248
Microbial detoxification of metals and radionuclides
Jonathan R Lloyd* and Derek R Lovley†
Microorganisms have important roles in the biogeochemical
cycling of toxic metals and radionuclides. Recent advances
have been made in understanding metal–microbe interactions
and new applications of these processes to the detoxification
of metal and radionuclide contamination have been developed.
Addresses
*Department of Earth Sciences, The University of Manchester,
Manchester M13 9PL, UK; jrlloyd@fs1.ge.man.ac.uk
† Department of Microbiology, The University of Massachusetts,
Amherst, Massachusetts 01035, USA
Current Opinion in Biotechnology 2001, 12:248–253
0958-1669/01/$ — see front matter
© 2001 Elsevier Science Ltd. All rights reserved.
Abbreviations
DIRB dissimilatory iron-reducing bacteria
SRB sulphate-reducing bacteria
Introduction
Industrial activities have led to large-scale contamination
of the environment with toxic heavy metals and radio-
nuclides. Chemical approaches are available for metal
remediation, but are often expensive to apply and lack the
specificity required to treat target metals against a back-
ground of competing ions. In addition, such approaches are
not applicable to cost-effective remediation of large-scale
subsurface contamination in situ. Biological approaches, on
the other hand, offer the potential for the highly selective
removal of toxic metals coupled with considerable opera-
tional flexibility; they can be used both in situ or ex situ in
a range of bioreactor configurations. Many such processes
utilise microorganisms that have key roles in the biogeo-
chemical cycling of toxic metals and radionuclides.
Advances in understanding the roles of microorganisms in
such processes, together with the ability to fine-tune their
activities using the tools of molecular biology, has led to
the development of novel or improved metal bioremedia-
tion processes. The aim of this article is to provide an
overview of the mechanisms by which microorganisms
interact with heavy metals and radionuclides, and to high-
light recent advances in the application of these processes
to the detoxification of metal contamination.
Biotechnological applications of metal-
resistant microorganisms
Microbes have evolved to deal with toxic metals using
several mechanisms, see [1•] for an overview. Perhaps the
best-studied metal-resistance system is encoded by genes
of the mer, or mercury resistance, operon. In this system,
Hg(II) is transported into the cell via the MerT transporter
protein, and detoxified by reduction to relatively non-toxic
volatile elemental mercury by an intracellular mercuric
reductase (MerA). This system is described in detail in a
recent review [2•] with new studies extending the model to
include two additional transporter proteins, MerF and
MerC [3]. Mercury reduction by a thermophilic Streptomyces
species has also been reported [4] along with a novel Fe2+-
dependent mechanism for mercury reduction in the
membrane fraction of Thiobacillus ferrooxidans, which may
involve cytochrome c oxidase [5]. A biosensor, constructed
by fusing the mer promoter region (merR) with bacterial
luminescence (lux) genes, has been used to monitor
bioavailable concentrations of mercury in soil [6].
Mercury-resistant bacteria have also been used to detoxify
mercury-contaminated water at the laboratory and pilot
scale. For example, a recombinant Pseudomonas putida
strain was used in a 500 mL reactor to treat sea and river
water containing added Hg2+ [5]. Reduced elemental mer-
cury was continuously stripped by a gas stream and then
captured in KMnO4 solution. Wagner-Dobler and cowork-
ers [7] used an alternative approach, capturing reduced
elemental mercury in a 20 mL immobilised-cell bioreactor
inoculated with a mercury-resistant P. putida species and
subsequently colonised with other mercury-resistant
strains. A companion study demonstrated the successful
removal of Hg2+ from chloralkali electrolysis water at the
laboratory scale [8], before the development of a pilot plant
for Hg2+ removal using this technology [9••]. In the latter
study, a 700 L reactor was packed with pumice granules of
particle size 4–6 mm and inoculated with seven mercury-
resistant Pseudomonas species. Acidic wastewater from a
chloralkali factory was neutralised and amended with
sucrose and yeast extract before introduction into the
bioreactor. Mercury concentrations of up to 10 mg/L were
successfully treated with a retention efficiency of 95%,
although effluent spikes above this concentration had a
deleterious (if reversible) effect on reactor performance.
