ELSEVIER Sedimentary Geology 126 (1999) 9–23

Ca-carbonates precipitation and limestone genesis

— the microbiogeologist point of view
Sabine Castanier Ł , Gaële Le Métayer-Levrel, Jean-Pierre Perthuisot
Laboratoire de Biogéologie et Microbiogéologie, Université de Nantes, 2 Rue de la Houssinière,
F-44072 Nantes Cédex 03, France
Received 9 March 1998; accepted 3 March 1999

Abstract

Experiments show that the production of carbonate particles by heterotrophic bacteria follows different ways. In
heterotrophy, the passive carbonatogenesis is generated by modifications of the medium that lead to the accumulation of
carbonate and bicarbonate ions and to the precipitation of solid particles. It is induced by several metabolic pathways of the
nitrogen cycle (ammonification of amino-acids, degradation of urea and uric acid, dissimilatory reduction of nitrates) and
of the sulphur cycle (dissimilatory reduction of sulphates). The active carbonatogenesis is independent of the mentioned
metabolic pathways. The carbonate particles are produced by ionic exchanges through the cell membrane following still
poorly known mechanisms. In autotrophy, non-methylotrophic methanogenesis and cyanobacterial photosynthesis also may
contribute to the precipitation of carbonates (autotrophic carbonates). As carbonatogenesis is neither restricted to particular
taxonomic groups of bacteria nor to specific environments, it has been an ubiquitous phenomenon since Precambrian
times. Carbonatogenesis is the response of heterotrophic bacterial communities to an enrichment of the milieu in organic
matter. After a phase of latency, there is an exponential increase of bacterial numbers together with the accumulation of
metabolic end-products. These induce a pH increase and an accumulation of carbonate and hydrogenocarbonate ions in the
medium. This phase ends into a steady state when most part of the initial enrichment is consumed and there is a balance
between death and growth in bacterial populations. Particulate carbonatogenesis occurs during the exponential phase and
ends more or less after the beginning of the steady state. The active carbonatogenesis seems to start first and to be followed
by the passive one which induces the growth of initially produced particles. In eutrophic conditions, the first solid products
are patches that appear on the surface of the bacterial bodies and coalesce until forming a rigid coating and=or particles
excreted from the cell. All these tiny particles assemble into biomineral aggregates which often display ‘precrystalline’
structures. These aggregates grow and form biocrystalline build-ups which progressively display more crystalline structures
with growth. In oligotrophic conditions, the primary solid products are rapidly smoothed in the crystalline structure and
leave no trace. In present aqueous environments, apart from deep ocean, the potential efficiency of heterotrophic bacterial
carbonatogenesis in Ca-carbonate sedimentation is much higher than autotrophic or abiotic processes. It much more likely
accounts for extensive apparently abiotic limestone formation than any of the latter. As far as biodetrital particles are
concerned, it may be observed that the shells and tests of organisms are built from the activity of cellular organites which
are nowadays considered by a number of biologists as endosymbiotic bacteria. Thus, apart from (probably mythical) purely

Ł Corresponding author. Fax: C33 2 51 12 54 63. E-mail: castznie@chimie.univ-nantes.fr

0037-0738/99/$ – see front matter  1999 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 7 - 0 7 3 8 ( 9 9 ) 0 0 0 2 8 - 7
10 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23

evaporitic and autotrophic ones, most limestones must be considered as principally of heterotrophic bacterial origin. As
the carbon of limestones is issued from organic matter, bacterial heterotrophic carbonatogenesis appears as a fundamental
phenomenon in the relationships between atmosphere and lithosphere during the biogeological evolution of the Earth.
 1999 Elsevier Science B.V. All rights reserved.

