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Many bacteria can reduce sulfur in small amounts, but

some bacteria can reduce sulfur in large amounts, in
essence, breathing sulfur.
LEARNING OBJECTIVE

 Describe the sulfur cycle

KEY POINTS

 The sulfur cycle describes the movement of sulfur through the geosphere and biosphere.
Sulfur is released from rocks through weathering, and then assimilated
by microbes and plants. It is then passed up the food chain and assimilated by plants
and animals, and released when they decompose.
 Many bacteria can reduce sulfur in small amounts, but some specialized bacteria can
perform respiration entirely using sulfur. They use sulfur or sulfate as an
electron receptor in their respiration, and release sulfide as waste. This is a common
form of anaerobic respiration in microbes.
 Sulfur reducing pathways are found in many pathogenic bacteria species. Tuberculosis
and leprosy are both caused by bacterial species that reduce sulfur, so the
sulfur reduction pathway is an important target of drug development.

TERMS

 assimilatory sulfate reduction
The reduction of 3'-Phosphoadenosine-5'-phosphosulfate, a more elaborated
sulfateester, leads also to hydrogen sulfide, the product used in biosynthesis (e.g., for
the production of cysteine because the sulfate sulfur is assimilated).
 extremophile
An organism that lives under extreme conditions of temperature, salinity, and so on.
They are commercially important as a source of enzymes that operate under similar
conditions.
FULL TEXT

The Sulfur Cycle
The sulfur cycle describes the movement of sulfur through the atmosphere,

mineral forms, and through living things. Although sulfur is primarily found
in sedimentary rocks or sea water, it is particularly important to living things

because it is a component of many proteins.

Sulfur is released from geologic sources through the weathering of rocks. Once

sulfur is exposed to the air, it combines with oxygen, and becomes sulfate

SO4. Plants and microbes assimilate sulfate and convert it into organicforms. As

animals consume plants, the sulfur is moved through the food chain and

released when organisms die and decompose.

Some bacteria – for example Proteus, Campylobacter, Pseudomonas and

Salmonella – have the ability to reduce sulfur, but can also use oxygen and

other terminal electron acceptors. Others, such as Desulfuromonas, use only

sulfur. These bacteria get their energy by reducing elemental sulfur to

hydrogen sulfide. They may combine this reaction with the oxidation of acetate,

succinate, or other organic compounds.

The most well known sulfur reducing bacteria are those in the domain Archea,

which are some of the oldest forms of life on Earth. They are

often extremophiles, living in hot springs and thermal vents where other

organisms cannot live. Lots of bacteria reduce small amounts of sulfates

to synthesize sulfur-containing cell components; this is known as assimilatory

sulfate reduction. By contrast, the sulfate-reducing bacteria considered here

reduce sulfate in large amounts to obtain energy and expel the resulting
sulfide as waste. This process is known as dissimilatory sulfate reduction. In a

sense, they breathe sulfate.

Sulfur metabolic pathways for bacteria have important medical implications.

For example, Mycobacterium tuberculosis (the bacteria causing tuberculosis)

and Mycobacterium leprae (which causes leoprosy) both utilize sulfur, so the

sulfur pathway is a target of drug development to control these bacteria.

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The Role of Micro-organisms in the Sulphur Cycle

Micro-organisms (most frequently bacteria) are often integrally involved in the chemical
alteration of minerals. Minerals, or intermediate products of their decomposition, may be
directly or indirectly necessary to their metabolism. The dissolution of sulphide minerals
under acidic conditions (ARD), the precipitation of minerals under anaerobic conditions,
the adsorption of metals by bacteria or algae, and the formation and destruction of
organometallic complexes are all examples of indirect micro-organism participation.
Where minerals are available as soluble trace elements, serve as specific oxidizing
substrates, or are electron donors/acceptors in oxidation-reduction reactions, they may
be directly involved in cell metabolic activity.

There are three categories of oxidation-reduction reactions for minerals with micro-
organisms:

Oxidation by autotrophic (cell carbon from carbon dioxide) or mixotrophic (cell carbon
from carbon dioxide or organic matter) organisms. Energy derived from the oxidation
reaction is utilized in cell synthesis.

