Genus

Proteobacteria/Deltaproteobacteria/Desulfobacterales/Desulfobacteraceae/

Desulfosarcina
Widdel 1981, 382VP (Effective publication: Widdel 1980, 382)
..........................................................................................................................................................................................
Jan Kuever, Department of Microbiology, Institute for Material Testing, Foundation Institute for Materials Science, D-28199,
Bremen Germany
Fred A. Rainey, Louisiana State University, Department of Biological Sciences, Baton Rouge, LA 70803, USA
Friedrich W. Widdel, Max Planck-Institut fr Marine Mikrobiologie, Abteilung Mikrobiologie, Celsiusstrasse 1, D-28359,
Bremen Germany

De.sul.fo.sarci.na. L. pref. de from; L. n. sulfur sulfur; or for CO2 fixation during autotrophic growth. Media
sarcina, Desulfosarcina sarcina-shaped sulfate reducer. containing a reductant and not less than 10 g/l NaCl
Irregularly shaped cells occurring singly and in and 2 g/l MgCl2 6H2 O are necessary for growth.
large, sarcina-like packets. Cells are 0.81.5 1.57.0 Optimal growth temperature, 2833 C. Thermophilic
m, occurring singly or in pairs. Spore formation is not species have not been described. The optimal pH
observed. Granules of poly--hydroxyalkanoic acid fre- range for growth is 7.27.6. Cells contain desulforu-
quently occur within the cells. Gram negative. Usually bidin as sulfite reductase, and cytochromes of the b and
nonmotile, but cells motile by means of a single polar c type. Colonies in anaerobic agar media are grayish to
flagellum may occur. Strictly anaerobic, having both yellowish, compact, and irregular in shape. Occur in
a respiratory and a fermentative type of metabolism. anoxic part of brackish water and marine habitats.
Chemoorganotrophs or chemoautotrophs, using for- The mol% G + C of the DNA is: 5159.
mate, acetate, propionate, butyrate, higher fatty acids, Type species: Desulfosarcina variabilis Widdel 1981,
other organic acids, alcohols, and benzoate or simi- 382 (Effective publication: Widdel 1980, 383.)
lar aromatic compounds as electron donors and also ..................................................................................
as carbon sources; these compounds are completely Irregularly shaped cells occurring singly and in large,
oxidized to CO2 . Chemoautotrophic growth occurs sarcina-like packets. Cells are 0.81.5 1.57.0 m, occur-
with H2 as the electron donor and CO2 as the carbon ring singly or in pairs. Spore formation is not observed.
source. Sulfate and other oxidized sulfur compounds Granules of poly--hydroxyalkanoic acid frequently occur
within the cells. Gram negative. Usually nonmotile, but cells
serve as terminal electron acceptors and are reduced to
motile by means of a single polar flagellum may occur. Strictly
H2 S. In the absence of an external electron acceptor,
anaerobic, having both a respiratory and a fermentative type
slow growth of the type species occurs by fermentation
of metabolism. Chemoorganotrophs or chemoautotrophs,
of lactate or pyruvate to acetate and propionate. Car- using formate, acetate, propionate, butyrate, higher fatty
bon monoxide dehydrogenase activity is commonly acids, other organic acids, alcohols, and benzoate or sim-
observed, indicating the operation of the anaerobic C1 ilar aromatic compounds as electron donors and also as
pathway (carbon monoxide dehydrogenase pathway, carbon sources; these compounds are completely oxidized
Wood pathway) for complete oxidation of acetyl-CoA, to CO2 . Chemoautotrophic growth occurs with H2 as the

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Bergeys Manual of Systematics of Archaea and Bacteria, Online 2015 Bergeys Manual Trust. This article is 2005 Bergeys Manual Trust.
DOI: 10.1002/9781118960608.gbm01020. Published by John Wiley & Sons, Inc., in association with Bergeys Manual Trust.
 2 Bergeys Manual of Systematics of Archaea and Bacteria

