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E. Rosenberg et al. (eds.), The Prokaryotes Alphaproteobacteria and Betaproteobacteria, DOI 10.1007/978-3-642-30197-1_396,
# Springer-Verlag Berlin Heidelberg 2014
Abstract
The family Acetobacteraceae is taxonomically included in the
order Rhodospirillales of the class Alphaproteobacteria, and 32
genera are validly described. The genera are basically classified
into two groups, an acetous group and an acidophilic group, in
the light of application, ecology, and phylogeny. The acetous
group comprises genera in acetic acid bacteria like Acetobacter,
Gluconacetobacter, Gluconobacter, Asaia, Granulibacter, and
Komagataeibacter. The acidophilic group consists of acidophilic
and neutrophilic genera like Acidiphilium and Roseomonas. In
the 1960s, taxonomy of acetic acid bacteria was significantly
Molecular Analyses
Phylogenetic Structure of the Family and Its
Genera
The family Acetobacteraceae is a member of the order
Rhodospirillales in the class Alphaproteobacteria (Sievers and
Swings 2005a). In 1980, the genera Acetobacter and
Gluconobacter were genetically separated on the basis of DNArRNA hybridizations and united in the family Acetobacteraceae
(Gillis and De Ley 1980). Moreover, the genera Acetobacter and
Gluconobacter were shown to constitute one large rRNA superfamily, which was shown as the alpha subclass of the
Proteobacteria (Stackebrandt et al. 1988), later named the class
Alphaproteobacteria (Garrity et al. 2005), together with the genera Agrobacterium, Rhizobium, Zymomonas, etc. On the basis of
5S rRNA gene sequencing, the genus Acidomonas was recognized
to form an independent lineage apart from the genera
Acetobacter and Gluconobacter in the family Acetobacteraceae
(Bulygina et al. 1992). The genus Gluconacetobacter was phylogenetically distinguished from the genera Acetobacter and
Gluconobacter by the partial 16S rRNA gene sequence analysis
(Yamada et al. 1997). Meanwhile, acetic acid bacteria were indicated to relate to acidophilic bacteria including the genera
Acidiphilium, Acidocella, and Rhodopila (Kishimoto et al. 1995;
Sievers et al. 1994a). Bacteriochlorophyll a-containing bacteria
such as the genera Rhodopila and Roseococcus were moderately
related to the genus Acidiphilium (Yurkov et al. 1994).
Therefore, the family Acetobacteraceae mainly consists of acetic
acid bacteria, acidophilic bacteria, and bacteriochlorophyll
a-containing bacteria.
Currently, at least 32 genera are thought to be the member of
the family Acetobacteraceae. The phylogenetic relationships of
the family Acetobacteraceae are shown in > Fig. 1.1. The 30
genera except for the genera Stella and Zavarzinia form
Genome Analyses
Approximately 20 complete and draft genome sequences of
the branching within the 16S rRNA gene tree of the family
Acetobacteraceae have been reported in public. The genome
10
. Fig. 1.1
Neighbour-joining tree of the family Acetobacteraceae and related bacteria based on 16S rRNA gene sequences. Phylogenetic analysis
was based on alignment of 1,219 bp 16S rRNA gene sequences of 114 strains. Solid circles at branching nodes indicate supporting
probabilities above 95 % by two phylogenetic analysis methods (NJ and ML), and open circles indicate probabilities above 85 % by either
analysis. Bar, 0.01 substitutions per nucleotide position
11
12
Phages
A bacteriophage A-1 was isolated from Gluconobacter oxydans
strain E and characterized to be of type A phage according to
Bradleys morphological classification (Bradley 1967; Jucker and
Ettlinger 1981; Robakis et al. 1985a; Schocher et al. 1979).
Abnormalities in the microbial process for the oxidation of
D-sorbitol to L-sorbose by G. oxydans were indicated to be
Biochemical Properties
Bioenergetics
Acetic acid bacteria, the obligate aerobes, are biochemically quite
unique. The Embden-Meyerhof pathway is not operative for
strains assigned to the genera Acetobacter and Gluconobacter,
but instead, the pentose phosphate cycle is functioning (Hauge
et al. 1955; Cheldelin 1961; De Ley 1960; Asai 1968). In addition,
the strains of the genus Gluconobacter lack the TCA cycle, differing from the strains of the genus Acetobacter, and the pentose
phosphate cycle acts as the only terminal oxidative catabolic
pathway. Glucose-6-phosphate dehydrogenase and 6phosphogluconate dehydrogenase reduce NAD, except for
NADP, in the cells of the acetic acid bacteria, and the two
dehydrogenases seem to play an important role in the respiratory chain phosphorylation (Cheldelin 1961).
Acetic acid bacteria are also known to accumulate
a large quantity of acetic acid from ethanol; dihydroxyacetone
from glycerol; D-gluconic acid, 2-keto-D-gluconic acid, and
D-Fructose
D-Fructose
D-Fructose
D-Fructose 6-phosphate
NADP
Fructose
dehydrogenase
5-Ketofructose reductase
NADPH2
5-Keto-D-fructose
Medium
5-Keto-D-fructose
Protoplasmic
membrane
5-Keto-D-fructose
The pentose
phosphate cycle
Cytoplasm
. Fig. 1.2
A presumed pathway for the catabolism of D-fructose in Gluconobacter species. 5-Ketofructose reductase is active only for
reduction, and the reverse reaction is scarcely found. The pH optimum of the reductase is 7.0 (The figure was cited from Yamada
et al. (1967b))
13
14
. Table 1.1
Membrane-bound primary dehydrogenases in acetic acid bacteria
Dehydrogenase
Substrate
Product
Alcohol
dehydrogenase
[EC 1.1.5.5]
Ethanol
Acetaldehyde
Aldehyde
dehydrogenase
[EC 1.2.99. 7]a
Acetaldehyde
Glucose
dehydrogenase
[EC 1.1.5.2]
DGlucose
Gluconate
2-dehydrogenase
[EC 1.1.99.3]a
Acceptor
Species
72 Kd (PQQ/heme c), 48 Kd
(3 hemes c), 15 Kd
Ubiquinone
G. suboxydans (1)
84 Kd (Iron sulfur?/
molybdopterin cytosine
dinucleotide?), 49 Kd
(3 hemes c), 17Kd
D-Glucono-d-lactone
80 Kd (PQQ)
Ubiquinone
G. suboxydans (1)
D-Gluconate
2-Keto-D-gluconate
64 Kd (FAD), 45 Kd
(3 hemes c), 21 Kd
Ubiquinone
G. dioxyacetonicus (1)
2-Keto-Dgluconate
dehydrogenase
[EC 1.1.99.4]a
2-Keto-D-gluconate
2,5-Diketo-D-gluconate
61 Kd(FAD), 47 Kd
(3 hemes c), 25 Kd
Ubiquinone
G. melanogenes (1)
Fructose
5dehydrogenase
[EC 1.1.99. 11]a
DFructose
5KetoDfructose
67 Kd (FADc), 51 Kd
(3 hemes c), 20 Kd
Ubiquinone
G. industrius (1, 7)
L-Sorbose
5-dehydrogenase
[EC 1.1.99.12]
L-Sorbose
5KetoDfructose
L-Sorbose
1-dehydrogenase
[EC 1.1.99.32]a
L-Sorbose
L-Sorbosone
L-Sorbosone
dehydrogenase
L-Sorbosone
2-Keto-L-gulonate
48 Kd?
Ubiquinone?
Sorbitol
dehydrogenase
[EC 1.1.99. 21]a
D-Sorbitol
L-Sorbose
63 Kd (FAD), 51 Kd
(3 hemes c), 17 Kd
Ubiquinone
G. suboxydans (1)
Glycerol
dehydrogenase
[EC 1.1.99. 22]a
Glycerol, meso
Erythritol, DArabitol,
DSorbitol, etc.
