Genus

Proteobacteria/Betaproteobacteria/Nitrosomonadales/Gallionellaceae/

Gallionella
Ehrenberg 1838, 166AL
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Lotta E-L. Hallbeck and Karsten Pedersen, Göteborg University, Department of Cell and Molecular Biology, Medicinare-
gatan 9 C, Box 462, S-405 30 Göteborg, Sweden

Gal.li.o.nel’ la. M.L. dim. ending -ella ; M.L. fem. n. Gal- Gram-negative, bean-shaped cells, usually 0.5–0.8 × 1.6–2.5
lionella named for B. Gaillon, a customs agent and zool- μm, that secrete an extracellular twisted stalk from the con-
ogist (1782–1839) in Dieppe, France. cave side, 0.3–0.5 μm in width and up to 400 μm or more
Gram-negative, bean-shaped cells, usually 0.5–0.8 in length. The stalk is composed of numerous 2 nm-wide
× 1.6–2.5 μm, that secrete an extracellular twisted fibers and is produced under microaerophilic conditions
when cells are in late exponential or stationary growth phase.
stalk from the concave side, 0.3–0.5 μm in width
Motile by means of a polar flagellum. Microaerophilic;
and up to 400 μm or more in length. The stalk is
chemolithotrophic growth can be obtained in vitro using
composed of numerous 2 nm-wide fibers and is pro-
oxygen and ferrous iron concentration gradients in a
duced under microaerophilic conditions when cells salt medium with CO2 as sole carbon source (Table 1).
are in late exponential or stationary growth phase. Mixotrophic metabolism has been demonstrated with glu-
Motile by means of a polar flagellum. Microaerophilic; cose, fructose, and sucrose. Can be found where anaerobic
chemolithotrophic growth can be obtained in vitro groundwater with ferrous iron reaches an oxygen-containing
using oxygen and ferrous iron concentration gradi- environment. Belongs to the Betaproteobacteria, family Gal-
ents in a salt medium with CO2 as sole carbon source lionellaceae, with one known species, G. ferruginea. Most closely
(Table 1). Mixotrophic metabolism has been demon- related species according to 16S rDNA sequence analysis is
strated with glucose, fructose, and sucrose. Can be the chemolithotroph Nitrosospira multiformis, distantly related
found where anaerobic groundwater with ferrous iron with a 16S rDNA sequence similarity of 90%.
reaches an oxygen-containing environment. Belongs The mol% G + C of the DNA is: 51–54.6 (Hanert, 1989).
Type species: Gallionella ferruginea Ehrenberg 1838, 166.
to the Betaproteobacteria, family Gallionellaceae, with
Number of validated species: 1
one known species, G. ferruginea. Most closely related
species according to 16S rDNA sequence analysis
Further descriptive information
is the chemolithotroph Nitrosospira multiformis, dis-
tantly related with a 16S rDNA sequence similarity The 16S rRNA gene of Gallionella ferruginea (strain Johan)
of 90%. has been sequenced between base numbers 47 and 1405
The mol% G + C of the DNA is: 51–54.6 (Hanert, (E. coli numbering) (Hallbeck et al., 1993). Phylogenetic
1989). analysis of these sequence data placed Gallionella among
Type species: Gallionella ferruginea Ehrenberg 1838, the Betaproteobacteria. G. ferruginea is distant from other
166. species in the tree, with a 10% sequence difference com-
.................................................................................. pared to the closest species, the chemolithotroph Nitrosospira

