6 Aerobic Degradation by

Microorganisms

WOLFGANG FRITSCHE
MARTIN HOFRICHTER
Jena, Germany

1 Introduction: Characteristics of Aerobic Microorganisms Capable of Degrading Organic

Pollutants 146
2 Principles of Bacterial Degradation 147
2.1 Typical Aerobic Degrading Bacteria 147
2.2 Growth-Associated Degradation of Aliphatics 148
2.3 Diversity of Aromatic Compounds – Unity of Catabolic Processes 149
2.4 Extension of Degradative Capacities 152
2.4.1 Cometabolic Degradation of Organopollutants 152
2.4.2 Overcoming the Persistence by Cooperation of Anaerobic and Aerobic
Bacteria 154
3 Degradative Capacities of Fungi 156
3.1 Metabolism of Organopollutants by Microfungi 157
3.1.1 Aliphatic Hydrocarbons 157
3.1.2 Aromatic Compounds 158
3.2 Degradative Capabilities of Basidiomycetous Fungi 160
3.2.1 The Ligninolytic Enzyme System 161
3.2.2 Degradation of Organopollutants 163
4 Conclusions 164
5 References 164
146 6 Aerobic Degradation by Microorganisms

List of Abbreviations
AsO arsenic-containing organic compounds
BTX benzene, toluene, xylenes
DBDs dibenzo-p-dioxines
DBFs dibenzofurans
DCA 3,4-dichloroaniline
DCP 2,4-dichlorophenol
DDT 1,1,1-trichloro-2,2b-bis(4-chlorophenyl)ethane
DNT 2,4-dinitrotoluene
KCN potassium cyanide
LiP lignin peroxidase
MnP manganese peroxidase
PAHs polycyclic aromatic hydrocarbons
PCBs polychlorinated biphenyls
PCP pentachlorophenol
TCC tricarboxylic acid cycle
TCE trichloroethene
TNT 2,4,6-trinitrotoluene

1 Introduction:
Characteristics of Aerobic
Microorganisms Capable of
Degrading Organic
Pollutants
The most important classes of organic pollu-
tants in the environment are mineral oil con-
stituents and halogenated products of petro-
chemicals. Therefore, the capacities of aerobic
microorganisms are of particular relevance for
the biodegradation of such compounds and
are exemplarily described with reference to
the degradation of aliphatic and aromatic hy-
drocarbons as well as their chlorinated deriva-
tives. The most rapid and complete degrada-
tion of the majority of pollutants is brought
about under aerobic conditions (RISER-RO-
BERTS, 1998).
The essential characteristics of aerobic mi-
croorganisms degrading organic pollutants are Fig. 1. Main principle of aerobic degradation of hy-
(Fig. 1): drocarbons: growth associated processes.