Dihydroxyacetone,
LErythrose, DXylulose,
LSorbose, etc.
80 Kd (PQQ), 14 Kd
Ubiquinone
G. industrius (3),
Acetic acid
A. aceti (1)
A. polyoxogenes (1),
A. europaeus (8)
58 Kd (FAD)
Ubiquinone
G. oxydans (1),
G. melanogenus (5)
For more details, see Matauahita et al. (2004), and Yakushi and Matsushita (2009, 2010)
a
The electron acceptor was proved to be ubiquinone. Therefore, the dehydrogenases will be classified as EC class 1.1.5
b
Unpublished data of Yakushi et al. (from Dr. Matsushita)
c
The prothsthetic group is assumed to be FAD, but it is not proved biochemically
References: (1), Matsushita et al. (1994); (2), Sato et al. (1969b); (3), Ameyama et al. (1985); (4), Adachi et al. (2001); (5), Sugisawa et al. (1991); (6), Sugisawa and
Hoshino (2002); (7), Kawai et al. (2013); (8), Thurner et al. (1997; and (9) Matsushita et al. (2003))
ADH
E
2H +
PQQ
c
bo 3
FDH
GDH
ACA
GL
5KF
FAD#
PQQ
c*
Periplasm
c*
CIO
Cytoplasmic
membrane
UQ
FAD#
2H +
KCN
+
1/2O 2
Cytoplasm
2H +
H2O
+
1/2O 2
NAD +
NADH
+
H+
NDH-II#
2H +
H2O
. Fig. 1.3
The membrane-bound dehydrogenases and the respiratory chain of Gluconobacter suboxydans. The dehydrogenases, e.g., alcohol
dehydrogenase, glucose dehydrogenase and fructose 5-dehydrogenase that contain prosthetic groups such as PQQ, FAD and heme c
generally reduce ubiquinone, an electron acceptor. The resulting electrons are then transferred either to the energy-producing
cytochrome o (bo3), a quinol oxidase or to the less energy-producing cyanide-insensitive quinol oxidase. For more details, see the
references (Matsushita et al. 2004; Yakushi and Matsushita 2009, 2010). Abbreviations: ADH alcohol dehydrogenase, GDH glucose
dehydrogenase, FDH fructose 5-dehydrogenase, E ethanol, ACA acetaldehyde, G D-glucose, GdL D-glucono-d-lactone, F D-fructose, 5KF
5-keto-D-fructose, FAD# the presence of FAD is assumed but not proved biochemically, NDH-II# Type-II NADH dehydrogenase, which is
not purified; bo3, cytochrome bo3; CIO cyanide-insensitive terminal quinol oxidase, c heme c; c* cytochrome c
6.0
3.0
40
30
1.5
20
1.0
10
0.5
10
20
30
40
Growth
(dry weight, mg/ml)
Concentration of ketohexose
and 5-keto-D- fructose (mg/ml)
4.0
pH
5.0
50
15
16
growth was generally seen (> Fig. 1.4; Sato et al. 1969a).
L-Sorbose accumulation reached the maximum 10 h later after
inoculation, and then the amount of L-sorbose decreased. The
production of 5-keto-D-fructose started 7 h later. After 16 h, the
first growth of the organism mostly stopped, and the organism
showed a temporarily stationary phase for approximately 10 h.
Of the two steps of the D-sorbitol oxidation, glycerol
dehydrogenase [EC 1.1.99.22] but not sorbitol dehydrogenase
[EC 1.1.99.21] first appeared to function (Soemphol et al. 2008),
and L-sorbose 5-dehydrogenase [EC 1.1.99.12] was secondly
concerned (Sato et al. 1969b), viz., L-sorbose production from
D-sorbitol and 5-keto-D-fructose production from L-sorbose.
The second growth began after 25 h from inoculation. The first
growth is due to the oxidations of D-sorbitol to L-sorbose and
then to 5-keto-D-fructose, which seems to be coupled with
the respiratory chain phosphorylation, and the second growth
is assumed to be due to the reduction of 5-keto-D-fructose
to D-fructose, followed by phosphorylation to D-fructose
6-phosphate and by flowing into the pentose phosphate cycle,
the terminal oxidative catabolic pathway of the organism. The
biphasic growth based on the oxidation of ethanol to acetic acid
and then to carbon dioxide and water can be seen in vinegar
production by an Acetobacter strain (Matsushita et al. 2004).
Nitrogen Fixation
Acetic acid bacteria capable of fixing molecular nitrogen seemed
to be restricted only to those of the three genera, Gluconacetobacter, Swaminathania, and Acetobacter (Pedraza 2008).
Acetobacter
diazotrophicus
(
Gluconacetobacter
diazotrophicus) was first reported as a nitrogen-fixing acetic
acid bacterium (Gillis et al. 1989; Cavalcante and Dobreiner
1988). Strains of the species were found inside sugarcane plant
tissues. As the second examples, Gluconacetobacter johannae and
Gluconacetobacter azotocaptans were found to be associated with
coffee plants (Fuentes-Ramrez et al. 2001). Swaminathania
salitolerans was described as a salt-tolerant, nitrogen-fixing, and
phosphate-solubilizing bacterium (Loganathan and Nair 2004).
Acetobacter peroxydans and Acetobacter nitrogenifigens were isolated, respectively, from wetland rice plants and kombucha tea
(Muthukumarasamy et al. 2005; Dutta and Gachhui 2006).
Recently, Asaia species, including Asaia bogorensis and
Asaia siamensis, were reported to have the capability of nitrogen
fixation (Samaddar et al. 2011). Nguyenibacter vanlangensis
found in rhizosphere or root of Asian rice plants showed the
growth on nitrogen-free LGI medium. The strains of the species
would be candidates able to fix molecular nitrogen (Vu et al. 2013).
The nitrogenase of Gluconacetobacter diazotrophicus was
immunochemically investigated from the viewpoint of the protection against molecular oxygen (Ureta and Nordlund 2002).
The nitrogenase in the organism could be protected by conformational mechanisms involving a putative FeSII protein operating when cells were subjected to sudden increases in oxygen
pressure, together with increased respiration of the organism
under diazotrophic condition.
Phenotypic Analyses
Currently, the family Acetobacteraceae is basically consists of two
groups, an acetous group and an acidophilic group. Additionally, the genus Frateuria is listed in this section. The genus
resembles biochemically the genera in acetic acid bacteria, but it
locates in the Gammaproteobacteria (Swings and Sievers 2005).
+/
+/
L-Arabinose
L-Sorbose
DXylose
5-Keto-D-gluconate
2,5-Diketo-D-glucoante
Acid from
*
*
2-Keto-D-gluconate
+
*
+/
D-Gluconate
Production of
+w
+w
+w
/+w
/+
+/+W
+/+w
+/+W
+/
Peritrichous
0.50.9 1.02.0
Ellipsoidal to rods
G. liquefaciens
Smooth,
entire,
creamy
Subpolar
Coccoid to
rods
E. medicaginis
+/
+/
Raised,
smooth,
entire,
shiny, white
or pink or
brown
Polar
0.61.0
1.03.0
Rods
G. oxydans
GluconoGluconacetobacter bacter
Peritrichous to lateral
0.41.0 0.82.5
Rods
A. bogorensis
Endobacter
+w
Polar
Lactate
Convex, circular,
smooth, entire,
beige to pink
Production of DHA
Acetate
Oxidation of
Colonial appearance
_
+
Mannitol agar
Glutamate agar
Growth on
Oxidase
Catalase
Polar
+/
Peritrichous
Motility
Flagella
0.41.0 1.03.0
Size (mm)
Rods
Rods
Ellipsoidal to rods
A.