electron donor and CO2 as the carbon source. Sulfate and Repeated subculturing from the culture fluid favors the
other oxidized sulfur compounds serve as terminal electron development of coccoidal or ellipsoidal cells which occur
acceptors and are reduced to H2 S. In the absence of an singly or in pairs (Figure 1A). Single cells of the type strain
external electron acceptor, slow growth of the type species are nonmotile, although some possess a single polar flagel-
occurs by fermentation of lactate or pyruvate to acetate and lum. Motile cells have been observed in enrichment cultures
propionate. Carbon monoxide dehydrogenase activity is of Desulfosarcina.
commonly observed, indicating the operation of the anaero- The temperature range for growth is 1538 C, with opti-
bic C1 pathway (carbon monoxide dehydrogenase pathway, mal growth occurring at 33 C. The pH range for growth is
Wood pathway) for complete oxidation of acetyl-CoA, or 6.79.0, with pH 7.4 being the optimal.
for CO2 fixation during autotrophic growth. Media con- D. variabilis reduces sulfate, sulfite, or thiosulfate to
taining a reductant and not less than 10 g/l NaCl and 2 H2 S. Elemental sulfur and nitrate cannot serve as electron
g/l MgCl2 6H2 O are necessary for growth. Optimal growth acceptors. D. cetonicum can use sulfur as electron acceptor.
temperature, 2833 C. Thermophilic species have not been Chemoautotrophic growth occurs with H2 as the electron
described. The optimal pH range for growth is 7.27.6. Cells donor and CO2 as the carbon source. The following com-
contain desulforubidin as sulfite reductase, and cytochromes pounds serve as both electron donors and organic carbon
of the b and c type. Colonies in anaerobic agar media are sources for chemoorganotrophic growth: formate, propi-
grayish to yellowish, compact, and irregular in shape. Occur onate, butyrate, valerate, 2-methylbutyrate, 3-methylbutyrate,
in anoxic part of brackish water and marine habitats. higher fatty acids up to 14 carbon atoms, lactate, pyruvate,
The mol% G + C of the DNA is: 5159. succinate, fumarate, ethanol, 1-propanol, 1-butanol, ben-
Type species: Desulfosarcina variabilis Widdel 1981, 382 zoate, and phenyl-substituted organic acids. Some species
(Effective publication: Widdel 1980, 383.) can additionally use acetone, toluene, m-cresol, or o-xylene.
Number of validated species: 3 Growth on acetate alone is very slow. All organic substrates
are completely oxidized to CO2 via the anaerobic C1 pathway
Further descriptive information (carbon monoxide dehydrogenase pathway, Wood pathway;
Schauder et al., 1986; Janssen and Schink, 1995a, 1995b;
D. variabilis forms large free-living cells (Figure 1A) and Harms et al. 1999b).
sarcina-like packets with irregular arrangement of cells or In the absence of an external electron acceptor, slow
distorted appearance (Figure 1B). These packets form pellets growth is possible with pyruvate or lactate, which are fer-
in liquid media and layers at the glass wall. Cells liberated mented to acetate and propionate. Fumarate can be slowly
from the packets (e.g., by squeezing) are irregularly shaped. fermented to acetate, propionate, and succinate; it does not

FIGURE 1. (A) Phase-contrast micrograph of Desulfosarcina variabilis, viable single cells from liquid medium. Bar = 10 m; (B)
phasecontrast micrograph of Desulfosarcina variabilis, cell packets from an agar colony. Bar = 10 m.

(A) (B)

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Bergeys Manual of Systematics of Archaea and Bacteria 3

act as terminal electron acceptor for anaerobic respiration. method is similar to that described for the genus Desulfobac-
Sugars are not fermented. ter, except that benzoate medium is added to the molten
Membrane-bound and soluble cytochromes of the b and c concentrated agar solution. Before inoculation of the agar
type have been identified in D. variabilis. Cells contain desul- medium, cell packets of Desulfosarcina should be broken
forubidin as sulfite reductase (Arendsen et al., 1993) and carefully with a homogenizer in medium under anoxic
menaquinone MK-7 as the predominant quinone (Collins conditions. In agar media, Desulfosarcina forms irregularly
and Widdel, 1986). Major cellular fatty acids of Desulfococcus shaped, compact colonies, which can be removed with a
multivorans strains are C15:0 anteiso and C16:0 (Kohring et al., sterile Pasteur pipette.
1994). Desulfosarcina ovata strains may be enriched and isolated
D. variabilis is sensitive toward light and has to be incu- with o-xylene as electron donor and carbon source. Pure
bated in the dark. Diffuse daylight in the laboratory may cultures are obtained by repeated application of the agar
inhibit growth completely. dilution method. The method is similar to that described
Desulfosarcina occurs in anoxic black mud of brackish for Desulfobacula toluolica (see genus Desulfobacula), except
water and marine habitats. that the overlying carrier phase contains 2% (v/v) o-xylene
Strains of Desulfosarcina may be cultured in a defined (Harms et al., 1999b).
medium1 with sulfate as electron acceptor and benzoate
as the electron donor and carbon source. For D. cetonicum, Maintenance procedures
brackish medium should be used. Ammonium ions are used
as the nitrogen source. For growth on benzoate, addition Pure cultures are maintained in liquid medium in screw-
of molybdate as a trace element is necessary. It is not yet capped bottles or in anoxically gassed, butyl rubber-sealed
known whether selenite is required, but selenite is added bottles, with storage at 25 C. Stock cultures are transferred
routinely to the medium. The type strains of D. variabilis and every 23 months. Desulfosarcina strains may be preserved
D. cetonicum do not require vitamins; other strains have not indefinitely by suspending the cells in anaerobic medium
been studied in this regard. When inoculation is done from containing 5% dimethylsulfoxide and storing in liquid
old cultures, initiation of growth is favored by the addition of nitrogen.
1030 mg sodium dithionite per liter of medium as a further
strong reductant, in addition to routinely added sodium Differentiation of the genus Desulfosarcina from
sulfide. other genera