When operated in combination with an activated carbon
filter, which also became colonised by bacteria, further
removal of mercury to below 10 µg/L was reported. Very
high loadings of mercury were retained in the reactor,
conservatively estimated at 31.5 kg for the 700 L vessel.
In addition to Pseudomonads, mercury-resistance genes are
also found in many other bacteria [2]. For example, the
Gram-negative soil bacterium Ralstonia eutropha (formally
Alcaligenes eutrophus CH34) is resistant to Hg(II) and other
toxic metals by virtue of an extensive battery of inducible
resistance determinants carried on two megaplasmids. This
organism continues to attract considerable interest for the
remediation of both toxic organic compounds and heavy
metals (reviewed in [10•]). Resistance to divalent metal
ions is provided by several efflux mechanisms (e.g. those
encoded by the czc and cnr genes [10•,11–13]). The appeal
of R. eutropha in environmental biotechnology, however,
lies in its ability to precipitate toxic cationic metals within
 Microbial detoxification of metals and radionuclides Lloyd and Lovley
an extracellular coat of insoluble metal carbonates. This is
formed by the generation of high concentrations of metal
ions at the cell surface in a zone of high local pH, caused by
the uptake of protons by metabolically active cells [10•]. In
addition, genes regulating expression of the metal-resis-
tance systems of R. eutropha have been fused to reporter
genes and used in biosensors [10•]. Alternative ‘protein-
based’ sensors have also been developed using
metal-resistance determinants from other bacteria. These
biosensors detect a change in capacitance when a metal
binds to a protein that has been coupled to an electrode
surface [14].
Extraction of metals from contaminated soil
and sediments
The reduction of As(V) to As(III) forms the basis of another
microbial resistance mechanism that has also been studied
in detail (e.g. [15,16]). This process has not proved useful for
bioremediation, because As(III) is more toxic and soluble
than As(V). However, arsenic-resistance genes have found
use in biosensors for As(III) [17•]. Dissimilatory As(V)
reduction, in which As(V) serves as a terminal electron
acceptor in respiration, could also be used to extract insolu-
ble As(V) from contaminated soil. For example, the
dissimilatory As(V)-reducing microorganism Sulfurospirillum
barnesii reduced and mobilised oxidised As(V) that was
coprecipitated on ferrihydrite or alumina phases [18]. Other
studies have confirmed the important roles that microorgan-
isms play in the cycling of arsenic. As(V) is the dominant
electron acceptor for carbon oxidation in some environ-
ments [19••], and a new organism has recently been isolated
that is able to use As(III) as an electron donor for aerobic
growth [20•], closing the biological arsenic cycle.
In addition to metal reduction, other biological mecha-
nisms that can be used for metal extraction include the
production of organic acids by heterotrophic organisms,
and the generation of sulphuric acid through bioxidation of
sulphur (e.g. by Thiobacillus spp.) [21••]. A recent develop-
ment has been the sequential extraction of copper from
contaminated soil by sulphur-oxidising bacteria, followed
by electrokinetic treatment using a DC current to further
acidify the soil and collect the mobilised metals at the
cathode [22]. Preacidification by sulphur-oxidising bacteria
reduced the power required for the electrokinetic step by
66%. In addition to mobilising metals through organic acid
production [23•], recent studies have also shown that
oxalate production by fungi can result in the precipitation
of metals as insoluble metal oxalates [24].
Biomineralisation; formation of insoluble
metal sulphides and phosphates
In addition to promoting the formation of metal carbonates
(see above), microbes can also act as foci for the precipita-
tion of highly insoluble metal sulphides and phosphates.
This can lead to the removal of toxic metals from solution,
but has not been conclusively shown to increase the resis-
tance of microorganisms to metals per se. Sulphide can be
249
produced biologically by several mechanisms. For exam-
ple, Escherichia coli was engineered to produce sulphide by
heterologous expression of the thiosulphate reductase
gene from Salmonella enteritica [25]. Induction of recombi-
nant protein synthesis led to enhanced sulphide
production from thiosulphate under anaerobic conditions,
leading to the precipitation of cadmium sulphide [25].
Metabolic engineering of an aerobic pathway for sulphide
production was also reported [26]. Concerns associated
with the release of recombinant microorganisms may,
however, limit the practical application of such technologies.