chiangmaiensis
Cells
A. methanolica
Ameyamaea Asaia
A. aceti
Acidomonas
Type species
Acetobacter
. Table 1.2
Characteristics of the genera in the acetous group
+w
+w
Convex,
smooth, entire,
yellow (non
diffusible)
+w
Coccobacillus
to rods
G. bethesdensis
Granulibacter
1
17
+/
DGalactose
DGlucose
D-Mannose
Dulcitol
1-Butanol
Production of water
soluble pigment(s)
Production of cellulose
Growth on N2 free
medium
Assimilation of
ammoniac nitrogen
Growth on methanol
2-Propanol
Acetate
1-Propanol
Glycerol
Ethanol
Xylitol
D-Sorbitol
meso Ribitol
D-Mannitol
myo -Inositol
/+
Raffinose
+ (D-glucose)
+
+
Endobacter
/+
+/
Trehalose
meso Erythritol
Sucrose
+/+W
Melibiose
+/
Ameyamaea Asaia
Maltose
DFructose
Lactose
L-Rhamnose
+
Acidomonas
+/
+/
+/
+/+W
+/+W
+/
+/
/+
/+
GluconoGluconacetobacter bacter
+ (D-glucose)
Granulibacter
Acetobacter
18
The Family Acetobacteraceae
Remarks
*not
accumulated in
culture media
6263
Isolation sources
5261
Q-10
62
Q-9
Major isoprenoid
quinone
C18:1 o7c
Pathogenicity
Flower of
red ginger
(Alpinia
purpurea)
66
Q-10
6061
Q-10
C18:1o7c; C16:0
A surface
sterilized
nodule of
alfalfa
(Medicago
sativa )
60.3
Q-10
6267
Q-10
C18:1o7c;
C18:1o7c
C19;0cycloo8c;
C16:0
3.5
Max. pH
Min. pH
5.07.0
2030 C
28 C
1530 C
2037 C
Opt. pH
C18:1o7c
Not grow at 42 C
1037 C
3.08.5
C18:1o7c
Range of growth pH
3.08.0
/+
Pantothenic acid
Growth at pH 3.0
Min. temperature
Max. temperature
Opt. temperature
Range of growth
temperature
Growth at 37 C
Growth on 30 % glucose
Requirement of acetic
acid
Nutritional requirement
Dahlia,
Chinese
bayberry,
strawberry,
rambutan,
kaki, grape,
spoiled
jackfruit
5462
Q-10
C18:1o7c
Nicotinic
acid/no
Lymph node
culture of
a chronic
granulomatous
patient
59.1
Q-10
C18:1o7c; M16:0
To humans
Not at pH 3.5
5.06.0
4.07.5
Not grow at
42 C
3537 C
2537 C
1
19
+
L-Sorbose
L-Arabinose +
Acid from
+/
5-Keto-Dgluconate
2,5,-Diketo-
D-Gluconate
+w
2-Keto-Dgluconate
D-Gluconate +
Production of
Production of
DHA
+/+w
+w
+
+w
Negligible or no
Smooth, entire,
creamy
Raised,
smooth,
entire, shiny,
pink
Straight rods
0.81.0 2.54.0
+w
+w
Lactate
Acetic acid
from ethanol
Acetate
Oxidation of
Raised, convex to
umbonate, smooth to
rough, entire to irregular,
butyrous
Mannitol
agar
Colonial
appearance
Glutamate
agar
Growth on
Oxidase
+
Rods
0.60.8 1.0
1.6
Peritrichous
Rods
0.60.8 1.01.6
Motility
0.81.0
1.02.0
Rods
N. vanlangensis S. floricola
Nguyenibacter Saccharibacter
Catalase
0.60.8
2.03.0
Neokomagataea
N. chiangmai- N. thailandica
ensis
Neoasaia
Flagella
Rods
Coccoid to rods
0.60.8 1.03.0
Size (mm)
K. baliensis
Cells
K. xylinus
Type species
Kozakia
+w
0.60.8
1.01.6
Rods
T. sakaeratensis
Tanticharoenia
+w
+w
Peritrichous
0.70.9
1.93.1
Straight rods
S. salitolerans
Suwaminathania
Komagataeibacter
20
The Family Acetobacteraceae
Sucrose
Trehalose
+
+
Raffinose
Dulcitol
+
+/
+w
Maltose
+w
+w
+w
+
+w
+
Melibiose
Lactose
+w
DGlucose
D-Mannose
DFructose
DGalactose +/
+w
+/+W
+w
+w
L-Rhamnose
+w
+w/
+
+
+
+w
+w
L-Sorbose
+/
+w
+d
+w/-
+/
+/
+w/-
+/+W
DXylose
Growth on
1-Butanol
2-Propanol
1-Propanol
+/
Glycerol
Ethanol
D-mannitol
D-sorbitol
meso
Erythritol
+
+
+w
+
Raffinose
Dulcitol
Trehalose
+/
Sucrose
+d
+
Melibiose
Maltose
Lactose
D-Mannose
DGlucose
DFructose
+
DGalactose +
L-Rhamnose
DXylose
1
21
+/
Growth on
30 % glucose
Growth at
37 C
Growth on
1.0 % KNO3
/+
Requirement
of acetic acid
Nutritional
requirement
Growth in the +
presence of
0.35 % acetic
acid
Production of
watersoluble
pigment(s)
Production of
cellulose
Growth on N2
free medium
Assimilation
of ammoniac
nitrogen
Growth on
methanol
Acetate
1-Butanol
2-Propanol
1-Propanol
+w
+w
+w
+ (mannitol)
+**
+
+/
Ethanol
Glycerol
+w
+w
Tanticharoenia
Suwaminathania
D-Sorbitol
D-Mannitol
Nguyenibacter Saccharibacter
+
Neokomagataea
Neoasaia
meso
Erythritol
Kozakia
Komagataeibacter
22
The Family Acetobacteraceae
C18:17c; C16:0
C18:1o7c
Q-10
5664
Major
isoprenoid
quinone
G+C content
of DNA
Production
of fructan
from
sucrose
Remarks
Flower of red
ginger
(Alpinia
purpureus)
63.1
Q-10
Flower of lantana
(Lantana camera ),
candle bush (Senna
alata)
5157
Q-10
Rhizosphere
and roots of
Asian rice
6869
Q-10
Wild rice
(Proteresia
coarctata )
Fix nitrogen
and solubilize
phosphate
Osmophilic. *no growth on ordinary
glutamate agar, but grown on the
medium supplemented with 7 %
glutamate.
5860
Q-10
Pollens of flowers
5253
Q-10
C18:0o7c/o9t/
ol2t
Symbols: +, positive in most species; +w, weakly positive; +/, positive in more than half species; /+, negative in more than half species; v, variable; and , negative in most species
Palm
Many kinds of vinegar,
brown
Kombucha, nata de coco,
cider, apple juice, beet juice, sugar, ragi
cherry
57
Q-10
Isolation
sources
G+C content
of type strain
C18:1o7c
Major cellular
fatty acid(s)
below 4.0
Min. pH
C18:1o7c;
C16;0
above 8.0
Max. pH
Pathogenicity
5.07.0
Opt. pH
2530 C
2033 C
4.07.5
2537 C
Range of
growth pH
Growth at pH
3.0
Min.
temperature
Max.
temperature
Opt.
temperature
Range of
growth
temperature
Soil in Thailand
6566
Q-10
1
23
24
NCCB
23001T
NBRC
LMG
1504T
T
T
14818 NCIMB 8621 ), isolated from beechwood
25
26
Type strain: RG1T ( LMG 23498T MTCC 6912T), isolated from kombucha tea. The G+C content of DNA of the type
strain is 64.1 mol%.
Cells are Gram negative and short rods with rounded ends,
measuring 0.50.8 by 1.52.0 mm, and are occasionally up to
4 mm in length. Cells occur singly, in pairs, or (rarely) in short
chains and are either motile with a single polar flagellum or
nonmotile. Cells with polar tuft flagella are found very rarely.
Endospores are not produced. Colonies are shiny, smooth, circular, convex, entire, beige to pink, and 13 mm in diameter on
PYM agar (pH 4.5) after 5 days at 30 C. Pellicles are produced in
PYM broth, but they are not real cellulose.