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Bergey’s Manual of Systematics of Archaea and Bacteria, Online © 2015 Bergey’s Manual Trust. This article is © 2005 Bergey’s Manual Trust.
DOI: 10.1002/9781118960608.gbm00988. Published by John Wiley & Sons, Inc., in association with Bergey’s Manual Trust.
 2 Bergey’s Manual of Systematics of Archaea and Bacteria
TABLE 1. Characteristics for identification of the genus
Gallionella
Characteristic
Cell morphology
Cell dimension, μm
Produce an extracellular twisted stalk
composed of numerous 2 nm-wide fibers
Grows with CO2 as sole carbon source
Grows with ferrous iron as energy source
Temperature range (∘ C)
Optimal temperature (∘ C)
pH range
Motile
multiformis. The remote position of G. ferruginea in relation
to other species and its utilization of iron as energy source
and electron donor compared to ammonia for the closest
gene cluster, Nitrosomonadaceae, indicate that a separate
family, Gallionellaceae, is justified. There is only one named
species at this time. The capacity for chemolithotrophic
iron oxidation among bacteria has been suggested to be
evolutionarily widespread (Lane et al., 1992). This is sup-
ported by, for instance, the phylogenetic distance (85% 16S
rRNA gene sequence similarity) between Gallionella and the
iron-oxidizing genus Thiobacillus.
The size, shape, and ultrastructure of Gallionella are illus-
trated in Figures. 1–3, and 4. Cells are curved, bean-shaped,
and may have a polar flagellum (Figure 1). A twisted extra-
cellular stalk is secreted from the concave side of the cell
(Figures. 1 and 3A). It consists of numerous 2 nm-wide
fibers at the point of excretion (Vatter and Wolfe, 1956).
The stalk becomes continuously encrusted with precipitated
ferric iron oxide (Figure 3B), which may totally cover old
stalks. The composition of the stalk has not been conclusively
demonstrated, but inorganic as well as organic composi-
tions have been suggested. Hanert (1989) proposed the
stalk to consist of colloidal ferric hydroxide, based on its
disappearance in 0.12% sodium thioglycolate. Hallbeck
and Pedersen (1995) found a higher carbon-to-nitrogen
ratio (C/N 6.8) in stalk-forming cultures compared to a
non-stalk-forming culture (C/N = 4.3), and concluded the
stalk to be composed of an extracellular carbon skeleton
without a dominating protein component. The cell wall is
Gram-negative (Figure 4D–E).
Two strains of Gallionella have been described (Table 2).
Strain BD from a drainpipe in Braunschweig was described
by Hanert (1989) and strain Johan from a 60 m-deep drink-
ing water well was described by Hallbeck and Pedersen
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FIGURE 1. G. ferruginea strain BD cell morphology and flagella
arrangement in cell suspension after the stalks were dissolved
Gallionella in sodium thioglycolate. Bar = 1 μm.
Bean shaped
0.5–0.8 × 0.8–2.5
+
+
+
5–25
17–20
5.0–6.5
+
(1990, 1991, 1995) and Hallbeck et al. (1993). Strain BD
was reported to have intracytoplasmic membranes (Hanert,
1989) while strain Johan does not (Figure 4). The isolation
procedure for Gallionella is by serial dilution and therefore
the possibility of contaminants in the cultures cannot be
conclusively excluded. Serial thin sectioning should show a
stalk connected to the sectioned cell, as in Figure 4A–C, to
confirm it to be a cell of Gallionella. Inclusions that might
be poly-β-hydroxybutyrate have been noted but apart from
this, strain Johan does not show any specific intracellular fine
structures (Figure 4D–E).
Gallionella can be cultured in vitro in screw-capped test
tubes using oxygen and ferrous iron concentration gradi-
ents in a salt medium with carbon dioxide as sole carbon
source (Figure 5). Solid-phase ferrous iron is placed on the
bottom of the tube in a fresh, autoclaved, and oxygen-free
salt medium. With this procedure, the concentration of
ferrous iron will decrease from the bottom to the top of the
tube as it dissolves and diffuses away from the solid phase.
Oxygen will decrease in concentration from the top of the
tube downwards as it diffuses into the salt medium from
Bergey’s Manual of Systematics of Archaea and Bacteria
FIGURE 2. Apical cell, region of stalk secretion, and ultrastruc-
ture of the stalk of attached G. ferruginea strain BD from the
drain pipe from which it was isolated. Bar = 0.5 μm. (Repro-
duced with permission from H.H. Hanert, Archives of Micro-
biology 60: 348–376, 1968, ©Springer-Verlag, Berlin.)
the air above the medium. The optimal growth temperature
is 17–20∘ C and temperatures above 25–30∘ C are lethal.