(1) Metabolic processes for optimizing the
contact between the microbial cells and
the organic pollutants. The chemicals
must be accessible to the organisms
having biodegrading activities. For ex-
ample, hydrocarbons are water-insolu-
ble and their degradation requires the
production of biosurfactants.
(2) The initial intracellular attack of organ-
ic pollutants is an oxidative process, the
activation and incorporation of oxygen
is the enzymatic key reaction catalyzed
by oxygenases and peroxidases.
(3) Peripheral degradation pathways con-
vert organic pollutants step by step into
intermediates of the central intermedi-
ary metabolism, e.g., the tricarboxylic
acid cycle.
(4) Biosynthesis of cell biomass from the
central precursor metabolites, e.g., ace-
tyl-CoA, succinate, pyruvate. Sugars re-
quired for various biosyntheses and
growth must be synthesized by gluco-
neogenesis.
A huge number of bacterial and fungal genera
possess the capability to degrade organic pol-
lutants. Biodegradation is defined as the bio-
logically catalyzed reduction in complexity of
chemical compounds (ALEXANDER, 1994). It is
based on two processes: growth and cometab-
olism. In the case of growth, organic pollutants
are used as sole source of carbon and energy.
This process results in a complete degradation
(mineralization) of organic pollutants as dem-
onstrated in Sect. 2.2. Cometabolism is defined
as the metabolism of an organic compound in
the presence of a growth substrate which is
used as the primary carbon and energy source.
The principle is explained in Sect. 2.4.
Enzymatic key reactions of aerobic biodeg-
radation are oxidations catalyzed by oxygen-
ases and peroxidases. Oxygenases are oxido-
reductases that use O2 to incorporate oxygen
into the substrate. Degradative organisms
need oxygen at two metabolic sites, at the in-
itial attack of the substrate and at the end of
the respiratory chain (Fig. 1). Distinct higher
fungi have developed a unique oxidative
system for the degradation of lignin based on
extracellular ligninolytic peroxidases and lac-
cases. This enzymatic system possesses increas-
2 Principles of Bacterial Degradation 147
ing significance for the cometabolic degrada-
tion of persistent organopollutants.Thus it is in
particular the domain of basidiomycetous fun-
gi, which requires a deeper insight and an ex-
tensive consideration. Therefore, this chapter
has been divided in two sections: bacterial and
fungal degradation capacities.
2 Principles of Bacterial
Degradation
2.1 Typical Aerobic Degrading
Bacteria
The predominant degraders of organopollu-
tants in the oxic zone of contaminated areas
are chemo-organotrophic species able to use a
huge number of natural and xenobiotic com-
pounds as carbon sources and electron donors
for the generation of energy. Although many
bacteria are able to metabolize organic pollu-
tants, a single bacterium does not possess the
enzymatic capability to degrade all or even
most of the organic compounds in a polluted
soil. Mixed microbial communities have the
most powerful biodegradative potential be-
cause the genetic information of more than
one organism is necessary to degrade the com-
plex mixtures of organic compounds present in
contaminated areas. The genetic potential and
certain environmental factors such as temper-
ature, pH, and available nitrogen and phos-
phorus sources, therefore, seem to determine
the rate and the extent of degradation.
The predominant bacteria of polluted soils
belong to a spectrum of genera and species
listed in Tab. 1. The composition of this list of
bacteria is determined by the fact whether
they can be cultured on nutrient-rich media.
We have to consider that the majority of bac-
teria present in soils cannot be cultured in the
laboratory yet.
Pseudomonads, aerobic gram-negative rods
that never show fermentative activities, seem
to have the highest degradative potential, e.g.,
Pseudomonas putida and P. fluorescens. Fur-
ther important degraders of organic pollutants
can be found within the genera Comamonas,
148 6 Aerobic Degradation by Microorganisms
Tab. 1. Predominant Bacteria in Soil Samples Pollut-
ed with Aliphatic and Aromatic Hydrocarbons, Po-
lycyclic Aromatic Hydrocarbons, and Chlorinated
Compoundsa
Gram-Negative Gram-Positive
Bacteria Bacteria
Pseudomonas spp. Nocardia spp.
Acinetobacter spp. Mycobacterium spp.
Alcaligenes sp. Corynebacterium spp.
Flavobacterium/ Arthrobacter spp.
Cytophaga group
Xanthomonas spp. Bacillus spp.
a
The reclassification of bacteria has been based on
phylogenetic markers resulting in changes of some
genera and species. This is why the names of species
are not mentioned.
Burkholderia, and Xanthomonas. Some spe-
cies utilize `100 different organic compounds
as carbon sources. The immense potential of
the pseudomonas does not solely depend on
the catabolic enzymes, but also on their capa-
bility of metabolic regulation (HOUGHTON and
SHANLEY, 1994). A second important group of
degrading bacteria are the gram-positive rhod-
ococci and coryneform bacteria. Many species,
now classified as Rhodococcus spp. had origi-
nally been described as Nocardia spp., Myco-
bacterium spp., and Corynebacterium spp.
Rhodococci are aerobic actinomycetes show-
ing considerable morphological diversity. A
certain group of these bacteria possess mycol-
ic acids at the external surface of the cell.
These compounds are unusual long-chain al-
cohols and fatty acids, esterified to the pepti-
doglycan of the cell wall. Probably, these lipo-
philic cell structures have a significance for the
affinity of rhodococci to lipophilic pollutants.
In general, rhodococci have high and diverse
metabolic activities and are able to synthesize
biosurfactants.
2.2 Growth-Associated
Degradation of Aliphatics
The aerobic initial attack of aliphatic and cy-
cloaliphatic hydrocarbons requires molecular
oxygen. Fig. 2 shows both types of enzymatic
reactions involved in these processes. It de-
pends on the nature of the substrate and the
enzymatic equipment of the involved microor-
ganisms, what kind of enzymatic reaction is
realized. n-Alkanes are the main constituents
of mineral oil contaminations (HINCHEE et al.,
1994). Long-chain n-alkanes (C10UC24) are
degraded most rapidly by mechanisms demon-
strated in Fig. 3. Short-chain alkanes (less than
C9) are toxic to many microorganisms, but
they evaporate rapidly from petroleum con-
taminated sites. Oxidation of alkanes is classi-
fied as being terminal or diterminal. The
monoterminal oxidation is the main pathway.
It proceeds via the formation of the corre-
sponding alcohol, aldehyde, and fatty acid.
b-Oxidation of the fatty acids results in the
formation of acetyl-CoA. n-Alkanes with an
uneven number of carbon atoms are degraded
to propionyl-CoA, which is in turn carboxylat-
ed to methylmalonyl-CoA and further con-
verted to succinyl-CoA. Fatty acids of a phys-
iological chain length may be directly incorpo-
rated into membrane lipids, but the majority of
degradation products is introduced into the
tricarboxylic acid cycle. The subterminal oxi-
dation occurs with lower (C3UC6) and longer
alkanes with the formation of a secondary al-
cohol and subsequent ketone. Unsaturated 1-
alkenes are oxidized at the saturated end of
the chains. A minor pathway has been shown
to proceed via an epoxide, which is converted
to a fatty acid. Branching, in general, reduces
the rate of biodegradation. Methyl side groups
do not drastically decrease the biodegradabili-
ty, whereas complex branching chains, e.g., the
tertiary butyl group, hinder the action of the
degradative enzymes.
Cyclic alkanes representing minor compo-
nents of mineral oil are relatively resistant to
microbial attack. The absence of an exposed
terminal methyl group complicates the pri-
mary attack. A few species are able to use cy-
clohexane as sole carbon source; more com-
mon is its cometabolism by mixed cultures.
The mechanism of cyclohexane degradation is
shown in Fig. 4. In general, alkyl side chains of
cycloalkanes facilitate the degradation.
Aliphatic hydrocarbons become less water
soluble with increasing chain length. Hydro-
carbons with a chain length of C12 and above
are virtually water insoluble. Two mechanisms
 2 Principles of Bacterial Degradation 149