Aerobic. Catalase positive. Oxidase negative. Acetic acid is
produced from ethanol. Acetate is oxidized, but lactate is not or
only weakly oxidized. Dihydroxyacetone is not produced from
glycerol. D-Gluconate is produced from D-glucose, but 2-keto-,
5-keto-, or 2,5-diketo-D-gluconate does not accumulate in culture
media. Acid is produced from L-arabinose, D-xylose, D-ribose,
D-galactose, D-mannose, D-glucose, glycerol, n-propanol, n-butanol, 2-methylpropan-1-ol, ethanol, and methanol, but not from
L-rhamnose, D-fructose, sucrose, maltose, lactose, raffinose, trehalose, D-mannitol, inositol, D-sorbitol, dulcitol, or soluble starch.
Production of acid from D-arabinose, D-ribose, melibiose, and
2-methylpropan-1-ol varies with strains. Methanol, ethanol,
D-glucose, D-mannose, glycerol, and succinic acid are utilized as
sole sources of carbon, but D-fructose, L-arabinose, maltose, trehalose, inositol, and D-mannitol are not utilized. Pantothenic acid
is an essential requirement for growth. Urease negative.
Growth occurs in the presence of 30 % glucose and 0.35 %
acetic acid. Growth occurs at the same extent between pH 3.0
and 8.0; acidotolerant. Growth occurs at 30 C, but not at 45 C.
27
28
Major cellular fatty acids are C18:1 and C16:0, and major
hydroxyl acids are C16:0 2-OH and C16:0 3-OH. Major ubiquinone is Q-10. The G+C content of DNA ranges from 62 to
63 mol%.
The type strain was isolated from a non-sterile fermentation
process for the production of single-cell protein (SCP) from
methanol with Candida species. Acidomonas strains were most
abundantly isolated from activated sludge samples, but not from
vegetables, fruit, decayed wood and leaves, manure, and paddy
soil (Yamashita et al. 2004).
. Table 1.3
Differential characteristics of Asaia species
A. bogorensis
A. lannensis
A. platicodi A. prunellae
A. siamensis
A. spathodea
Flagella
Peritrichous
Lateral
Peritrichous
Peritrichous
Lateral
Lateral
Peritrichous
Peritrichous
Acetic acid
from ethanol
(w)
(w)
Acid from
L-Rhamnose
Dulcitol
+(w)
Growth in the
presence of
0.35 % acetic
acid
Growth on
30 % glucose
Growth at
37 C
+
+
Growth pH 3.0 +
Isoprenoid
quinone
Q-10
Q-10
Q-10
Q-10
Q-10
Q-10
Q-10
+
Q-10
58.9
60.3
60.8
60.0
58.9
59.3
59.7
Symbols: +, positive in most strains; w, weakly positive; v, variable with strains; and , negative in most strains
29
30
31
32
D-galactose, D-glucose, D-mannose, melibiose, D-mannitol, glycerol, and ethanol, but not from lactose or dulcitol. Acid production
varies with species from D-arabinose, L-sorbose, L-rhamnose, maltose, sucrose, raffinose, L-arabitol, meso-erythritol, D-mannitol,
and meso-ribitol. Growth occurs on D-glucose and D-mannitol,
but does not on lactose or maltose. Growth on other sugars and
sugar alcohols differs with the species. Growth on meso-ribitol
and L-arabitol was once used for differentiation of
Gluconobacter oxydans from Gluconobacter cerinus and
Gluconobacter frateurii (Mason and Claus 1989; Katsura et al.
2002; Sievers and Swings 2005d). However, this relation is not
clear because some strains show a weak or very weak reaction
toward one of the polyols. The production of water-soluble pigment is found in the strains of G. oxydans, G. kanchanaburiensis,
G. sphaericus, G. uchimurae, and G. wancherniae. Strains in
several species require nicotinic acid for growth.
Optimum temperature for growth is between 25 C and
30 C. Majority of species grow at 35 C, and a few species
grow at 37 C. Optimum pH for growth is around pH 5.5.
Majority of species grow at pH 3.0 or 3.5. Major cellular fatty
acid is C18:17c. Major ubiquinone is Q-10. The G+C content of
DNA ranges from 54.0 to 61.5 mol%.
Gluconobacter strains are isolated from a variety of
fruits, including kaki (persimmon), jujube, peach, a
variety of citrus fruits, strawberry, cherry, apple, apricot, grape,
fig, etc., and flowers and other sugar-rich environments.
33
34
Gluconobacter thailandicus
Tanasupawat, Thawai, Yukphan,
Moonmangmee, Itoh, Adachi, and Yamada 2004
The characteristics are the same as those described for the
species (Tanasupawat et al. 2004).
Type strain: F-149 T ( BCC 14116T JCM
12310T NBRC 100600T PCU 225T TISTR 1533T), isolated
from a flower of Indian cork tree (Millingtonia hortensis), Bangkok, Thailand. The G+C content of DNA of the type strain is
55.8 mol%.
35
36
37
38
39
40
medium. g-Pyrone compound is weakly produced. Watersoluble pigment is produced. Levan-like polysaccharides are
produced from sucrose.
Growth occurs weakly on 30 % glucose and weakly in
the presence of 0.35 % acetic acid. Growth does not occur on
1.0 % KNO3. Major cellular fatty acid is C18:17c. Major quinone is Q-10. The G+C content of DNA ranges from 68.1 to
69.4 mol%.
Nguyenibacter strains were isolated from the rhizosphere and
roots of Asian rice collected in Vietnam.
41
42
Motility
NA
Thermophilic
Acidophilic
NA
NA
NA
72
Carotenoids
Optimum temperature
Optimal growth pH
Oxidase
Motility
C18:1
Q-10
70.371.0
Oxidase
66.3
Q-9/10, MK-9/10,
RQ-9/10
72.4
Q-10
69.170.4
NA
+
C18:17C
+
C16:0, C19:17c, Cyclo
C19:0 8c
Slightly
alkaliphilic
NA
Neutrophilic
Mesophilic
Q-9
7075
Thermophilic
6573
Q-10
C16:0,
C18:17c
VA
70.2
Q-9
Neutrophilic Neutrophilic
Mesophilic
NA
NA
NA
+
Q-10
58.765.6
C16:0, C18:17c/C18:16C
C18:1
Neutrophilic
Acidophilic
NA
Mesophilic
Mesophilic
Cocci
Belnapia
Rods
Acidocella
Roseomonas Rubritepida
Cocci, short
rods
Mesophilic
69.169.8
Roseococcus
Short rods
Rhodovarius
60.561.9
Q-10
NA
Cyclo C19:0 8c
Q-10
Acidophilic
Mesophilic
Acidophilic
Mesophilic
NA
C18:1
Acidophilic
6370
Q-10
C18:1
Acidophilic
Mesophilic
Neutrophilic
Optimal growth pH
Mesophilic
NA
Bacteriochlorophyll a
Mesophilic
Anaerobic phototrophy
Carotenoids
Red, purple
Optimum temperature
Colonial color
Cocci
Cocci
Cell form
Paracraurococcus Rhodopila
NA
Bacteriochlorophyll a
NA
Anaerobic phototrophy
Acidisphaera
Coccobacilli
Acidisoma
Red, purple
Rods
Acidiphilium
Colonial color
Rods
Cell form
Acidicaldus
. Table 1.4
Characteristics of the genera in the acidophilic group
Mesophilic
NA
NA
NA
NA
NA
Rods
Zavarzinia
68.2
NA
C18:17C
Neutrophilic
Mesophilic
Rods
Humitalea
69.373.5
NA
NA
66.1
Q-10
NA
Neutrophilic Neutrophilic
Mesophilic
NA
NA
Flat star
Stella
70.5
Q-10
C18:1
Neutrophilic
Mesophilic
NA
Cocci
Craurococcus
1
43
44
45
46
47
48
Cells are Gram negative and short rods, 1.01.2 mm long. Long
filaments are formed in liquid culture. Strictly aerobic,
chemoorganotrophic, mesophilic, and neutrophilic. Colonies
are red. Catalase positive. Oxidase positive. BChl is absent, but
carotenoids are present.