Iron sulfide or iron carbonate can be used as a source of
ferrous iron. The culture grows to a maximum of 5 × 106
cells/ml culture (Figure 3C), which makes many traditional
techniques for strain characterization impossible. Strain BD
develops small, circular, colonies attached to the wall of the
tube, 3–5 d after inoculation. Strain Johan predominantly
forms a ring at a specific level in the concentration gradient.
The cells of strain Johan colonize as new rings from upper
levels in the tube as the culture gets old, i.e., 10 days or
more. Eventually, most of the tube will be filled by a brittle
mass of stalks, iron oxides, and cells. Hallbeck and Pedersen
(1990) have demonstrated that Gallionella strain Johan is
free living in vitro in its exponential growth phase and does
not produce stalks until late exponential and stationary
phases. The generation time at optimal temperature for
strain Johan is 8.3 h. Stalk production continues for many
days in stationary phase. Some cell division may still occur but
not at the growth rate observed during the first 4–5 d after
inoculation. The maximum mean stalk length per cell of
strain Johan has been determined to be 60 μm in a 16-day-old
culture. Individual cells may produce much longer stalks;
400 μm has been reported for a single cell of strain BD. With
prolonged subculturing on iron carbonate as energy source,
some of the Gallionella strain Johan cultures were found to
have irreversibly lost their ability to form a stalk (Hallbeck
et al., 1993). Their identity was confirmed by 16S rRNA gene
sequencing. They still form a ring of oxidized iron as the
stalk-forming variant does, but the ring is very thin.
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3
Gallionella is chemolithotrophic with ferrous iron as
energy source and electron donor and with carbon dioxide as
sole carbon source. CO2 fixation by strain Johan was revealed
using hydrogen [14 C]-carbonate (Hallbeck and Pedersen,
1991), while in vivo activity of the Calvin cycle key enzyme,
ribulose bisphosphate-carboxylase, has been reported for
strain BD (Hanert, 1989). Mixotrophic metabolism has been
shown on glucose, fructose, and sucrose. Growth did not
occur on these sugars without ferrous iron and carbon diox-
ide. Ammonium and nitrate can be used as nitrogen sources;
the capacity for nitrogen fixation is unknown.
The environment where stalk-forming Gallionella can
be found, commonly attached to surfaces, is slowly flowing
groundwater that is rich in ferrous iron but has a low organic
carbon content. Typical places to search for Gallionella are in
drainpipes, storage basins for groundwater from deep wells,
in tunnels, and on rock walls with seeping groundwater. A
common feature of these environments is that cold (below
20∘ C), reduced, anaerobic, and ferrous-iron-bearing ground-
water reaches an oxygen-containing atmosphere. Such
environments are suitable for chemolithotrophic growth
with ferrous iron as energy source and electron donor, and
oxygen as electron acceptor. A slow flow, supplying ferrous
iron and possibly carbonate from the groundwater to the
attached cells, seems to be an absolute prerequisite for
growth of Gallionella. In this situation, the stalk may act as a
holdfast and prevent the cells from being washed out to a
more oxidized environment without ferrous iron.
Hallbeck and Pedersen (1995) have demonstrated an
additional function for the stalk. The iron oxidation that
occurs in a typical Gallionella environment can be differ-
entiated into two parts: a) The respiratory iron oxidation
performed by the cells in their energy metabolism and b)
the nonmetabolic iron oxidation induced by the increas-
ing oxygen tension as the anoxic groundwater reaches the
atmosphere. Ferrous iron reacting with oxygen participates
in a chain of reactions yielding highly reactive oxygen such
as perhydroxyl (HO2 ), hydrogen peroxide (H2 O2 ), and the
hydroxyl radical (HO) (Stumm and Morgan, 1996). The sur-
vival of stalk-forming Gallionella strain Johan (Sta+ ) in media
with low and high potential for oxygen radical formation was
compared with a variant of strain Johan that irreversibly lost
the ability to form a stalk (Sta− ) (Hallbeck and Pedersen,
1995). It was found that Gallionella Sta+ survived longer (9
weeks) than Sta− (6 weeks) in cultures with a high potential
for oxygen radical formation. It was therefore suggested that
the stalk of Gallionella protects the cells against the toxic
oxygen species discussed above, by directing the oxidation of
iron to the stalk. This phenomenon could be compared to
 4 Bergey’s Manual of Systematics of Archaea and Bacteria