Fig. 2. Initial attack on xe-

nobiotics by oxygenases.
Monooxygenases incorpo-
rate one atom of oxygen of
O2 into the substrate, the
second atom is reduced to
H2O. Dioxygenases incor-
porate both atoms into the
substrate.

are involved in the uptake of these lipophilic
substrates: the attachment of microbial cells at
oil droplets and the production of biosurfac-
tants (HOMMEL, 1990). The uptake mechanism
linked to attachment of the cells is still un-
known, whereas the effect of biosurfactants
has been studied well (Fig. 5). Biosurfactants
are molecules consisting of a hydrophilic and a
lipophilic moiety. They act as emulsifying
agents, by decreasing the surface tension and
by forming micelles. The microdroplets may be
encapsulated in the hydrophobic microbial
cell surface. The products of hydrocarbon deg-
radation, introduced to the central tricarboxyl-
ic acid cycle, have a dual function. They are
substrates of the energy metabolism and build-
ing blocks for biosynthesis of cell biomass
and growth (Fig. 1). The synthesis of amino
acids and proteins needs a nitrogen and sulfur
source, that of nucleotides and nucleic acids a
phosphorus source. The biosynthesis of the
bacterial cell wall requires activated sugars
synthesized by gluconeogenesis.
Products of growth-associated degradation
are CO2, H2O, and cell biomass. The cells act as
the complex biocatalysts of degradation. In ad-
dition, cell biomass may be mineralized after
exhaustion of the degradable pollutants in a
contaminated site.
2.3 Diversity of Aromatic
Compounds – Unity of Catabolic
Processes
Aromatic hydrocarbons, e.g., benzene, tolu-
ene, ethylbenzene and xylenes (BTEX com-
pounds), and naphthalene belong to the large
volume petrochemicals, widely used as fuels
and industrial solvents. Phenols and chloro-
phenols are released into the environment as
150 6 Aerobic Degradation by Microorganisms
products and waste materials from industry.
Aromatic compounds are formed in large
amounts by all organisms, e.g., as aromatic
amino acids, phenols, or quinones. Thus, it is
not surprising that many microorganisms have
evolved catabolic pathways to degrade aromat-
ic compounds. In general, man-made organic
chemicals (xenobiotics) can be degraded by
microorganisms, when the respective molecu-
les are similar to natural compounds. The di-
versity of man-made aromatics shown in Fig. 6
can be converted enzymatically to natural in-
termediates of the degradation: catechol and
protecatechuate. In general, benzene and re-
lated compounds are characterized by a higher
thermodynamic stability than aliphatics are.
Fig. 3. Peripheral pathways
of alkane degradation. The
main pathway is the termi-
nal oxidation to fatty acids
catalyzed by a n-alkane
monoxygenase, b alcohol
dehydrogenase and c al-
dehyde dehydrogenase.
Only few reports on bacteria capable of
attacking benzene have been published
(SMITH, 1990). The first step of benzene oxida-
tion is a hydroxylation catalyzed by a dioxygen-
ase (Fig. 2).The product, a diol, is then convert-
ed to catechol by a dehydrogenase. These ini-
tial reactions, hydroxylation and dehydrogena-
tion, are also common to pathways of degrada-
tion of other aromatic hydrocarbons.
The introduction of a substituent group on-
to the benzene ring renders alternative mecha-
nisms possible to attack side chains or to oxid-
ize the aromatic ring. The versatility and
adaptability of bacteria is based on the exis-
tence of catabolic plasmids. Catabolic plasmids
have been found to encode enzymes degrading