The major cellular fatty acids are cyclo C19:08c and cyclo
C17:0. The major quinone is Q-10. The major polar lipids are
phosphatidylglycerol, phosphatidylethanolamine, phosphatidyl
di-methylethanolamine, phosphatidylcholine, and unknown
aminolipids. The G+C content of the genomic DNA is
72.4 mol%. The type species is Rhodovarius lipocyclicus.
49
50
51
52
53
54
Ecology
The genera Acetobacter and Gluconobacter were the core genera
in acetic acid bacteria for a long time. Members of the genus
Acetobacter have niches in alcoholic and acidic environments,
and they are isolated from vinegar, wine, beer, sake, cider, fruits,
and other alcoholic materials. The main substance of vinegar is
acetic acid, and its concentration ranged from approximately
4 % to 6 %. Malt vinegar contains 4.35.9 % of acetic acid, cider
vinegar 3.99.0 %, wine vinegar 4.47.4 %, spirit vinegar
11.512.5 %, rice vinegar 4.005.24 %, and Balsamic vinegar
6.2514.88 % (Adams 1998). By contrast, the members of the
genus Gluconobacter are isolated from sugary materials, fruits,
flowers, foods, and other sources. However, the concept of acetic
acid bacteria has extended to all of the bacteria in the family
Sources of Isolation
Members of the genus Gluconobacter were first isolated from
kaki (Diospyros kaki Thunb.) and other fruits (Asai 1934), and
isolation sources extend to a variety of flowers and fruits (Kondo
and Ameyama 1958). The genus Komagataeibacter is separated
from the genus Gluconacetobacter and has niches in acetous
materials such as a variety of vinegar, kombucha, and fruit
juice. Gluconacetobacter strains were reported to be isolated
from vinegar, fruits, dried fruits, rhizosphere of coffee plants,
pink sugarcane mealybug, and even in a tumulus in Japan.
Asaia strains are widely distributed in such flowers as orchid
tree, plumbago, astilbe, Asian dayflower, heliconia, spider lily,
balloon flower, self-heal, crown flower, ixora, and African tulip
tree. Further, an Asaia strain was isolated from spoiled fruitflavored bottle water (Moore et al. 2002). Acidomonas
methanolica strains were abundantly isolated from activated
sludge samples, but not from vegetables, fruits, decayed wood
and leaves, manure, and paddy soil (Yamashita et al. 2004).
Strains of the genera Ameyamaea, Neoasaia, and
Neokomagataea were isolated from a variety of flowers.
A Tanticharoenia sakaeratensis strain was isolated from soil. An
osmophilic acetic acid bacterium, Saccharibacter floricola, was
isolated from pollens in Japan.
55
56
. Table 1.5
Diversity of acetic acid bacteria relevant to isolation sources
Sources
Genera
Species
Fermented foods
Acetobacter
A. pasteurianus
41
A. orleanensis
16
A. lovaniensis
14
15
A. indonesiensis
14
A. tropicalis
A. syzygii
Gluconobacter
Gluconacetobacter
Others
1
4
Total
26
2
31
27
1
2
15
6
2
9
83
60
157
G. oxydans
21
G. frateurii
12
16
14
46
14
24
23
67
G. hansenii
Gluconactobacter sp.
8
1
A. bogorensis
A. siamensis
Asaia sp.
K. baliensis
Frateuria
F. aurantia
Total
20
49
51
13
13
3
2
Kozakia
6
6
19
Asaia
Flowers
41
A. cibinogensis
A. orientalis
Fruits
65
13
3
0
67
4
16
13
16
118
98
99
16
331
sources of carbon (> Table 1.7). Thus, 345 strains were isolated
from 776 samples consisting of 684 from flowers, 58 from fruits,
and 34 from fermented foods (Suzuki et al. 2008).
The isolates were classified into the genera Acetobacter,
Gluconobacter, Gluconacetobacter, Asaia, Saccharibacter, and
Frateuria based on phenotypic characteristics, 16S rRNA
sequence analysis, and DNA-DNA hybridization (Suzuki et al.
2008). Asaia strains are most abundantly isolated, followed by
Acetobacter strains, Gluconobacter strains, and Gluconacetobacter. A few strains of the genera Saccharibacter and Frateuria
are isolated (> Table 1.10). No Acidomonas strains are found.
The Asaia strains are obtained by the use of ME VI and from
flowers as expected (> Table 1.10). Further, acetic acid bacteria
are distributed evenly across Japan (> Fig. 1.5). In addition,
the strains are isolated from Akita, Niigata, Tokyo, and
Yamanashi located in the temperate zone in Japan. Three new
species of the genus Asaia were identified based on the characterization of the isolates from flowers collected in the northern
area (Suzuki et al. 2010). Consequently, the genus Asaia is not
Insects
Mosquitoes
A mosquito, Anopheles stephensi, is a vector of Plasmodium sp.,
the pathogen of Asian malaria. The total number of cases of
malaria is estimated to be 300500 million in the world, and two
. Table 1.6
Diversity of acetic acid bacteria relevant to country origins
Country
Genera
Acetobacter
Gluconobacter
Gluconacetobacter
Asaia
Indonesia
Thailand
The Philippines
Total
A. pasteurianus
11
21
41
A. orleanensis
11
12
26
A. lovaniensis
27
31
A. indonesiensis
20
27
A. tropicalis
A. syzygii
A. cibinogensis
A. orientalis
15
2
6
2
9
91
20
G. oxydans
17
G. frateurii
35
46
52
12
67
46
157
21
11
G. hansenii
Gluconactobacter sp.
3
20
12
A. bogorensis
25
17
51
A. siamensis
13
Asaia sp.
3
19
11
67
37
Kozakia
K. baliensis
Frateuria
F. aurantia
16
Total
3
4
0
4
16
16
16
208
51
72
331
57
58
. Table 1.7
Enrichment media for acetic acid bacteria
Enrichment media at pH 3.5
Ingredient
DGlucose
ME I
1.0
ME II
ME III
ME IV
ME V
D-Sorbitol
ME VI
0.15
2.0
1.0
D-Mannitol
2.0
Dulcitol
2.0
1.0
Methanol
2.0
Ethanol
0.5
Peptone
1.5
0.5
0.5
0.5
0.5
0.5
Yeast extract
0.8
0.3
0.3
0.3
0.3
0.3
Acetic acid
0.3
Cycloheximide
0.001
0.001
0.001
0.001
0.2
0.01
0.001
. Table 1.8
Diversity of acetic acid bacteria relevant to enrichment media
Enrichment media
Genera
Acetobacter
Gluconobacter
Gluconacetobacter
ME I
Frateuria
ME IV
ME V
Total
A. pasteurianus
23
12
41
21
26
A. lovaniensis
16
A. indonesiensis
15
27
A. tropicalis
12
15
31
A. syzygii
A. cibinogensis
A. orientalis
94
32
33
G. oxydans
11
G. frateurii
10
24
46
21
32
67
157
21
G. hansenii
Gluconactobacter sp.
6
0
20
A. bogorensis
32
17
51
A. siamensis
13
Asaia sp.