FIGURE 3. Light microscopy images of cells and stalks of G. ferruginea strain Johan. A, Normarski microscope image of a stalked
cell from an environmental sample. Bar = 10 μm. B, Normarski microscope image of stalks with precipitated iron from a
culture. Bar = 50 μm. C, Fluorescent microscopy image of an acridine orange stained culture, filtered on a 0.2 μm membrane
filter. Note the abundant cells among the stalks. Bar = 50 μm.

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Bergey’s Manual of Systematics of Archaea and Bacteria 5

FIGURE 4. Ultra thin section of G. ferruginea strain Johan in stationary phase. A–C, Serial thin sections of a cell with an adja-
cent stalk. Bars = 0.5 μm. D, Cross section of strain Johan with cell wall outer membrane (see next page). Bar = 0.2 μm. E,
Longitudinal section of strain Johan. Bar = 0.25 μm. (see next page)

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 6 Bergey’s Manual of Systematics of Archaea and Bacteria

FIGURE 4. (Continued)

the action of the protein ferritin, proposed to perform iron
oxidation in both procaryotic and eucaryotic cells (Artymiuk
et al., 1991). It is not known whether the iron oxidation on
the stalk is enzymatic or the stalk acts as a surface catalyst
for the oxidation reaction. Thus, the stalk acts as a holdfast
that allows Gallionella to colonize and survive in an ecological
niche, with high ferrous iron content and some oxygen, that
is unavailable for bacteria without a defense system against
the oxygen radicals formed during inorganic oxidation of
ferrous iron.
Enrichment and isolation procedures
Various media for enrichment and cultivation of Gallionella
have been tested. To create proper growth conditions, fer-
rous iron and CO2 must be in the medium. Lieske (1911)

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Bergey’s Manual of Systematics of Archaea and Bacteria 7

designed a culture medium composed of carbonic water and TABLE 2. Differential characteristics of Gallionella
metallic iron. The use of iron sulfide as a source of reduced ferruginea strains Johan and BD
iron was first suggested by Van Niel (Vatter and Wolfe, 1956). Gallionella ferruginea Gallionella ferruginea
Kucera and Wolfe (1957) published a growth medium com- Characteristic strain Johan strain BD
posed of a salt solution and iron sulfide, which made it pos- Cells bean-shaped + +
sible to obtain pure cultures of Gallionella. The salt solution Cell dimensions 0.5–0.8 × 1.6–2.5 0.5–0.7 × 0.8–1.8
was initially prepared with tap water, because the medium (μm)
lacked a crucial component. This component was later found Temperature range 5–25 nd
to be calcium. Since then the ferrous iron medium has been for growth (∘ C)
widely used with some minor modifications. Wolfe’s modi- Optimal 20 17
fied medium is made as follows: Screw-capped tubes (180 × temperature (∘ C)
16 mm) are filled with 10 ml salt medium consisting of 1.0 g pH range for 5.0–6.5 nd
NH4 Cl, 0.4 g MgSO4 ⋅7H2 O, 0.1 g CaCl2 ⋅2H2 O, 0.05 g K2 HPO4 exponential
growth
and 1 liter double-distilled water. The salt medium is auto-
claved, chilled to 5∘ C and infused with sterile filtered CO2 to Generation time in 8.3 h nd
exponential
pH 4.6–4.8. A ferrous sulfide or ferrous carbonate precipi- growth
tate (0.5 ml) is added slowly to the bottom of the tubes with a
Motility without + +
Pasteur pipette and the tubes are left for four to six hours to stalks
allow gradient conditions to establish before inoculation. The Motility with stalks − −
ferrous sulfide and ferrous carbonate have to be prepared in
Stalks not produced + nd
the laboratory. Ferrous sulfide is prepared by dissolving 7.8 in exponential
g FeSO4 (N6 H4 )2 SO4 (Mohr’s salt) and 4.8 g sodium sulfide growth phases
separately, each in 200 ml boiling, distilled water and subse- Length of stalks average 60/cell up to 400/cell
quently pouring the ferrous solution into the sulfide solution. (μm)
Use two 500 ml beakers, mix with a glass rod. Fill the beaker Maximum cell 5 × 106 nd
up to the top and seal with a rubber stopper to avoid oxidation number in in vitro
culture (cells/ml)
of the ferrous iron. Let the iron sulfide sediment for at least
four hours. Decant and wash with boiling water five times. Colony form in vitro Ring on the tube wall Circular colonies
Centrifuge at high velocity, collect the iron sulfide in small Growth with CO2 as + +
bottles, fill up with water and close with an airtight lid. Ster- sole carbon
source
ilize at 121∘ C for 20 min. Store cool in airtight vials. Ferrous
Growth with ferrous + +
carbonate is prepared by dissolving 3.9 g FeSO4 (NH4 )2 SO4
iron as sole
and 1.0 g anhydrous Na2 CO3 separately, each in 100 ml boil- energy source
ing distilled water, and subsequently pouring the carbonate a For symbols, see standard definitions; nd, not determined.
solution into the ferrous solution, preferably under a nitro-
gen atmosphere. The precipitated ferrous carbonate is then
washed five times with boiling double-distilled water and ster-
of maximally 10−6 (30–50 parallel cultures starting with a
ilized in closed vials at 121∘ C for 20 min. Store cool in airtight
10−4 serial dilution). Five to ten serial transfers, each starting
vials. It is recommended that all solutions used for prepa-
from one colony, are necessary to achieve pure cultures in
ration of the medium and source of ferrous iron are filter
sterilized (0.2 μm) to remove any cells or particles that may this manner. This procedure requires up to 10 weeks but is a
give a background during microscopic counts. very certain method for continually reducing the number of
Isolation by serial dilution and serial transfer has been contaminants and obtaining a pure culture. Purity is checked
applied successfully to Gallionella strains BD and Johan microscopically and by use of a variety of heterotrophic and
(Figure 5). For strain BD, an enrichment test tube with autotrophic media i.e., yeast extract bouillon, nutrient agar,
colonies attached to the wall of the test tube is washed Nitrosomonas medium, and Thiobacillus ferrooxidans medium.
several times (Hanert, 1989). One colony is subsequently Stalk material of strain Johan can be collected, suspended in
suspended in 10 ml fresh sterile medium and inoculated into new medium, diluted and inoculated according to the strain
new test tubes with Wolfe’s modified medium at dilutions BD procedure.