Enrichment and isolation procedures Desulfosarcina differs from many other anaerobic sulfate-
reducing bacteria by its formation of dense sarcina-like cell
For the selective enrichment of Desulfosarcina strains, packets. A characteristic physiological property of Desulfos-
benzoate-sulfate medium is used. Other aromatic com- arcina is its growth with benzoate and phenyl-substituted
pounds or acetone will favor growth of species other than the fatty acids. This property is also shared by members of the
type species, D. variabilis. Bottles or tubes (100-, 50-, or 20-ml genera Desulfococcus and Desulfobacterium; however, species of
capacity) can serve as culture vessels and are kept anoxic the latter genera are coccoid or small rods under all cultural
with tight screw caps (if completely filled), or by providing conditions and never form cell packets.
an anoxic headspace of N2 /CO2 (90:10) and sealing with
butyl rubber stoppers. The medium is inoculated with black Taxonomic comments
anoxic mud from brackish or seawater sediments; 25%
of the total volume should be added. Enrichment cultures The closest relatives of the genus Desulfosarcina are members
are incubated at 2530 C and mixed by shaking every of the genus Desulfonema, and Desulfococcus multivorans. A
second day. After intense formation of H2 S has occurred signature sequence for all described Desulfosarcina species
(generally after 1020 d), transfers to fresh medium are except D. cetonicum is 5 -AGGCCACCCTTGATCCAA-3
made. Enrichment cultures should be mixed well before (oligonucleotide probe DSC193), which binds to the 16S
subculturing. rRNA corresponding to position 5 -193210-3 of the
Pure cultures of Desulfosarcina strains are obtained by Escherichia coli 16S rRNA sequence (Ravenschlag et al.,
repeated application of the agar dilution method. The 2000).

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 4 Bergeys Manual of Systematics of Archaea and Bacteria

Desulfosarcina ovata
List of species of the genus Desulfosarcina sp. nov.
...................................................................................
Desulfosarcina variabilis
ova.ta. M.L. adj. ovatus from L. n. ovum egg; L. fem. adj. ovata
Widdel 1981, 382VP (Effective publication: Widdel
1980, 383.) referring to the oval (egg-like) cell shape.
................................................................................... Characteristics are as described in Table 1. The type strain
va.ri.abi.lis. L. adj. variabilis changeable, variable. was isolated from a North Sea oil tank in Wilhelmshaven (Ger-
The morphology is as depicted in Figure 1A and B. Char- many) with o-xylene as sole organic substrate.
acteristics are as described for the genus and listed in Table 1. The mol% G + C of the DNA is: 51.0 (Tm ).
Occurs in anoxic mud of brackish water and marine habitats. Type strain: oXyS1, DSM 13228, JCM 12297.
The mol% G + C of the DNA is: 51.2 (Tm ). GenBank accession number (16S rRNA): Y17286.
Type strain: 3be13, Montpellier, DSM 2060.
GenBank accession number (16S rRNA): M34407, M26632.
End note

Desulfosarcina cetonicum 1. The defined medium is prepared as described in the foot-

comb. nov. (Desulfobacterium cetonicum Galushko and note of the section on the genus Desulfobacter. The follow-
Rozanova 1994, 370.) ing additions are recommended (g/l): NaCl, 13.0 and
...................................................................................
MgCl2 6H2 O, 2.0. After autoclaving of the medium, the
ceto.ni.cum. L. adj. cetonicum pertaining to ketones; Desul- following additional components are added aseptically
fosarcina cetonicum a sulfate-reducing bacterium oxidizing from sterile stock solutions (per liter of medium): sodium
ketones. benzoate solution (150 g/l), 5.0 ml; sodium selenite solu-
Characteristics are as described in Table 1. The type strain tion (Na2 SeO3 5H2 O, 3 mg/l and NaOH, 0.5 g/l), 1.0 ml.
was isolated from flooded oil deposits of the Apsheron penin- The pH of the medium is adjusted to 7.37.5.
sula (Azerbaijan) using butyrate.
The mol% G + C of the DNA is: 59.0 (Tm ).
Type strain: 480, DSM 7267, JCM 12296, VKM B-1975.
GenBank accession number (16S rRNA): AJ237603.