Keywords: biomineralisation; bacteria; experiments; metabolism; limestones

1. Introduction are present in the medium, such a depletion favours

In nature carbonate precipitation may theoreti-
cally occur following several known processes: (i)
abiotic chemical precipitation from saturated solu-
tions by evaporation, temperature increase and=or
pressure decrease; (ii) external or internal skele-
ton production by eukaryotes; (iii) lowering of CO2
pressure under effect of autotrophic processes (pho-
tosynthesis, methanogenesis); (iv) fungal mediation
(Callot et al., 1985; Verrecchia and Loisy, 1997);
(v) heterotrophic bacterial mediation. As a matter
of fact, bacterial contribution to limestone formation
has been suspected for years (Drew, 1910a,b; Keller-
man, 1915; Berkeley, 1919; Lipmann, 1924; Molish,
1924; Nadson, 1928; Krumbein, 1968, 1974, 1978)
but remained controversial until recent experiments
in microbiogeological laboratories investigated the
metabolic pathways involved, the modes and condi-
tions of solid particles formation, and evaluated bac-
terial carbonate productivity (Krumbein, 1979a,b;
Castanier, 1987; Riege et al., 1991; Le Métayer-
Levrel, 1996; Castanier et al., 1997, 1999).
2. The metabolic pathways of bacterial
Ca-carbonate formation
The production of Ca-carbonate particles through
bacterial mediation follows different ways.
2.1. Autotrophic pathways
In autotrophy, three metabolic pathways are in-
volved: non-methylotrophic methanogenesis (Marty,
1983), anoxygenic photosynthesis and oxygenic pho-
tosynthesis. All three pathways use CO2 as carbon
source to produce organic matter. Thus, they induce
CO2 depletion of the medium or of the immedi-
ate environment of the bacteria. When calcium ions
calcium-carbonate precipitation (Fig. 1).
2.2. Heterotrophic pathways
In heterotrophy two bacterial processes may oc-
cur, often concurrently.
2.2.1. Passive precipitation
Passive precipitation or passive carbonatogenesis
operates by producing carbonate and bicarbonate
ions and inducing various chemical modifications in
the medium that lead to the precipitation of calcium
carbonate. Two metabolic cycles can be involved: the
nitrogen cycle and the sulphur cycle.
In the nitrogen cycle, passive bacterial precipi-
tation follows three different pathways: (i) the am-
monification of amino-acids in aerobiosis (i.e. in the
presence of gaseous or dissolved oxygen), in the
presence of organic matter and calcium); (ii) the dis-
similatory reduction of nitrate (in anaerobiosis (i.e.
in the absence of oxygen) or microaerophily (i.e. in
the presence of very low amounts of oxygen), in
the presence of organic matter, calcium and nitrate);
and (iii) the degradation of urea or uric acid (in
aerobiosis, in the presence of organic matter, cal-
cium, and urea or uric acid). Both urea and uric acid
result from eukaryotic activity, notably that of ver-
tebrates. These three pathways induce production of
carbonate and bicarbonate ions and, as a metabolic
end-product, ammonia, which induces pH increase
(Fig. 2). When the HC concentration decreases, the
carbonate–bicarbonate equilibria are shifted towards
the production of CO23 ions. If calcium ions are
present, calcium-carbonate precipitation occurs. If
Ca2C (and=or divalent cations) are lacking in the
medium, carbonate and bicarbonate ions accumulate,
and the pH increase and bacterial activity may favour
zeolite formation. This happens in soda lakes, e.g. in
Kenya (Castanier et al., 1993).
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23
Fig. 1. Bacterial calcium-carbonate production in autotrophy.
In the sulphur cycle, bacteria use a single metabolic
pathway: the dissimilatory reduction of sulphate
(Fig. 3). The environment must be anoxic, and rich
in organic matter, calcium and sulphate. Using this
pathway, bacteria produce carbonate, bicarbonate ions
and hydrogen sulphide. If calcium ions are present,
the precipitation of Ca-carbonates depends on the hy-
drogen sulphide behaviour. If the hydrogen sulphide
degasses, this induces pH increase and, Ca-carbonate
precipitation. On the other hand, hydrogen sulphide
may be used by other bacteria. If anoxygenogenic
sulphide phototrophic bacteria are involved, the hy-
drogen sulphide is oxidised into sulphur which forms
intra-cellular or extra-cellular deposits. Hydrogen sul-
phide up-take induces pH increase favouring calcium-
carbonate precipitation. If autotrophic sulphide-oxi-
dising aerobic bacteria are involved, they produce sul-
phate ions. Together with hydrogen ions from water
this gives sulphuric acid, the pH decreases and no
solid Ca-carbonate appears. If hydrogen sulphide is
11
neither used by bacteria nor discharged, pH decreases
and Ca-carbonates cannot precipitate.
2.2.2. Active precipitation
Active precipitation or active carbonatogenesis
is independent of the other previously mentioned
metabolic pathways. The carbonate particles are pro-
duced by ionic exchanges through the cell membrane
by activation of calcium and=or magnesium ionic
pumps or channels, probably coupled with carbonate
ion production. Numerous bacterial groups are able
to operate such processes.
In all experiments carbonatogenesis appears to be
the response of heterotrophic bacterial communities
to an enrichment of the milieu in organic matter. Af-
ter a phase of latency, there is an exponential increase
of bacterial strengths together with the accumulation
of metabolic end-products. These induce an accumu-
lation of carbonate and hydrogenocarbonate ions in
the medium and, by different ways, a pH increase
12 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23
Fig. 2. Passive bacterial precipitation of calcium carbonate in the nitrogen cycle.
that favours carbonate precipitation. This phase ends
into a steady state when most part of the initial en-
richment is consumed. Particulate carbonatogenesis
occurs during the exponential phase and ends more
or less after the beginning of the steady state. In most
cases, the active carbonatogenesis seems to start first
and to be followed by the passive one which in-
duces the growth and shape modifications of initially
produced particles.
3. Relationships between bacteria, minerals and
environmental conditions
3.1. In situ experiments with eutrophication
The solid products of bacterial carbonatogenesis
were first studied in eutrophicated karstic originated
waters from a natural pool situated at Les Cugnes
near Les Eyzies (Castanier, 1987). They were col-
lected on glass plates suspended directly in the natu-
ral environment and in jars containing natural water
(Figs. 4–7). Both received nutritional amendments in
order to intensify the several metabolic pathways of
bacterial carbonatogenesis.
The first solid products are probably amorphous
and perhaps hydrated at the beginning (Castanier et
al., 1988). They appear on the surface of the bacterial
bodies as patches or stripes that extend and coalesce
until forming a rigid coating (cocoon) (Fig. 4). In
other cases, solid particles formed inside the cellu-
lar body, are excreted from the cell (excretates). All
these tiny particles, including more or less calcified
bacterial cells, assemble into biomineral aggregates
which often display ‘precrystalline’ or rather ‘pro-
crystalline’ structures (Fig. 5). Sometimes the bacte-
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23 13