Electron acceptance by minerals (reduction) for heterotrophic (cell carbon from
organic matter) and mixotrophic bacteria. Chemical energy is used to create new cell
material from an organic substrate.

Electron donation by minerals (oxidation) for bacterial or algal photosynthesis
(reaction is fuelled by photon energy).

Natural Oxidation in the Sulphur Cycle

Oxidation of sulphur or sulphides for energy production is restricted to the bacterial
genus Thiobacillus, the genus Thiomicrospira, and the genus Sulfolobus. These
bacteria all produce sulphuric acid (i.e. hydrogen ions, H+, and sulphate ions, SO4-- ) as
a metabolic product. Extensive reviews of these bacteria and their behaviour have been
written by Brierley (1978) and Trudinger (1971).

It is these bacteria that are known to accelerate the generation of Acid Rock Drainage
(ARD) from pyritic and pyrrhotitic rocks under suitable conditions. Evangelou & Zhang
(1995) report that sulphide oxidation catalysed by bacteria may have reaction rates six
orders of magnitude (i.e. 1,000,000 times) greater than the same reactions in the
absence of bacteria. Photomicrographs 1, 2 and 3, from LeRoux, North & Wilson
(1973), illustrate the shape and appearance of T. ferrooxidans: The bacteria develop
flagella only if they are required for mobility in accessing energy sources.

Photomicrographs 1 (left), 2 (centre) and 3 (right): Thiobacilli from bacterial generator (no flagella)
- left & centre - and grown on ferrous iron (flagella) - right. Magnification about x5000 (left),
x20,000 (centre), x15000 (right) (from LeRoux, North & Wilson, 1973)

ARD is the product formed by the atmospheric (i.e. by water, oxygen and carbon
dioxide) oxidation of the relatively common iron-sulphur minerals pyrite and pyrrhotite in
the presence of (catalysed by) bacteria (Thiobacillus ferrooxidans), and any other
products generated as a consequence of these oxidation reactions.

An important reaction involving T. ferrooxidans is the oxidation of ferrous to ferric iron
(Fe++ to Fe+++)

4Fe++ + O2 + 4H+ = Fe+++ + 2H2O

Ferric iron is a powerful oxidizing agent. Even at a Fe+++/Fe++ ratio of 1:1,000,000, a
Redox potential of greater than +0.4 V is generated which is sufficient for the attack of
most base metal sulphides (Dutrizac & MacDonald, 1974). The general equation for the
ferric ion reaction with base metal sulphides is:

MS + nFe+++ = Mn+ + S + nFe++
Consequently T. ferrooxidans, in generating Fe+++, is indirectly responsible for the
dissolution of base metal sulphide minerals and the mobilization of metallic cations such
as Cu++, Zn++, Pb++ and Cd++. Base metal sulphides react only very slowly with sulphuric
acid in the absence of ferric iron (Roman & Benner, 1973).

The importance of Redox potential in determining metal solubility and transport can be
clearly seen for copper in the Eh-pH diagram for the Cu-H2O-O2-S-CO2 system (Figure
2, Garrels & Christ, 1965). The effects of bacteria upon the rate of dissolution of copper
from chalcopyrite are highly pronounced, as demonstrated by Malouf & Prater (1961),
Figure 3.

Figure 2: Eh-ph Diagram for Cu-H2O-O2-S-CO2 System (from Garrels and Christ, 1965)
Figure 3: Effects of Bacteria upon the Rate of Disolution of Copper from Chalcopyrite (from Malouf
and Prater, 1961)

In general, for substantial metal mobilization from base metal sulphides the following
conditions must be met:

Ferric iron for sulphide oxidation

T. ferrooxidans and oxygen for ferrous to ferric oxidation

pH compatible with T. ferrooxidans habitat requirements, typically pH 1.5-3.5 (Roman
& Benner, 1973)

The typical habitat pH of T. ferrooxidans of 1.5 to 3.5 is not one that develops
spontaneously. It is currently believed (Béchard, 1996) that these conditions are
produced by a consortium of bacteria acting in succession. Such a succession may
include T. thioparus at neutral pH, giving way to dominance by metallogenium bacteria
under mildly acid conditions (pH 3.5 to 4.5) (Walsh & Mitchell, 1972), and finally T.
ferrooxidans dominance at low pH.