Fig. 4. Peripheric metabolic pathway of cycloaliphat-
ic compounds (cycloparaffins).
naturally occurring aromatics such as cam-
phor, naphthalene, and salicylate. Most of the
catabolic plasmids are self-transmissible and
have a broad host range. The majority of gram-
negative soil bacteria isolated from polluted
areas possess degradative plasmids, mainly the
so called TOL plasmids. These pseudomonads
are able to grow on toluene, m- and p-xylene,
and m-ethyltoluene. The main reaction in-
volved in the oxidation of toluene and related
arenes is the methyl group hydroxylation. The
methyl group of toluene is oxidized stepwise
to the corresponding alcohol, aldehyde, and
carboxylic group. Benzoate formed or its alky-
2 Principles of Bacterial Degradation 151
lated derivatives are then oxidized by toluate
dioxygenase and decarboxylated to catechol
(SMITH, 1990).
The oxygenolytic cleavage of the aromatic
ring occurs via o- or m-cleavage. The signifi-
cance of the diversity of degradative pathways
and of the few key intermediates is still under
discussion. Both pathways may be present in
one bacterial species. “Whenever an alternative
mechanism for the dissimilation of any com-
pound becomes available (ortho- versus meta-
cleavage of ring structures, for example) control
of each outcome must be imposed” (HOUGH-
TON and SHANLEY, 1994). The metabolism of a
wide spectrum of aromatic compounds by one
species requires the metabolic isolation of
intermediates into distinct pathways. This kind
of metabolic compartmentation seems to be
realized by metabolic regulation. The key en-
zymes of the degradation of aromatic sub-
strates are induced and synthesized in appre-
ciable amounts only when the substrate or
structurally related compounds are present.
Enzyme induction depends on the concentra-
tion of the inducing molecules. The substrate
specific concentrations represent the thresh-
old of utilization and growth and are in the
magnitude of µM. A recent report on the regu-
lation of TOL catabolic pathways has been
published by RAMOS et al. (1997).
Fig. 7 shows the pathways of the oxygenolyt-
ic ring cleavage to intermediates of the central
metabolism. At the branchpoint catechol ei-
ther is oxidized by the intradiol o-cleavage, or
the extradiol m-cleavage. Both ring cleavage
reactions are catalyzed by specific dioxygen-
ases. The product of the o-cleavage – cis,cis-
muconate – is transferred to the instable enol-
lactone, which is in turn hydrolyzed to oxoad-
ipate. This dicarboxylic acid is activated by
transfer to CoA, followed by the thiolytic
cleavage to acetyl-CoA and succinate. Proto-
catechuate is metabolized by a homologous set
of enzymes. The additional carboxylic group is
decarboxylated and, simultaneously, the dou-
ble bond is shifted to form oxoadipate enol-
lactone. The oxygenolytic m-cleavage yields 2-
hydroxymuconic semialdehyde, which is me-
tabolized by the hydrolytic enzymes to for-
mate, acetaldehyde, and pyruvate. These are
then utilized in the central metabolism. In gen-
eral, a wealth of aromatic substrates is degrad-
152 6 Aerobic Degradation by Microorganisms
ed by a limited number of reactions: hydroxy-
lation, oxygenolytic ring cleavage, isomeriza-
tion, hydrolysis.The inducible nature of the en-
zymes and their substrate specificity enable
bacteria with a high degradation potential, e.g.,
pseudomonads and rhodococci, to adapt their
metabolism to the effective utilization of sub-
strate mixtures in polluted soils and to grow at
a high rate.
2.4 Extension of Degradative
Capacities
2.4.1 Cometabolic Degradation of
Organopollutants
Cometabolism, the transformation of a sub-
stance without nutritional benefit in the pres-
ence of a growth substrate, is a common phe-
nomenon of microbial activities. It is the basis
of biotransformations (bioconversions) used
Fig. 5. Involvement of
biosurfactants in the
uptake of hydrocar-
bons. The figure demon-
strates the emulsifying
effect of a rhamnolipid
produced by Pseudo-
monas spp. within the
oil–water interphase
and the formation of
micelles. Lipid phases
are printed in bold.
in biotechnology to convert a substance to a
chemically modified form. Microorganisms
growing on a particular substrate gratuitously
oxidize a second substrate (cosubstrate). The
cosubstrate is not assimilated, but the product
may be available as substrate for other organ-
isms of a mixed culture.
The prerequisites of cometabolic transfor-
mations are the enzymes of the growing cells
and the synthesis of cofactors necessary for en-
zymatic reactions, e.g., of hydrogen donors (re-
ducing equivalents, NADH) for oxygenases.
The principle is shown in Fig. 8. The example
demonstrated in Fig. 8 has been used in field
experiments for the elimination of trichloro-
ethylene (THOMAS and WARD, 1989). Methan-
otrophic bacteria used in this experiment can
utilize methane and other C1 compounds as
sole sources of carbon and energy. They oxi-
dize methane to CO2 via methanol, formalde-
hyde, and formate. The assimilation requires
special pathways, and formaldehyde is the
intermediate assimilated. The first step of