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 8 Bergey’s Manual of Systematics of Archaea and Bacteria
FIGURE 5. Isolation procedure for G. ferruginea strain BD
(arrow, one-colony culture). (Reproduced with permission
from H.H. Hanert, Archives of Microbiology 60: 348–376,
1968, ©Springer-Verlag, Berlin.)
Maintenance procedures
Maintaining pure stock cultures of Gallionella strain BD and
Johan over the past years was performed using the described
culture conditions with serial dilution transfers every four to
eight weeks. Preservation of Gallionella culture material for
at least 13 weeks by freezing at −80∘ C in 15% glycerol has
been reported by Nunley and Krieg (1968). This procedure,
however, did not result in survival of strain Johan.
Taxonomic comments
The genus Gallionella is characterized by its chemolithotrophic
growth with ferrous iron, and its production of a twisted stalk
consisting of a bundle of numerous fibers that makes Gal-
lionella very easy to identify. Gallionella has been described
under several different names such as Spirophyllum ferrugineum
(Ehrenberg, 1836; Adler, 1904), Didymohelix ferrugineum (Grif-
fith, 1853), Gloeosphaera ferruginea (Rabenhorst, 1854), and
Gallionella filamenta (Balashova, 1967a, b). Most, if not all,
of the attempts to categorize species of the genus Gallionella
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under various names arise from observed differences in the
appearance of the stalk. The current phylogenetic position
of the family Gallionellaceae is based on only a single 16S rRNA
gene sequence of Gallionella.
More than 160 years ago, in 1836, Ehrenberg first dis-
covered the stalks of Gallionella when he studied ochre
masses. In this description (Ehrenberg, 1836), Gallionella
was referred to as a fossil infusorian and he called it “die
Eisenochertierchen”, the small iron ochre animal. Haeckel
(1866) presented a phylogenetic tree that for the first time
included the kingdom Monera for unicellular organisms
and Zopf (1879) first included Gallionella with the Bacteria.
Gallionella has been observed and described by many more
scientists since these early days, and there has been contin-
uous discussion about its morphology and physiology, but
most studies have focused on the stalks. Beger and Bring-
mann (1953) and van Iterson (1958) have summarized the
first 100 years of discussion about the intriguing charac-
teristics of Gallionella. Winogradsky (1888, 1922) proposed
an autolithotrophic life for the so-called “iron bacteria”,
including Gallionella. He mentioned Leptothrix, Cladothrix,
and Gallionella as examples of lithotrophic iron bacteria.
Adler (1904) found only small amounts of Gallionella in fresh
water from Karlspader, but when the water was left in bottles
for several days they “ausserordentlich stark vermehrt” i.e.,
they had grown, or more correctly, the stalks had become
elongated. Lieske (1911) succeeded in the cultivation of
Gallionella in carbonic water with metallic iron as ferrous iron
source. In 1924–1929, Cholodny made microscopic studies
on cover slips that he had submerged in habitats of Gallionella
(Cholodny, 1924, 1929). He sketched admirable pictures of
cells attached to the end of stalks, and he showed that the
stalk was excreted by the cell and was not a living part of it.
Teichmann (1935) made cultures according to Lieske (1911)
and found a great number of bean-shaped cells in the fluid.
This observation influenced Pringsheim (1949b) to suggest
that “It is not impossible that motile cells are formed under