Fig. 3. Passive bacterial precipitation of calcium carbonate in the sulphur cycle.

rial bodies themselves present an angular crystalline
shape as if they would contain a growing single crys-
tal (Fig. 4, 4). Such a feature could be compared
with the cubic-shaped bacteria that are found in hy-
persaline environments (Gerdes et al., 1985). At this
stage of evolution and in numerous cases, the calcified
bacterial cells tend to arrange themselves into nearly
crystalline structures (Fig. 5, 5) and sometimes into
dendritic or fibroradial fabrics (Fig. 6) the latter being
considered as possibly a precursor to ooids (Castanier
et al., 1989). The angles of such structures are gener-
ally close to but neither exactly those of the rhombo-
hedral system nor those of the orthorhombic one as if
bacteria dislike the crystallographic physical rules.
The primary aggregates grow and form secondary
biocrystalline assemblages or build-ups which pro-
gressively display more crystalline structures with
growth (Fig. 7). Tetrahedral assemblages and pentag-
onal faces are often observed. This phase should cor-
respond to the passive carbonatogenesis. Different
types of crystallogenetic sequences seem to follow
the different metabolic pathways (Castanier, 1987).
This has to be explored further but if it proves true,
it could lead to the reconstitution of palaeomicro-
biotopes from the examination of carbonate particles
as long as they are not disturbed or changed by
diagenesis.
14 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23 15