Non-engineered organisms, such as a strain of Klebsiella
planticola, have also been reported to precipitate cadmium
by the formation of sulphide from thiosulphate and may
provide a more attractive option [27]. An alternative
approach that continues to receive much attention relies
on the use of sulphate-reducing bacteria (SRB) to convert
sulphate to sulphide under anaerobic conditions (e.g. [28]
and reviewed in [21••]). This approach forms the basis of
one of the few commercial metal biotreatment processes,
operated at the Budelco Zinc Refinery in the Netherlands.
Phosphate is an essential nutrient required for the synthe-
sis of nucleic acids, ATP and other important molecules
and is generally not released in quantity by living organ-
isms. However, two approaches have been studied that
couple biologically liberated phosphate to the formation of
metal phosphate biominerals. The first approach relies on
the accumulation of high concentrations of phosphate at
the cell surface of a Citrobacter species, with phosphate
cleaved from glycerol 2-phosphate. The acid-phosphatase
catalysing this process is associated with the outer mem-
brane and exocellular lipopolysaccharide (LPS) [29•].
Mineral formation was initiated by nucleation at the phos-
phate groups of the LPS, with crystal growth driven by
enzymatically generated phosphate. Cells precipitated
large quantities of uranium and cadmium phosphates, and
the accumulation of ‘tethered’ crystals may have prevented
fouling of the cell surface. An alternative approach that
does not rely on the supply of an organic phosphate donor,
harnesses the cycling of phosphate by Acinetobacter john-
sonii to metal removal [30]. Polyphosphate is synthesised
as an energy source under aerobic conditions; under anaer-
obic conditions it is degraded yielding ATP and promoting
the precipitation of metal phosphates.
Engineering biosorption
Biosorption, the metabolism-independent sorption of heavy
metals and radionuclides to biomass, has been studied in
detail for more than a decade. This research topic has been
reviewed in detail recently and will not be described here
[21••,31]. This is partly because of considerable duplication
and superficiality in the literature and reduced interest in
commercialising this type of technology [21••]. Some new
studies have applied the tools of molecular biology to
enhance metal sorption and do warrant attention. For exam-
ple, a mouse metallothionein was targeted to the outer
membrane of a metal-resistant R. eutropha isolate (see
250 Environmental biotechnology
above) [32••]. The engineered strain accumulated more
Cd2+ than its wild-type counterpart, and offered tobacco
plants some protection from Cd2+ when inoculated into con-
taminated soil. A surface display technique has also been
used successfully to generate ZnO-binding peptides fused
to fimbrae on the surface of E. coli cells [33]. Finally, Gram-
positive bacteria (Staphylococci) have also been engineered to
produce surface-exposed peptides that are able to bind Hg2+
and Cd2+ [34]. Future studies should compare the perfor-
mance of such engineered systems with other more
traditional biosorbants.
Metal remediation through biodegradation of
associated organic compounds
Citrate has found use as a chelating agent in decontamina-
tion operations, forming highly recalcitrant and mobile
metal–citrate complexes. Cultures of Pseudomonas aerugi-
nosa and P. putida were, however, able to grow using a
range of metal–citrate complexes as carbon source, with
metal precipitation promoted by the addition of inorganic
phosphate [35•]. A unique aspect of this study was the use
of P. putida to treat Ni–citrate waste generated by cleaning
a bioinorganic ion-exchange column previously used to
remove nickel. Another industrial chelating agent ethyl-
enediaminetetraacetate (EDTA) also forms very strong
complexes with divalent and trivalent metal ions that are
degraded by bacterial strain DSM 9103 [36]. The
biodegradation of toxic organotin compounds used as bio-
cides and antifouling agents has also received recent
attention (e.g. the degradation of tributyl tin to dibutyl and
monobutyl tin, reviewed in [37]). Degradation of triphenyl
tin by a small soluble extracellular compound secreted by
a fluorescent Pseudomonad has also been reported [38].