0
Kozakia
ME III
A. orleanensis
20
Asaia
ME II
K. baliensis
24
67
4
139
103
38
34
F. aurantia
Total
39
16
16
17
331
. Table 1.9
Habitats of acetic acid bacteria
Genera
Acetobacter
Fermented foods
Flower
Others
A. orleanensis
A. lovaniensis
Palm seed,
coconut
juice
A. indonesiensis
Palm wine
Palm seed
A. tropicalis
Coconut
juice
A. syzygii
Vinegar
Flower
Montana, coconut
Soybean
curd
A. orientalis
tempeh
Canna flower
G. oxydans
Nata de coco
Sapodilla, papaya,
orange, soursop,
mangosteen
G. frateurii
Sapodilla
Asaia
Flowers
A. pasteurianus
A. cibinogensis
Gluconobacter
Fruits
G. hansenii
Nata de coco
Nata de coco
A. bogorensis
Tape ketan
Pineapple
Bauhinia, plumbago,
ixora, lantana, African
tulip, coconut flower,
alamanda
A. siamensis
Asaia sp.
K. baliensis
Frateuria
F. aurantia
Cincau
Gluconactobacter sp.
Kozakia
Soybean
curd
Ragi, palm
brown
sugar
Lime, menteng,
cempedak, kemaris
Coconut flower
59
60
. Table 1.10
Relationships between enrichment media and isolation sources
Enrichment media
Isolation sources
Me I
Me II
Me-VI
Flowers
Fruits
Fermented foods
Total
Acetobacter
56
19
21
24
29
53
96
Gluconobacter
24
22
25
21
46
Gluconacetobacter
25
13
16
18
41
Asaia
38
119
152
157
Saccharibacter
Frateuria
Total
145
22
178
211
63
71
345 strains
Genus
. Fig. 1.5
Distribution of acetic acid bacteria in Japan Suzuki et al. (2008)
74 % of Acinetobacter. The prevalence of the genus Acinetobacter correlated with mosquito gender and the prevalence of the
genus Asaia with the interaction between mosquito gender and
the sampling sites. Male mosquitoes were more infected than
female mosquitoes for each bacterial genus (Minard et al.
2013a). This study emphasizes the need to gain a global view
Flies
Drosophila melanogaster (fruit fly) is widely used in research in
genetics and developmental biology. In view of bacterial community in the fly, adult flies were collected from 11 sites in the
USA, and the DNAs from bacteria were isolated from male flies
at each of the 11 different locations. Bacterial 16S rRNA gene
sequences libraries recovered were compared, and the bacteria
from these sequences were grouped into 74 different taxa, spanning the phyla Proteobacteria, Bacteroidetes, and Firmicutes. In
addition, many groups were found in the genus Gluconacetobacter in the order Alphaproteobacteria (Corby-Harris et al.
2007). However, conflicts are reported on the bacterial taxa,
particularly the genera Gluconobacter and Gluconacetobacter,
which were found in Drosophila samples. The samples were
collected around the world, and after extraction of DNA, the
16S rDNA PCR amplification and sequencing were done. As
a result, the family Acetobacteraceae was not a major family, and
both the groups were not detected in the samples (Chandler
et al. 2011).
Further, commensalism with microorganisms in gut of
the Drosophila has attracted attention to biologists. In the
light of interests, bacteria were examined by DGGE fingerprints
of PCR-amplified 16S rRNA fragments for the midgut of
reared wild-type flies and Cad-RNAi flies that carried the
UAS-Cad-RNAi construct to mimic the loss of function via
RNA interference (RNAi). Lactobacillus plantarum, Lactobacillus
brevis, and Acetobacter pomorum were commonly detected
in both the wild-type flies and Cad-RNAi flies. However,
a bacterium in the family Acetobacteraceae EW911 (A911)
[a dominant commensal member in wild-type flies, controls,
1.5 105 colony-forming units (CFUs) per gut] was
markedly diminished and maintained at very low level in the
Cad-RNAi gut (900 CFUs per gut), whereas Gluconobacter
sp. EW707 (G707) (a minor commensal member in the
control gut, 800 CFUs per gut) emerged as a dominate
commensal in the Cad-RNAi gut (1.7 104 CFUs per gut)
(Ryu et al. 2008).
Olive fruit fly (olive fly, Bactrocera oleae) is one of the
major pests of the olive tree. Wild specimens of B. oleae
Mealybug
Pink sugarcane mealybug (Saccharococcus sacchari) is
common to all countries that grow sugarcane and almost the
extensive species of mealybug found on commercially grown
sugarcane. A small number of acetic acid bacteria were isolated
from pink sugarcane mealybugs and were identified
as Gluconobacter oxydans (Ashbolt and Inkerman 1990).
In addition, Gluconacetobacter sacchari, G. liquefaciens, and
G. diazotrophicus strains were isolated from pink
sugarcane mealybugs collected in Australia and Hawaii (Franke
et al. 1999).
Bee
Honeybees (Apis mellifera) are domesticated bees reared in
apiaries and are used for producing honey and for pollination
commercial crops. Isolation of Gluconobacter strains was
attempted with enrichment approach from not only 411 samples, mostly bees, but also flowers visited bees, honey, and other
materials from the hive and some wasps in three regions in
Belgium. As a result, 53 strains were isolated, and the strains
were studied based on phenotypic characteristics. Further, the
strains were classified into two groups, Group A and Group
B with polyacrylamide gel electrophoresis of soluble cell proteins. As a result, 37 strains in Group A were identified as
Gluconobacter oxydans subsp. suboxydans (now Gluconobacter
oxydans) and 15 strains in Group B as a member of the genus
Gluconobacter (Lambert et al. 1981). Bacteria associated with
worker adults of two subspecies of Apis mellifera in Southern
Africa, Apis mellifera capensis and Apis mellifera scutellata, were
screened by analysis with high-fidelity PCR of 16S rRNA
sequences. As a result, the genus Gluconacetobacter was detected
in the abdomen of the workers with the genera Lactobacillus,
Bifidobacterium, Serratia, and three other genera (Jeyaprakash
et al. 2003).
In addition, the structural diversity of bacteria found in the
gut of three different bee species, Apis mellifera carnica (honeybee), Bombus terrestris (bumblebee), and Osmia bicornis (red
mason bee), was studied for understanding how specific insects
select for their own bacterial communities. The bees were collected at a flowering oilseed rape field for three successive years
61
62
Leafhopper
Flavescence doree (FD) is a grapevine disease that afflicts several
wine production area in Europe. FD is caused by Candidatus
Phytoplasma vitis and the insect vector of a leafhopper,
Scaphoideus titanus, which is widespread in vineyards. Focusing
on Asaia sp. and S. titanus, the association with the bacteria and
the leafhopper was studied in the light of better understanding
of a possible role of the bacteria.
S. titanus adult individuals were collected in vineyards with
heavy symptoms of FD from different area in Piedmont region
in Italy. Further, feeds that were fed on 300 ml of sucrose 5 % in
TE pH 8.0 with or without rifampicin (200 mg ml1) were used
in order to simulate the natural situation of insects feeding on
plant. Quantitative real-time PCR was performed on insects or
feeds DNA with appropriate primers.
Data demonstrated that Asaia sp. was a significant part of the
insect microflora. The distributing pattern of the bacteria inside
the insect body was investigated through the use of Asaia
expressing a green fluorescent protein. Asaia strain SF2.1 tagged
with Gfp was able to colonize efficiently salivary glands and
female and male reproduction organs of S. titanus (Crotti et al.
2008).
Plant Hopper
Granulibacter bethesdensis
Scientists in an American group isolated a novel Gram-negative
rod from cervical and supercervical lymph nodes of a patient
with the chronic granulomatous disease (CGD). The disease is
a rare inherited disease of the phagocyte NADPH oxidase system, which leads to defective production of superoxide and
hydrogen peroxide. Normally, the phagocyte kills bacteria and
fungi ingested by reactive oxygen compounds produced, but the
phagocyte of the patient with the disease is unable to kill the
bacteria and fungi because of deficiency of the phagocyte. Therefore, the patient with the disease suffers from recurrent lifethreatening infection with bacteria and fungi. The scientists
attempted to make clear the pathogenicity of the strains and
introduced the bacteria into mouse models of CGD. As
a consequence, they obtained evidence that the organism causes
the similar disease in the patient, and the organism was reisolated from the mouse. Therefore, Kochs postulates were
fulfilled as a new pathogen (Greenberg et al. 2006a).