certain conditions”, a conclusion in accordance with current
knowledge of free-living cells in exponential growth phase.
Beger and Bringmann (1953) made comparisons between
earlier drawings of the stalk of Gallionella and their own
electron microscopy studies and proposed that the genus
Gallionella consisted of five species. Vatter and Wolfe (1956)
presented electron microscopy images of cells with stalks and
in 1957, Kucera and Wolfe (1957) introduced an excellent
growth medium containing iron sulfide as the source of
ferrous iron. Wolfe could have succeeded in working with
Gallionella but he concluded that “these organisms (iron
bacteria) are too difficult to be profitable” (Wolfe, 1964).
Bergey’s Manual of Systematics of Archaea and Bacteria
In 1958, a thesis on Gallionella was presented by van Iterson
(1958). Excellent electron microscopy images of the organ-
ism were presented and it was suggested that the stalk was
a living part of the organism with sporangia in the form of
membrane sacs on the stalk. Balashova (1967a,1967b, 1968)
and Balashova and Cherni (1970) made electron microscopy
observations of the stalk and concluded that the stalk might
have zoogloeal forms and budding cells on the stalks. Hanert
(1989) made detailed studies on stalk elongation using single
cells and measurements of iron oxidation in both natural
samples and lab cultures. Intracytoplasmic membranes and
evidence for autotrophic growth was presented. In 1990,
Lütters-Czekalla (1990) reported growth of Gallionella BD
with reduced sulfur compounds as electron donor instead
of ferrous iron. This observation remains to be confirmed.
In 1990–1995 Hallbeck and Pedersen (1990, 1991, 1995)
and Hallbeck et al. (1993) reported the 16S rDNA sequence
of Gallionella and showed that Gallionella has a free-living
stage without a stalk in exponential growth phase. They
demonstrated chemolithotrophic growth with CO2 as the
sole carbon source and mixotrophic metabolism. They also
demonstrated an organic composition of the stalk and found
that the stalk was important for survival in the environments
where Gallionella is found. A thesis summarizing these results
was published in 1993 (see further reading).
List of species of the genus Gallionella
Gallionella ferruginea
Ehrenberg 1838, 166AL
...................................................................................
fer.ru.gi’ ne.a. L. fem. adj. ferruginea rust-colored.
Morphology and description the same as those of the
genus. Organisms occur in groundwater habitats, espe-
cially in places where iron-bearing groundwater reaches an
oxygen-containing environment. Microaerophilic, possibly
facultatively anaerobic, chemolithotrophic, and mixotrophic.
Gram negative. G + C buoyant density was calculated by using
a micromethod with novel collimating optics (Hanert 1989).
(E. coli B was used as a reference.)
The mol% G + C of the DNA is: 51–54.6. (Bd).
Type strain: no culture isolated.
GenBank accession number (16S rRNA): L07897.
Acknowledgments
We are grateful to Professor W.G. Ghiorse for many valuable
discussions about Gallionella and its life style and to Professor
Grant Ferris for valuable comments on the manuscript. Our
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9
work with Gallionella was supported by the Swedish Natural
Science Research Council.
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Further reading
Hallbeck, L. 1993. On the biology of the iron-oxidizing and
stalk-forming bacterium Gallionella ferruginea, Thesis,
Göteborg University, Göteborg Sweden.