Dissimilatory reduction of metals
Dissimilatory iron-reducing bacteria (DIRB) have an
important role in oxidising organic contaminants (e.g.
aromatic hydrocarbons) in the subsurface and can also
immobilise contaminant metals by reduction to less soluble
forms (reviewed in [31,39]). Geobacter spp. are the dominant
organisms selected when Fe(III)-reduction is stimulated in
the subsurface by the addition of various electron donors
and/or electron-shuttling compounds [40••]. Other organ-
isms recently shown to conserve energy through
Fe(III)-reduction include hyperthermophilic bacteria and
archaea. In common with Geobacter species, one such organ-
ism, Pyrobaculum islandicum, also reduced toxic metals
including U(VI), Tc(VII), Cr(VI) and Co(III) [41]. Possible
roles in the formation of uranium deposits and magnetite in
hydrothermal environments were highlighted.
In addition to the direct enzymatic reduction of contaminant
metals, Fe(III)-reducing bacteria can also effect the solubili-
ty of toxic metals via indirect mechanisms. For example,
Tc(VII) was reduced abiotically by nanocrystals of biogenic
magnetite (a mixed Fe(III)/(II) oxide formed by DIRB),
resulting in the precipitation of TcO2 on the surface of the
mineral [42••]. Given the solubility of enzymatically reduced
Tc(IV) noted in one recent study [43], the Fe(II)-mediated
mechanism may be more useful for immobilising Tc in the
subsurface. Zinc incorporation into biogenic magnetite has
also been reported [44]. In addition to promoting metal
immobilisation, Fe(III) reduction may also result in the
liberation of toxic metals associated with insoluble Fe(III)
oxides [21], a process that may be accelerated by aqueous
flux [45]. In order to model Fe(III) reduction accurately, a
prerequisite for use in field applications, it is important to
gain a detailed understanding of the mechanisms of Fe(III)
reduction. A membrane-bound NADPH-dependent Fe(III)
reductase was purified from the Fe(III)-reducing bacterium
Geobacter sulfurreducens [46], consistent with a requirement
for direct contact between Fe(III)-reducing bacteria and
insoluble Fe(III) oxides [47,48].
The ability to reduce toxic Cr(VI) to Cr(III), which can
form insoluble hydroxides, is not restricted to Fe(III)-
reducing bacteria. Several new isolates with potentially
useful properties have been isolated from contaminated
sites in Pakistan [49,50]. In addition, biofilms of SRB also
reduced and precipitated Cr(VI). Cr(VI) reduction was
thought to be enzymatic; reduction by sulphide was
discounted because sulphate reduction was inhibited
dramatically in the presence of chromate [51]. A microbial
consortium was also able to couple the oxidation of phenol
to Cr(VI) reduction, opening the way for treatment of
mixed wastes containing toxic organic contaminants and
metals [52]. Purification of a new chromate reductase from
P. putida was also reported [53].
U(VI) is another priority pollutant reduced by phylogenet-
ically distinct bacteria. Biological reduction of U(VI) to
insoluble U(IV) was stimulated by the addition of ethanol
and trimetaphosphate to contaminated groundwaters [54].
SRB were thought to be responsible for U(VI) reduction,
but a detailed analysis of the microbes involved was not
presented. An SRB-containing microbial mat was
immobilised onto silica particles and used to clean U(VI)-
contaminated groundwater [55] and kinetic data for
biological U(VI) reduction have also been obtained for
another mixed microbial culture containing SRB [56].
U(VI) reduction was enhanced by the addition of sulphate,
but abiotic U(VI) reduction by sulphide was discounted.
The mechanism of U(VI) reduction by Fe(III)-reducing
bacteria has also been investigated. A novel screening
method was used to identify a mutant of Shewanella
putrefaciens unable to reduce U(VI) [57•]. Evidence was
presented to suggest that the mechanism of U(VI)
reduction was distinct from those of Fe(III) and Mn(IV)
reduction, but may share components of the nitrite-reduc-
ing pathway. Uranium has also been shown to shuttle
electrons to Fe(III) oxides in pure cultures of the DIRB
Geobacter metallireducens [58], which may partly explain
inhibition of U(VI) reduction by ferric hydroxides added to
cultures of Shewanella alga BrY [55]. Uranium was not an
effective electron shuttle in sediment samples, however,
and may not perform this role in the subsurface [58].
 Microbial detoxification of metals and radionuclides Lloyd and Lovley
Another actinide that has attracted attention is the mobile
and long-lived α-emitter 237Np present in low-active
nuclear wastes. Removal is ineffective using chemical-
based techniques, but biotreatment of 237Np was possible
using a combination of the biological reduction of Np(V)
by S. putrefaciens followed by precipitation of Np(IV)
phosphate by a Citrobacter sp. [59•].