3.2. Observations of karstic helictite
Bacterial calcite from a helictite (Collett, 1878)
of Clamouse Cave, formed in an oligotrophic envi-
ronment, was also investigated (Le Métayer-Levrel,
1996; Le Métayer-Levrel et al., 1997). Two micro-
organisms are present (Fig. 8). The first one exhibits
chains of spheres budding or ellipsoids ca. 0.5 to 1
µm in diameter. Cells are usually covered by a thick
carbonate cocoon. From its morphological charac-
ters, this form should be attributed to the genus
‘Isosphaera’ which is the only budding, non-photo-
synthetic bacterium forming chains of spherical cells
(Chafetz and Meredith, 1983; Giovannoni and Cas-
tenholz, 1989). It is a gliding bacterium. Here how-
ever, it could be another species than the only known
species ‘Isosphaera pallida’ Giovannoni, Schabtach
and Castenholz, which lives naturally in hot springs
and in the laboratory within a temperature range of
40–55ºC.
The second organism exhibits more or less anas-
tomosed ribbons 1 µm wide, wearing very numerous
finger-shaped refringent, strongly calcified excres-
cences 0.1 to 0.3 µm wide. This organism could
be the mobile gliding species Vitreoscilla filiformis
Strohl which grows better in nutrients-poor than in
nutrients-rich milieus (Strohl, 1989). Such a species
is undoubtedly well adapted to the nutritional envi-
ronment of caves.
Collected images show remarkable relationships
between these two organisms and carbonate crystals
the outer shapes of which are generally non-rhombo-
hedral (Fig. 8). The ‘Isosphaera’ chains, when they
are short and constituted of probably young ellip-
soidal cells, project perpendicularly from the crystal
faces and are often situated parallel to a crystalline
plan. They seem either to spring out of automor-
phous cavities of the crystalline structure or to rise
directly from the crystalline mass. In this case, a pad
ringing round the base of the filament is often ob-
served at the surface of the crystal. This ring is usu-
ally irregular but sometimes forms a well rounded
areola that connects regularly with the crystalline
surface. This suggests that this pad is a carbonate de-
posit, perhaps amorphous, due to bacterial activity,
which is progressively but rapidly incorporated into
the carbonate crystal.
The Vitreoscilla filaments flatten against crystals
faces and are gradually incorporated into the crys-
tal masses. The filaments and notably the calcified
excrescences are rapidly smoothed in the crystalline
structure. Images show that the quasi-totality of the
mass of crystals is from bacterial origin.
Surfaces of unetched carbonate crystals do not
show clear relationships between the crystalline
structures and the orientation of the filaments. How-
ever, after partial dissolution of crystals with EDTA,
SEM imagery clearly shows that the filaments act as
axes of nucleation to series of piled calcite rhom-
bohedra. When the crystals have almost totally been
removed by dissolution, there remains a network of
more or less calcified filaments.
These observations and experiments show that in
both nutritional conditions bacteria play a major role
in crystallisation, both in supplying carbonate matter
and in its physical structure (Perthuisot et al., 1997).
After eutrophication, bacterial activity is very high
at the beginning and early solid products, as well as
biomineral aggregates, have poorly defined crystal
structure which is overwhelmed by biological luxuri-
ant processes. For example, such conditions are suit-
able for the formation of round bodies with a radial