Fig. 6. Degradation of a broad
spectrum of aromatic natural
and xenobiotic compounds into
two central intermediates: cate-
chol and protocatechuate.
methane oxidation is catalyzed by methane
monooxygenase, which attacks the inert CH4.
It is an unspecific enzyme that also oxidizes
various other compounds, e.g., alkanes, aro-
matic compounds, and trichloroethylene
(TCE). The proposed mechanism of TCE
transformation according to HENRY and
GRBIĆ -GALLIĆ (1994) is shown in Fig. 8.TCE is
oxidized to an epoxide excreted from the cell.
The unstable oxidation product breaks down
to compounds, which may be used by other mi-
croorganisms. Methanotrophic bacteria are
aerobic indigenous bacteria, in soil and aqui-
2 Principles of Bacterial Degradation 153
fers, but methane has to be added as growth
substrate and inducer for the development of
methanotrophic biomass. The addition of
methane as substrate limits the application for
bioremediation.
Cometabolism of chloroaromatics is a wide-
spread activity of bacteria in mixtures of in-
dustrial pollutants. KNACKMUSS (1997) demon-
strated that the cometabolic transformation of
2-chlorophenol gives rise to dead end metab-
olites, e.g., 3-chlorocatechol. This reaction
product may be auto-oxidized or polymerized
in soil to humic-like structures. Irreversible
154 6 Aerobic Degradation by Microorganisms
binding of dead end metabolites may fulfill the
function of detoxification. The accumulation
of dead end products within microbial commu-
nities under selection pressure is the basis for
the evolution of new catabolic traits (REIN-
ECKE, 1994).
2.4.2 Overcoming the Persistence
by Cooperation of Anaerobic and
Aerobic Bacteria
As a rule, recalcitrance of organic pollutants
increases with increasing halogenation. Substi-
Fig. 7. The two alternative pathways
of aerobic degradation of aromatic
compounds: o- and m-cleavage, a
phenol monoxygenase, b catechol
1,2-dioxygenase, c muconate lacton-
izing enzyme, d muconolactone iso-
merase, e oxoadipate enol-lactone
hydrolase, f oxoadipate succinyl-
CoA transferase, g catechol 2,3-di-
oxygenase, h hydroxymuconic semi-
aldehyde hydrolase, i 2-oxopent-4-
enoic acid hydrolase, j 4-hydroxy-2-
oxovalerate aldolase.
tution of halogen as well as nitro and sulfo
groups at the aromatic ring is accomplished by
an increasing electrophilicity of the molecule.
These compounds resist the electrophilic at-
tack by oxygenases of aerobic bacteria. Com-
pounds that persist under oxic conditions are,
e.g., PCBs (polychlorinated biphenyls), chlori-
nated dioxins, some pesticides, e.g., DDT.
To overcome the relatively high persistence
of halogenated xenobiotics, reductive attack of
anaerobic bacteria is of significance. The deg-
radation of environmental pollutants by an-
aerobic bacteria is the subject of Chapter 7,
this volume. Because of the significance of re-
ductive dehalogenation for the first step in the