Finally, the radiotoxicity of effluents containing high-activity
radionuclides (e.g. 235U, 99Tc and 241Pu) may also adversely
effect the microbial component of a bioprocess [31].
Deinococcus radiodurans, an organism that is exceptionally
resistant to ionizing radiation, is being developed using
genetic approaches for the treatment of both metals and
organic contaminants [60]. Recent studies have also shown
that D. radiodurans is able to reduce Cr(VI) directly,
and U(VI) and Tc(VII) using the electron shuttle
anthraquinone-2,6-disulfonate [61•]. Although this organ-
ism may be useful for ex situ treatment, it may not be
applicable for in situ applications as it might not compete
successfully with organisms indiginous to the subsurface.
Conclusions and future directions
Significant advances have been made in understanding the
roles of microorganisms in mineral cycling, and in the
application of these processes to the bioremediation of
metals and radionuclides. Additional advances are expect-
ed with the use of new techniques. For example, genomic
approaches are set to revolutionise many aspects of biolo-
gy and will undoubtedly make an impact in the arena of
environmental biotechnology. Whole genome transcription
profiles have already been screened to uncover potentially
useful metal detoxification strategies in E. coli [62••].
Given the imminent availability of complete genome
sequences for several environmentally relevant microor-
ganisms, this type of approach will surely prove useful for
determining the precise mechanisms of environmentally
relevant metal–microbe interactions.
Acknowledgements
The authors thank the Department of Energy Natural and Accelerated
Bioremediation Research Program for financial support.
References and recommended reading
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have been highlighted as:
• of special interest
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This study showed that Pseudomonas species are able to grow using citrate
complexed to metals (Cd, Zn, Cu, Fe, Co or Ni), with metal precipitation pro-
moted in the presence of inorganic phosphate. P. putida was also used in a
fill-and-draw reactor to treat nickel–citrate-contaminated wastewater.
36. Satroutdinov AD, Dedyukhina EG, Chistyakova TI, Witschel M,
Minkevich IG, Eroshin VK, Egli T: Degradation of metal–EDTA
complexes by resting cells of the bacterial strain DSM 9103.
Environ Sci Technol 2000, 34:1715-1720.
37. Gadd GM: Microbial interactions with tributyltin compounds:
detoxification, accumulation, and environmental fate. Sci Total
Environ 2000, 258:119-127.
38. Inoue H, Takimura O, Fuse H, Murakami K, Kamimura K, Yamaoka Y:
Degradation of triphenyltin by a fluorescent pseudomonad. Appl
Environ Microbiol 2000, 66:3492-3498.
39. Lovley DR, Anderson RT: Influence of dissimilatory metal reduction
on fate of organic and metal contaminants in the subsurface.
Hydrogeol J 2000, 8:77-88.
40. Snoeyenbos-West O, Nevin KP, Anderson RT, Lovley DR: Enrichment
•• of Geobacter species in response to stimulation of Fe(III)
reduction in sandy aquifer sediments. Microbial Ecol 2000,
39:153-167.
Laboratory and field experiments were conducted to determine which micro-
bial populations responded to stimulation of Fe(III) reduction in aquifer sed-
iments. Molecular studies showed that the addition of electron donors or
electron-shuttle compounds resulted in an increase in 16S rDNA sequences
assigned to Geobacter species, proportional to the degree of stimulation of
Fe(III) reduction.
41. Kashefi K, Lovley DR: Reduction of Fe(III), Mn(IV), and toxic metals
at 100°C by Pyrobaculum islandicum. Appl Environ Microbiol 2000,
66:1050-1056.
42. Lloyd JR, Sole VA, Van Praagh CV, Lovley DR: Direct and Fe(II)
•• mediated reduction of technetium by Fe(III)-reducing bacteria.
Appl Environ Microbiol 2000, 66:3743-3749.
In this study, Tc(VII) reduction and precipitation by magnetite, formed during
the reduction of Fe(III) oxides by Geobacter sulfurreducens, was far more
efficient than direct enzymatic reduction of the radionuclide. Studies with
enrichment cultures of Fe(III)-reducing bacteria and sediment samples in
which Fe(III) reduction had been stimulated, confirmed environmental rele-
vance of Fe(II)-mediated reduction of Tc(VII).