Fig. 4. First stages of Ca-carbonate bacterial precipitation in eutrophic conditions. (1) Bacterial cells with carbonate pimples (white
arrows) and surrounding excretates (black arrow). Carbonate pimples cannot be confused with endospores because they appear on
dividing cells (1). Carbonate pimples are sometimes located at one tip (2) or both tips (3) of the cells. Pimples form sometimes in the
central zone of the cell against the inner part of the membrane (4). (2) Rods and cocci totally covered by carbonate (black arrows). The
black triangle points at a rod which bears a tail made of excretates and a lateral pimple. The white arrow points at a ghost of bacterial rod
which was destroyed by SEM preparation and which appears as a print into its produced carbonate material. (3) Cocooned rods (a couple
of which is issued from a recent division) and surrounding excretates (white arrow) on a background of weakly calcified bacteria. (4)
Carbonated bacterial bodies with hexagonal crystalline shapes (white arrows) on a background of poorly calcified bacterial bodies with
excretates. The black arrow points at a calcified dense cluster of bacterial bodies. (5) Couple of young bacterial cells. The crack of the
upper one (black arrow) shows the rigidity of the carbonate cocoon. (6) Part of a colony of cocooned cells showing the pores (arrows)
which maintain nutritional and gaseous exchanges between cells in spite of the cocoons. A broken cocoon let appear the inside carbonate
products.
16 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23

Fig. 5. Early biomineral aggregates in eutrophic conditions. (1) Triangular loose aggregate composed of more or less calcified bacterial
cells cemented by aggregates. White arrow points at a heavily cocooned cell. (2) Tiny aggregate composed of a few heavily carbonated
bacterial cells arranged in an angular particle possibly prefiguring a rhombohedron. (3) Angular biomineral aggregates emerging from
a bacterial colony. Some of them (left down) display nearly rhombohedral shapes. Notice divisions of background bacterial cells. (4)
Tiny cluster displaying a subhexagonal arrangement. (5) Biocrystalline network composed of carbonated rods arranged following three
directions. The left outer rim of the structure is more or less cemented (white arrow). While developed on a glass plate, this figure recalls
some palissadic calcite cements in limestones.

internal structure, i.e. ooids (Castanier et al., 1989). rules soon overcome the biological primary disorder.
On the contrary, in oligotrophic conditions, bacterial Thus, observations of recently formed or unmodified
production rate is low so that the crystallographic carbonate bacterial grains could give information on
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23 17

Fig. 6. Early original biomineral aggregates in eutrophic conditions. (1) Fibroradial fabric growing laterally as well as perpendicularly
to the glass plate around a single axis. Carbonate needles are composed of carbonated bacterial cells. Such a fabric recalls the internal
structure of ooids. (2) Dendritic fabric constituted by calcified bacteria among which vibrioid forms.