Fig. 8. Cometabolic degradation of trichlorometh-
ane by the methane monoxygenase system of
methanotrophic bacteria.
degradation of higher halogenated com-
pounds, this process has been announced. Re-
ductive dehalogenation effected by anaerobic
bacteria is either a gratuitous reaction or a new
type of anaerobic respiration (ZEHNDER, 1988).
The process reduces the degree of chlorination
and, therefore, makes the product more ac-
cessible to mineralization by aerobic bacteria.
The potential of a sequence of anaerobic
and aerobic bacterial activities for the mineral-
ization of chlorinated xenobiotics is described
in Fig. 9. PCBs, which are selected as an exam-
ple for degradation of halogenated com-
pounds, are well-studied objects (TIEDJE et al.
1993; SYLVESTRE and SANDOSSI, 1994; BEDARD
and QUENSEN, 1995). The scheme demon-
strates the principle of enzymatic dehalogena-
tion mechanisms. The realization of the reac-
tions depends on the structure of the chemical
compounds as well as on the microorganisms
and conditions in a polluted ecosystem. We
have to distinguish between the general degra-
dation potential and the actual conditions nec-
essary for its realization. Reductive dehalog-
enation, the first step of PCB degradation, re-
quires anaerobic conditions and organic sub-
strates acting as electron donors. The PCBs
2 Principles of Bacterial Degradation 155
have the function of an electron acceptor to al-
low the anaerobic bacteria to transfer elec-
trons to these compounds. Anaerobic bacteria
capable of catalyzing reductive dehalogena-
tion seem to be relatively ubiquitous in nature.
Most dechlorinating cultures are mixed cul-
tures (consortia). Aanaerobic dechlorination
is always incomplete, products are di- and
monochlorinated biphenyls. These products
can be metabolized further by aerobic micro-
organisms. The substantial reduction of PCBs
by sequential anaerobic and aerobic treatment
has been demonstrated in the laboratory (AB-
RAMOWICZ, 1990).
The principle of aerobic microbial dehalog-
enation reactions of chloroaromatics are de-
scribed in Fig. 9. Hydrolytic dechlorination has
been elucidated using 4-chlorobenzoate as
substrate for Pseudomonas and Nocardia spp.
A halidohydrolase is capable of replacing the
halogen substituent by a hydroxy group origi-
nating from water. This type of reaction seems
to be restricted to halobenzoates substituted in
the p-position. Dechlorination after ring cleav-
age is a common reaction of the o-pathway of
chlorocatechols catalyzed by catechol 1,2-
dioxygenases to produce chloromuconates.
The oxygenolytic dechlorination is a rare for-
tuitous reaction catalyzed by mono- and di-
oxygenases. During this reaction, the halogen
substituent is replaced by oxygen of O2.
Higher chlorinated phenols, e.g., pentachlo-
rophenol, have been widely used as biocides.
Several aerobic bacteria that degrade chloro-
phenols have been isolated (Flavobacterium,
Rhodococcus). The degradation mechanism
has been elucidated in some cases (MCALLIS-
TER et al., 1996). Thus, Rhodococcus chloro-
phenolicus degrades pentachlorophenol
through a hydrolytic dechlorination and three
reductive dechlorinations, producing trihy-
droxybenzene (APAJALAKTI and SALKINOJA-
SALONEN, 1987). The potential of these bacte-
ria is limited to some specialists and specific
conditions. Therefore, the use of polychlorinat-
ed phenols has been banned in many coun-
tries.