The metabolic activity of T. ferrooxidans is temperature dependent, peaking at about
30-35 degrees Celsius, and falling with both increasing and decreasing temperature
(Roman & Benner, 1973).

Leduc & Ferroni (1994) have demonstrated that T. ferrooxidans strains are site-specific.

From the above discussion it is clear that consideration of bacterial behaviour is most
important in understanding the process of ARD generation. This is particularly so when
"kinetic" tests are used to predict the rate of generation of ARD in the field. Only if the
bacterial conditions of testwork are identical to those in the field, can rates of ARD
generation and/or metal solubilization be taken from laboratory kinetic testwork and
used to predict field behaviour with any degree of confidence.
Natural Reduction in the Sulphur Cycle

The direct reduction of sulphate ions to hydrogen sulphide is effected in nature by
specialized, strictly anaerobic bacteria of the
genera Desulfovibrio and Desulfotomaculum.

These sulphate reducing bacteria (SRB) are heterotrophic (cell carbon from organic
compounds) organisms that utilize sulphate, thiosulphate, S2O3--, sulphite, SO3--, or
other reducible sulphur-containing ions as terminal electron acceptors in their
respiratory metabolism. In the process these sulphur-containing ions are reduced to
hydrogen sulphide.

The bacteria require an organic substrate which is usually a short chain acid such as
lactic or pyruvic acid. In nature such substrates are generated by the fermentation
activities of other anaerobic bacteria on more complex organic substrates. Thus in
natural systems, the specific requirement for a short chain acid by the SRB is met by
the availability of a complex organic source and a mixed bacterial system. Lactate is
used by the SRB during anaerobic respiration to produce acetate according to the
reaction (Cork and Cusanovich, 1979):

2CH3CHOHCOO- + SO4-- = 2CH3COO- + 2HCO3- + H2S

This is the major natural process for the conversion (destruction) of sulphate ion.
However, the process may be adapted to a controlled engineering process by the use of
anaerobic reaction vessels and carbon monoxide, CO, and hydrogen, H 2, or partially
oxidized propane or natural gas, as the energy source for the bacteria (Warkentin and
Rowley, 1994). This process of sulphate ion to hydrogen sulphide conversion is the first
stage of a pilot plant that operated in 1996 at the former Britannia mine site in British
Columbia, where the second stage utilized the hydrogen sulphide generated to
precipitate copper (Cu++), Zinc (Zn++), and cadmium (Cd++) ions from ARD as metallic
sulphides (Warkentin and Rowley, 1994). Further information, including details of the
performance of this pilot plant may be found on the NTBC Research Corporation web
site.

Steffen, Robertson and Kirsten (B.C.) Inc. (1991) have proposed the use of the
underground mine at Faro, YT, as a giant underground SRB reactor to convert ARD
sulphate ion to hydrogen sulphide, and precipitate zinc contained in the ARD as zinc
sulphide. The proposed system would use the mixed bacterial system present in liquid
cow manure as an SRB source, and sugar as a bacterial carbohydrate (energy) source.

SRB activity in natural wetlands is capable of metal sulphide precipitation from ARD as
a result of sulphate to hydrogen sulphide reduction, and this concept may be extended
to constructed wetlands. Dr. André Sobolewski treats this subject in considerable detail
on his web site Wetlands for Treatment of Mine Drainage. Reviews of this subject have
also been published by the Mine Environment Neutral Drainage (MEND) program
(MEND, 1990, 1993).
Other Micro-organism Reactions in the Sulphur Cycle

Sulphate ion is taken up from soil by plants, which incorporate it into protein, and plant
protein is consumed by animals that convert plant protein to animal protein. Death of
plants and animals allows bacterial decomposition of protein in remains to produce
hydrogen sulphide and other products , in processes involving many fungi,
actinomycetes and bacteria such as the heterotroph Proteus vulgaris.

Some bacteria can function in the transition zone between aerobic and anaerobic
environments,. Hydrogen sulphide may be oxidized to sulphur by such bacteria which
deposit elemental sulphur in their cells while using oxygen as the terminal electron
acceptor.

Hydrogen sulphide may also be oxidized to sulphate photosynthetically by the bacteria,
Chromtiacceae and Chlorobiaceae.

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