their original nutritional microenvironment. Besides,
they should give indications on the used metabolic
pathways used by bacteria in eutrophic conditions.
4. Heterotrophic bacterial productivity and
geological implications.
Quantitatively, the production of solid carbonate
depends essentially upon the strains in the bacte-
rial population, the environmental conditions (tem-
perature, salinity, etc.), the quality and quantity of
available nutrients, and time. In the laboratory, all
heterotrophic metabolic pathways were tested; and a
number of experiments were carried out with Bacil-
lus cereus, which is a heterotrophic bacterium able
to undertake the ammonification of amino-acids and
dissimilatory nitrate reduction. In the experiment
presented in Fig. 9, the medium initially contains 4
g organic matter per litre (Castanier, 1984; Le Mé-
tayer-Levrel, 1996). As usual, the phase of latency
is followed by an exponential increase in bacte-
rial growth together with the accumulation in the
medium of metabolic end-products: carbonate, bi-
carbonate and ammonia ions. This phase is followed
by the steady state. Ca-carbonate precipitation oc-
curs during the exponential phase and ends more or
less after the beginning of the steady state.
In the above-mentioned experiment (Fig. 9), with
a nutritive input of 4 g l 1 of organic matter, 2.4
g of calcite were obtained per litre per day (Le
Métayer-Levrel, 1996). The ‘carbonatogenic yield’
(or calcium-carbonate yield) may be defined as the
ratio of the weight of organic matter input to the
weight of calcium carbonate produced. In the case
presented here it is 0.6. We presently keep in the
laboratory about a hundred carbonatogenic strains
collected from various natural environments. Most
of them display carbonatogenic yields around 0.5,
sometimes more, the lowest yield measured being
0.2. Mentioned numbers concern experiments with
monospecific strains. In nature, carbonatogenesis is
generally carried out by plurispecific populations
so that sedimented organic matter may be totally
mineralised into carbonates.
As far as limestones genesis is concerned, skele-
tons and tests produced by animals and algae lead to
the mass of ‘biodetrital limestones’. In the case of
algae, photosynthesis may contribute to mineral pro-
duction. Fungal contribution remains poorly known
and quantitatively limited in marine or paralic en-
vironments where fungi are very scarce. Abiotic
processes, such as evaporation, temperature increase,
CO2 degassing, as well as photosynthesis, are limited
by either carbonate–bicarbonate ion concentrations,
or by the ionic strength of solutions. Furthermore,
eukaryotic photosynthesis is balanced by respiration
so that only cyanobacteria and other photosynthetic
18 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23

Fig. 7. Biocrystalline build-ups. (1) Triangular build-up. Notice the incorporation of bacterial cells into the structure basis. (2) and
(3) Several kinds of cask-shaped build-ups. Notice the incorporation of bacterial cells into the whole structures. (4) Isodiametric
biocrystalline particles, with pentagonal faces. Constituting bacterial cells have locally been removed by preparation and appear as holes
at the surface of crystals. (5) Rhombohedral build-up. Compare to Fig. 5, 2. (6) Large biocrystalline assemblage produced by and solely
composed of different kinds of carbonatogenic bacteria. (7) Close-up of (6) displaying a colony of calcified small bacteria.
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23 19

Fig. 8. Relationships between crystals and bacteria in a cave concretion (helictite from Clamouse, France) in oligotrophic conditions.
(1) Calcite crystal from which short ‘Isosphaera’ chains project. Notice relationships between the orientation of bacterial chains and
crystalline planes and on the surface of particle remains of Vitreoscilla excrescences on crystal surfaces; see (4). (2) ‘Isosphaera’ chain
emerging from a crystal. Notice the basal budding rings and the granulous and rigid aspect of the heavily calcified outer membrane
(cocoon). (3) Bacterial filaments (probably Vitreoscilla) flattening themselves against a mineral particle into which they are integrated.
At the surface of the particle, ghosts of filaments still stand out while several excrescences are still emerging from the mineral mass,
the whole of which is from bacterial origin. (4) Terminal phase of the incorporation into a crystal of calcite of a (calcified) bacterial
filament. Its constitutive equipment is being smoothed in the thread of the crystalline structure. The initial outline of the filament and its
excrescences are still recognisable. The bacterial origin of crystalline material should leave no trace unless initial excrescences (ca. 0.2
µm wide) could be depicted after etching.