Further, this organism was isolated from the same patient three
more times, and a similar Gram-negative rod was isolated from
other two patients. The group was unable to identify the strains by
conventional methods and commercial identification kits at that
time. Therefore, the scientists employed phylogenetic analysis of
16S rRNA sequence, IT region, and RecA protein-encoding gene.
Thus, they identified the bacterium as a new genus and a new
species and named them Granulibacter bethesdensis in the family
Acetobacteraceae (Greenberg et al. 2006b).
The Granulibacter bethesdensis strains have phenotypic characteristics common to acetic acid bacteria, produce acid from
D-glucose and ethanol, and oxidize lactate and acetate weakly.
Further, the strains prefer high concentrations of D-glucose (e.g.,
5 % of D-glucose) and grow on glutamate agar and mannitol agar.
Interestingly, they assimilate methanol (Greenberg et al. 2006b).
Further, genomic sequences revealed genes relevant to methanol
utilization. Therefore, Granulibacter strains have the characteristics of both acetic acid bacteria and facultative methylotrophs
(Greenberg et al. 2007). This study should be emphasized as the
Asaia bogorensis
Scientists at the University of Pennsylvania isolated
a bacterium from the peritoneal dialysis fluid of a patient with
a medical history of end-stage renal disease secondary to diabetes mellitus, but they were unable to identify the organism by
conventional biochemical tests. Therefore, the scientists
performed 16S rRNA gene sequencing of the organism and
identified it as Asaia bogorensis. They concluded that the organism from multiple samples made it unlikely that the organisms
represented a contaminant and may cause peritonitis (Snyder
et al. 2004).
Other case was presented on Asaia bogorensis as well. A man
with a history of intravenous drug abuse was transferred to
University of Helsinki Hospital, Finland, who was suspected to
have systemic infection. A Gram-negative rod was isolated from
his blood and not identified by conventional routine tests and
commercial identification systems. However, this bacterium was
identified as Asaia bogorensis on the basis of 16S rRNA gene
analysis (Tuuminen et al. 2006).
These reports suggest the potential opportunistic pathogens
of Asaia bogorensis. Both groups of Pennsylvania University and
Helsinki University emphasized the usefulness of sequencing of
16S rRNA gene for the identification of unusual bacteria from
the clinical specimen.
Asaia lannensis
Following the isolation of Asaia bogorensis, Gram-negative
bacilli were isolated from a bloodstream infection on a child
with cancer and bone marrow transplantation. However, the
bacteria were unable to be identified with commercial identification systems but were identified as Asaia lannensis according
to 16S rRNA gene sequencing. This would be the first case of
bacteremia caused by Asaia lannensis (Abdel-Haq et al. 2009).
Further, two cases of nosocomial infection with Asaia
lannensis were reported. Case 1 is of a 15-month-old female
and Case 2 is of a 5-year-old female both with congestive heart
failure due to idiopathic dilated cardiomyopathy. In Case 1, the
patient was presumed to have bacteremia. Isolates from blood
cultures of both patients showed the same micro- and macroscopic features. The organisms were strictly aerobic, catalase
positive, and oxidase negative and produced acid from D-xylose,
D-glucose, and D-mannitol. However, three commercial identification systems failed to identify the bacteria, and the bacteria
were identified as Asaia lannensis according to 16S rRNA gene
sequencing (Juretschko et al. 2010).
63
64
Gluconobacter spp.
Members of the genus Gluconobacter are widely distributed in
fruits and flowers. However, several strains were found in human
blood and sputa (Alauzet et al. 2010). A Gram-negative rod was
isolated from peripheral blood samples of a male patient with
a history of intravenous drug abuse. Conventional methods and
commercialized systems failed to identify it. However, the
strain was identified as Gluconobacter sp. by sequencings of 16S
rRNA gene, 16S-13S tRNA gene, and dnak gene. In other case,
four Gram-negative, catalase-positive, and oxidase-negative rods
were isolated from sputa of a boy suffering from cystic fibrosis
(CF). Phenotypic identification was unsuccessful for the strains,
but the strains were identified as Gluconobacter spp. according to
the above molecular means. In addition, a Gram-negative, catalase-positive, and oxidase-negative strain was isolated from the
sputum of a girl with CF. The strain was not identified by
phenotypic methods while identified as Gluconobacter sp. by
the molecular means. In addition, strains of Gluconobacter
spp. were isolated from bloodstream infection associated with
endocardial lesions in a female intravenous drug abuser (Bassetti
et al. 2013). However, the abovementioned Gluconobacter strains
were unable to be identified at the level of species.
Acidomonas methanolica
The genus Acidomonas comprises one species, Acidomonas
methanolica, which is a known methanol-utilizing acetic acid
bacterium. Until now, the strains are most abundantly isolated
from sewage samples (Yamashita et al. 2004). However, an
Acidomonas methanolica strain was isolated from an excised
Roseomonas spp.
The genus Roseomonas includes Gram-negative coccobacilli
characterized by pink-pigmentation and their inability to utilize
methanol as a carbon source (Rihs et al. 1993). Strains of this
genus have since been isolated from human blood, cerebrospinal
fluid, sputum, genitourinary site, and other clinical specimens
(Gilardi and Faur 1984; Wallace et al. 1990; Struthers et al. 1996;
Han et al. 2003). Cases of catheter-related bacteremia due to
Roseomonas gilardii and Roseomonas mucosa were reported
(Alcala et al. 1997; Vasallo et al. 1998; Bard et al. 2010). Further,
blood isolates were obtained from patients with clinically significant underlying disease who had central venous catheters in
place, and the majority was associated with polymicrobial catheter infections (Lewis et al. 1997). Cases of peritonitis caused by
Roseomonas gilardii and Roseomonas fauriae were reported in
patients undergoing continuous ambulatory peritoneal dialysis
(Sandoe et al. 1997; Bibashi et al. 2000). In addition, peritonitis
caused by Roseomonas mucosa was reported in an adolescent
infected with HIV on continuous cycling peritoneal dialysis
(Boyd et al. 2012). Roseomonas species were reported to have
an overall low pathogenic potential for humans, but the species,
in particular R. gilardii, may be significant pathogens in persons
with underlying medical complications (Struthers et al. 1996)
A case of chronic postoperative endophthalmitis due to
Roseomonas spp. was reported, which might result in
misdiagnosis and delayed treatment and caused ocular damage
and severe visual loss. This was related to a case with postoperative endophthalmitis secondary to Roseomonas infection (Chen
et al. 2009). Natural reservoir of Roseomonas strains is not
known. However, the isolates from water were described (Rihs
et al. 1993), and domestic water supplies were noticed for
Roseomonas infection (Sandoe et al. 1997; Bibashi et al. 2000).
In addition, Roseomonas vinacea was isolated from a soil sample
collected from the Qinghai-Tibet Plateau, China (Zhang et al.
2008).
Application
Vinegar
The word vinegar is derived from vin, aigre, and
vinaigre in French, literally meaning sour wine (Kersters
et al. 2006). In the past, wine making and vinegar preparation
were always linked. As early as 4000 B.C., vinegar was already
mentioned in Babylonian writings, as a product of the date
palm, the culture of which was highly developed (Nickol
1979). In Babylon, fruit honey, viz., sugary syrup made from
figs, apricots, grapes, and dates, was used in the kitchen and for
65
66
Alcoholic Beverages
Starting in the Pasteurs time, it was known that acetic acid
bacteria were involved in the acetification of wine (ValleryRadot 1924). The microbiologists at that time already knew
that these bacteria could be isolated from the pellicle that formed
on wine when it was exposed to air. They were also aware of the
fact that the acetification required oxygen (Behrens 1896;
Fuhrmann 1905; Wermischeff 1893). The high alcohol tolerance
of acetic acid bacteria was already observed. Many acetic acid
bacteria were isolated from wine must, but not from either intact
or injured grapes (Fuhrmann 1905). Acetobacter pasteurianus
(75 % of the isolated strains) and A. aceti (19 %) strains were
isolated from wines in southern France (Dupuy 1957). Acetic
acid bacteria were present in about half of the samples of ripe
grapes, and they multiplied on injured grapes (Peynaud and
Domercq 1961).
