flagella are not observed.
Gram-stain-negative. Endospores not formed. Strictly anaer-
obic. The substrates, which support growth include H2,
formate, methanol, glucose, syringate, pyruvate, lactate,
betaine, ethanol, propanol, glycerol, and acetoin. According
to 16S rDNA sequence data and substrate specificity, the
organism could be affiliated with the genus Acetobacterium.
However, its morphology is not typical for the genus Acetobac-
terium: the organism was described as Gram-stain-negative,
and flagella were not observed. Temperature and pH char-
acteristics of growth, as well as the G+C content of the DNA
are lacking.
Isolated from a subsurface sandstone.
Type strain: CN-E.
Acknowledgements
We are grateful to N.A. Kostrikina, who took the electron
micrograph presented in this description.
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Further reading
Dierert, G. and G. Wohlfarth. 1994. Metabolism of
homoaceto gens. Antonie van Leeuwenhoek. 66: 209221.
Drake, H.L. 1994. Acetogenesis, acetogenic bacteria, and the
Acethyl-CoA Wood/Ljungdahl pathway: past and current
perspectives. In Drake (Editor), Acetogenesis, Chapman &
Hall, New York, London, pp. 360.
Schuppert, B. and B. Schink. 1990. Fermentation of
methoxyac etate to glycolate and acetate by newly iso-
lated strains of Ace tobacterium sp. Arch. Microbiol. 153:
200204.