bacteria probably play a geological role through this
process which results in the formation of stromato-
lites. The most critical geological question is whether
extensive thicknesses of limestone and early carbon-
ate cements are formed primarily by biotic or abiotic
means. Therefore, it is necessary to compare the ef-
ficiency of heterotrophic bacterial carbonatogenesis
and inorganic precipitation.
At present, from the nutrient-poor open ocean
environment towards littoral and lagoonal environ-
ments, organic matter sedimentation varies between
20 and 10,000 g m 2 y 1 (Basson et al., 1977; Allen
et al., 1979). In such conditions and assuming a cal-
cium-carbonate yield of 0.5, a calcite density of 2.5,
bacterial carbonatogenesis is able to produce, in a
year, a CaCO3 layer the thickness of which would
be between 4 µm and 2 mm. Heterotrophic bac-
terial carbonatogenesis thus may form a limestone
layer from 4 to 2000 m thickness in one million
years (Fig. 10). As a comparison, with the present-
day composition of seawater, and assuming a mean
oceanic evaporation rate of 150 mm y 1 , such a
physical process would produce a CaCO3 layer 15
µm thick, i.e. 15 m in one million years.
Most extensive limestones have formed on shelves
and platforms. On present continental shelves, the or-
20 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23
Fig. 9. Laboratory experiment using the heterotrophic bacterium Bacillus cereus.
ganic matter deposition is comprised between 150 and
1000 g m 2 y 1 .
Such considerations, when applied to real exam-
ples from the geological record, suggest that het-
erotrophic bacterial carbonatogenesis much more
likely accounts for extensive apparently abiotic
(azoic) limestone formation than any other process
(Castanier et al., 1997, 1999).
5. Discussion and conclusions
The environmental conditions of heterotrophic
bacterial metabolic pathways are diverse (aerobiosis,
anaerobiosis, microaerophily). However, carbonate
precipitation always appears to be the response of
heterotrophic bacterial communities to enrichment
of the environment in organic matter.
Nutritional conditions play a major role in the
relationships between crystals and bacteria. Obser-
vations of recently formed or unmodified carbonate
bacterial grains could give information on their orig-
inal nutritional microenvironment and possibly (in
eutrophic conditions) on metabolic pathways of bac-
terial carbonatogenesis. In most cases, however, the
primary bacterial origin of carbonate is blurred by
subsequent diagenesis.
The yield of heterotrophic bacterial carbonatoge-
nesis and the scale of solid carbonate production by
heterotrophic bacteria are potentially much higher
than autotrophic or chemical sedimentation in ma-
rine, paralic or aqueous continental environments.
Furthermore, bacterial heterotrophic carbonatogen-
esis is neither restricted to particular taxonomic
groups of bacteria nor to specific environments so
that it probably has been a ubiquitous phenomenon
since Precambrian times. It just requires organic mat-
ter enrichment. Thus, heterotrophic bacterial carbon-
atogenesis much more plausibly accounts for appar-
ently abiotic limestone deposition and for carbonate
cementation than does any other single process. The
origin of oolites is here included as an outcome
 S. Castanier et al. / Sedimentary Geology 126 (1999) 9–23
Fig. 10. The potential deposition rates of limestones through aerobic heterotrophic bacterial processes.
of heterotrophic activity (Krumbein, 1979a, 1983;
Castanier et al., 1989).
As far as biodetrital particles are concerned, it can
be observed that carbonate shells and tests of organ-
isms result from the activity of their mitochondria
(or chloroplasts). Other organelles may be involved
such as Golgi apparatus (Hemleben et al., 1986; de
Vrind-de Jong and de Vrind, 1997). Since these cel-
lular organites are nowadays considered by a number
of biologists as endosymbiotic bacteria (Margulis
and Sagan, 1986), apart from purely evaporitic and
autotrophic origins, the great majority of limestones
can be considered as being of heterotrophic bacte-
rial origin and the carbon they contain as principally
coming from primary organic matter.
Such considerations provide renewed conceptions
over genesis of azoic limestones, including oolitic
ones, their deposition rates, their initial consistency
and stability (slumping), and their diagenesis. Fi-
nally, bacterial heterotrophic carbonatogenesis ap-
pears as a fundamental phase in the relationships
21
between atmosphere and lithosphere and in the bio-
geological — or geophysiological (Krumbein and
Schellnhuber, 1992) — evolution of the Earth.
Acknowledgements
We are deeply indebted to A. Maurin who long
ago showed us the good track, to G. Camoin who
invited us to the good place, and to W.E. Krumbein
and V.A. Pedone who gave a good help to this paper.
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