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Bergeys Manual of Systematics of Archaea and Bacteria 5

TABLE 1. Morphological and physiological characteristics of described species of the genus

Desulfosarcinaa

Species Desulfosarcina variabilis Desulfosarcina cetonicumb Desulfosarcina ovatab

Morphology Oval rod, packages Oval Rod
Cell size (m) 11.5 1.52.5 0.81.2 1.82.7 0.81.0 2.54.0
Motility (sp) nr
Mol% G + C content 51 59 51
Sulfite reductase DR nr nr
Major menaquinone MK-7 nr nr
Optimal temperature 33 30 32
( C)
Oxidation of substrate C C C
Electron donors used for
sulfate reduction:
H2 +c nr nr
Formate +c +c +
Acetate (+) + +
Fatty acids: C atoms 314 316 4nr
Isobutyrate nr nr
2-Methylbutyrate (+) nr nr
3-Methylbutyrate (+) nr nr
Ethanol + + +
Lactate + + +
Pyruvate + + +
Fumarate + nr
Succinate + +
Malate +
Benzoate + + +
Acetone nr + nr
1Butanol + + nr
m-Cresol nr +
Phenylacetate + nr nr
1Propanol + + nr
Toluene nr + +
o-Xylene nr nr +
Fermentative growth on:
Fumarate + nr nr
Lactate + nr nr
Pyruvate + nr nr
Compounds used as
elelectron acceptors:
Sulfate + + +

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 6 Bergeys Manual of Systematics of Archaea and Bacteria

TABLE 1. (Continued)

Species Desulfosarcina variabilis Desulfosarcina cetonicumb Desulfosarcina ovatab

Sulfite + nr nr
Sulfur + nr
Thiosulfate + + nr
Nitrate nr
Other electron nr nr nr
acceptors
Growth factor nr
requirement
NaCl requirement 15 nrd
(g/l)
Literature Widdel, 1980 Galushko and Rozanova, 1991; Mueller et al., 1999 Harms et al., 1999b
a Symbols: sp, single polar flagellum; nr, not reported; C, complete oxidation; +, good growth; (+), poor growth; , no growth;
DR, desulforubidin.
b So far not a validated species.
c Autotrophic growth.
d So far only grown in marine medium.

bacterium, Desulfobacterium cetonicum. Arch. Microbiol. 163:

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publication of new names and new combinations previ- Anaerobic degradation of m-cresol by Desulfobacterium
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Harms, G., K. Zengler, R. Rabus, F. Aeckersberg, D. Minz, Ravenschlag, K., K. Sahm, C. Knoblauch, B.B. Jorgensen
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Janssen, P.H. and B. Schink. 1995a. Metabolic pathways Schauder, R., B. Eikmanns, R.K. Thauer, F. Widdel and
and energetics of the acetone-oxidizing, sulfate-reducing G. Fuchs. 1986. Acetate oxidation of carbon dioxide

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This article is 2005 Bergeys Manual Trust. Published by John Wiley & Sons, Inc., in association with Bergeys Manual Trust.
Bergeys Manual of Systematics of Archaea and Bacteria 7

in anaerobic bacteria via a novel pathway not involving from saline environments. Description of Desulfobacter
reactions of the citric acid cycle. Arch. Microbiol. 145: postgatei gen. nov., sp. nov. Arch. Microbiol. 129: 395400.
162172.

Widdel, F. 1980. Anaerober Abbau von Fettsuren und Further reading

Benzoesure durch neu Isolierte Arten Sulfat-reduzierender Bak-
terien, Universitt zu Gttingen, Lindhorst/Schaumburg-
Galushko, A.S. and E.P. Rosanova. 1991. Desulfobacterium
Lippe Gttingen.
cetonicum spec. nov., a sulfate-reducing bacterium oxidiz-
Widdel, F. and N. Pfennig. 1981. Studies on dissimilatory ing fatty acids and ketones. Mikrobiologiya 60: 102107.
sulfate-reducing bacteria that decompose fatty acids. I.
Isolation of new sulfate-reducing bacteria with acetate

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This article is 2005 Bergeys Manual Trust. Published by John Wiley & Sons, Inc., in association with Bergeys Manual Trust.