Gluconobacter strains occurred profusely on ripe grapes of
the Bordeaux region (73 % of all strains isolated), whereas
Acetobacter and Pseudomonas strains constituted, respectively,
only 12 and 15 % of the isolates (Blackwood et al. 1969).
Gluconobacter strains were present on intact and injured grapes
but not during the fermentation. The ketogenesis by
Gluconobacter strains on grapes led to a higher requirement for
SO2 in the sulfitation process. Gluconobacter but no Acetobacter
strains were isolated from dried and mature grapes and vines
(Passmore and Carr 1975). G. oxydans strains were isolated from
grapes (Ameyama 1975). G. oxydans was reported as the dominating acetic acid bacteria in fresh grape must, and A.
pasteurianus and Gluconacetobacter liquefaciens were dominated
during the fermentation process in South Africa (Du Toit and
Lambrechts 2002).
Palm wine is a typical tropical beverage from the fermentation of sugary palm sap and a whitish, effervescent, acidic alcoholic beverage, which is the product of a mixed alcoholic, lactic
acid, and acetic acid fermentation. It is popular beverages in
Africa, South America, and the Far East. As a first step, the sugar
of the sap is fermented to ethanol within 812 h by Zymomonas
strains and yeasts, thus creating a medium highly suitable for the
development of acetic acid bacteria. A rather complex flora was
present in palm wines (Swings and De Ley 1977). During palm
sap fermentation, the acetic acid bacteria appear after 23 days.
Acetic acid bacteria utilizing D-glucose and/or sucrose might be
present in earlier stages of the palm sap fermentation (Okafor
1975). Acetobacter species were isolated from palm wines
(Faparusi 1973; Okafor 1975) and from the immature spadix
of the palm tree (Faparusi 1973). A. pasteurianus, A. lovaniensis,
A. indonesiensis, and A. tropicalis were isolated from palm wine
(Simonart and Laudelout 1951; Lysdiyanti et al. 2003b) and
Gluconacetobacter xylinus from the leaflets of the palm tree and
the surrounding air (Faparusi 1973). Gluconobacter oxydans was
found on the floret of the palm tree, in the tap hole, and in palm
sap (Faparusi 1973, 1974).
Acetic acid bacteria occasionally cause deterioration of alcoholic beverage. Acetobacter pasteurianus strains were isolated
from hiochi sake samples with the smell of acetic acid, applying
Cocoa Fermentations
Cocoa is one of many foods that need microbial fermentation or
microbial curing for flavor development. The main reason for
the fermentation of cocoa is to induce biochemical transformation within the cacao beans that leads to the formation of the
color, the aroma, and the flavor of precursors of cocoa. These
properties are further developed during drying, roasting, and
final processing of the well-fermented cocoa beans to produce
cocoa powder and chocolate (Hansen et al. 1998; Thompson
et al. 2001; Cleenwerck et al. 2007; Romero-Cortes et al. 2012).
Of the microbial flora in the fermentation, yeasts, lactic acid
bacteria, acetic acid bacteria, and spore-forming bacteria were
identified (Schwan and Wheals 2004). During the fermentation
of cocoa beans, yeasts dominate for the first 24 h. Since the pulp
of cocoa beans disappears and acetic acid bacteria start to dominate, the temperature rises to 50 C, and the resulting heat and
the resulting acid are due to chemical reactions in the cocoa
beans known as the microbial fermentation or the microbial
curing (Schwan 1998). The effect of acetic acid produced and
rise in temperature cause the death of the seed embryo as well as
the end of fermentation (Cleenwerck et al. 2007). In the cocoa
fermentation process, Acetobacter ghanensis and Acetobacter
tropicalis strains were reported (Cleenwerck et al. 2007;
Romero-Cortes et al. 2012).
Cocoa wine is made by fermentation of cocoa seeds and is
a popular drink in Nigeria (Bassir 1968; Kersters et al. 2006). The
cocoa beans are ground and boiled in sugared water and allowed
to cool to approximately 50 C, before inoculum is added. The
alcohol concentration of the cocoa wine is 9.511.6 % and is
significantly higher than that of palm wine (Bassir 1968).
Acetobacter and Gluconobacter strains were isolated from cocoa
wine (Bassir 1968). To improve the reliability and quality of the
cocoa fermentation process, a defined and mixed starter culture
including Acetobacter and Gluconobacter strains was successfully
used (Schwan 1998).
Nata
Nata is a white- to creamy-colored gelatinous or agar-like film
grown on surface of juice made from coconut, pineapple, sugarcane, or other fruit wastes in the Philippines, Thailand, Indonesia, and other Southeast Asian countries. Nata de coco is
produced from coconut water or coconut milk. The following
ingredients are added to coconut water: to be final concentrations 2 % of glacial acetic acid; 15 % of sucrose; 0.5 % of
ammonium dihydrogen phosphate, giving pH 5.05.5; and
10 % of 48 h culture inoculum. After fermentation at 2831 C
67
68
. Fig. 1.6
The Reichstein-Grussner process for the production of L-ascorbic acid. The biological oxidation of D-sorbitol to L-sorbose is catalyzed by
G. oxydans. The other processes are no biological steps
. Fig. 1.7
The conversion of D-sorbitol to 2-keto-L-gulonic acid by G. oxydans. The conversion of 2-keto-L-gulonic acid to L-ascorbic acid is a no
biological step to yield vitamin C
cross-linking agents, such as glutaraldehyde and polyethyleneimine on L-sorbose production by immobilized G. oxydans
cells, were studied (Park et al. 1998). The theoretically maximal
productivity of the biological oxidative step was achieved by
using a G. oxydans mutant that was selected under conditions
of substrate inhibition (De Wulf et al. 1996).
Several strategies including recombinant DNA technologies
are being developed to shift the synthesis of ascorbic acid from
chemical procedures to purely bioconversion routes, where the
synthesis of 2-keto-L-gulonic acid is a key intermediate
(Deppenmeier et al. 2002). Some Gluconobacter strains convert
D-sorbitol to 2-keto-L-gulonic acid via L-sorbosone.
The 2-keto-L-gulonic acid can be converted nonbiologically
to L-ascorbic acid (> Fig. 1.7; Hoshino et al. 1990; Saito et al.
1997).
Overexpression of the L-sorbose dehydrogenase and the
L-sorbosone dehydrogenase genes in G. oxydans resulted in
improved yields of 2-keto-L-gulonic acid (Saito et al. 1997). In
this respect, the application of Ketogulonigenium strains belonging to the family Rhodobacteraceae may open new perspectives
for the biological conversion of L-sorbose to 2-keto-L-gulonic
acid (Urbance et al. 2001).
. Fig. 1.8
Synthesis of 1-deoxynojirimycin and miglitol from D-glucose. The conversions of 1-amino-D-sorbitol to 6-amino-L-sorbose and of
N-hydroxyethyl-1-amino-D-sorbitol to N-hydroxyethyl-6-amino-L-sorbose are the regioselective oxidation by a G. oxydans strain. The
other conversion are no biological steps. The first step is a chemical reductive amination of D-glucose and the last step is a chemical
stereoselective ring closure
Gluconobacter as Biosensor
The cells and enzymes of G. oxydans find applications as biosensors for the estimation of the concentrations of various
aldoses, polyalcohols, ethanol, glycerol, etc. (Lusta and
Reshetilov 1998; Macauley et al. 2001; Tkac et al. 2000, 2001).
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