43. Wildung RE, Gorby YA, Krupka KM, Hess NJ, Li SW, Plymale AE,
McKinley JP, Fredrickson JK: Effect of electron donor and solution
chemistry on products of dissimilatory reduction of technetium by
Shewanella putrefaciens. Appl Environ Microbiol 2000,
66:2451-2460.
44. Cooper DC, Picardal F, Rivera J, Talbot C: Zinc immobilization and
magnetite formation via ferric oxide reduction by Shewanella
putrefaciens 200. Environ Sci Technol 2000, 34:100-106.
45. Roden EE, Urrutia MM, Mann CJ: Bacterial reductive dissolution of
crystalline Fe(III) oxide in continuous-flow column reactors. Appl
Environ Microbiol 2000, 66:1062-1065.
46. Magnuson TS, Hodges-Myerson AL, Lovley DR: Characterization of
a membrane-bound NADH-dependent Fe3++ reductase from the
dissimilatory Fe3++-reducing bacterium Geobacter sulfurreducens.
FEMS Microbiol Lett 2000, 185:205-211.
47. Nevin KP, Lovley DR: Lack of production of electron-shuttling
compounds or solubilization of Fe(III) during reduction of
insoluble Fe(III) oxide by Geobacter metallireducens. Appl Environ
Microbiol 2000, 66:2248-2251.
48. Lloyd JR, Blunt-Harris EL, Lovley DR: The periplasmic 9.6 kDa
c-type cytochrome of Geobacter sulfurreducens is not an electron
shuttle to Fe(III). J Bacteriol 1999, 181:7647-7649.
49. Badar U, Ahmed N, Beswick AJ, Pattanapiptpaisal P, Macaskie LE:
Reduction of chromate by microorganisms isolated from metal
 Microbial detoxification of metals and radionuclides Lloyd and Lovley
contaminated sites of Karachi, Pakistan. Biotechnol Letts 2000,
22:829-836.
50. Shakoori AR, Makhdoom M, Haq RU: Hexavalent chromium
reduction by a dichromate-resistant Gram-positive bacterium
isolated from effluents of tanneries. Appl Microbiol Biotechnol
2000, 53:348-351.
51. Smith WL, Gadd GM: Reduction and precipitation of chromate by
mixed culture sulphate-reducing bacterial biofilms. J Appl
Microbiol 2000, 88:983-991.
52. Chirwa ES, Wang Y-T: Simultaneous chromium(VI) reduction and
phenol degradation in an anaerobic consortium of bacteria. Wat
Res 2000, 34:2376-2384.
53. Park CH, Keyhan M, Wielinga B, Fendorf S, Matin A: Purification to
homogeneity and characterization of a novel Pseudomonas
putida chromate reductase. Appl Environ Microbiol 2000,
66:1788-1795.
54. Abdelouas A, Lutze W, Gong W, Nuttall EH, Strietelmeier BA,
Travis BJ: Biological reduction of uranium in groundwater and
subsurface soil. Sci Tot Environ 2000, 250:21-35.
55. Bender J, Duff MC, Phillips P, Hill M: Bioremediation and
bioreduction of dissolved U(VI) by microbial mat consortium
supported on silica gel particles. Environ Sci Technol 2000,
34:3235-3241.
56. Spear JR, Figueroa LA, Honeyman BD: Modeling reduction of
uranium U(VI) under variable sulfate concentrations by sulfate-
reducing bacteria. Appl Environ Microbiol 2000, 66:3711-3721.
57. Wade R Jr, DiChristina TJ: Isolation of U(VI) reduction-deficient
• mutants of Shewanella putrefaciens. FEMS Microbiol Letts 2000,
184:143-148.
Chemical mutagenesis was combined with a novel screening technique to
identify a U(VI) reduction-deficient (Urr) mutant that was unable to grow
anaerobically using U(VI) or NO2– as an electron acceptor. This is the first
253
reported isolation of a respiratory mutant unable to grow using U(VI) as an
electron acceptor.
58. Nevin KP, Lovley DR: Potential for nonenzymatic reduction of
Fe(III) during microbial oxidation of organic matter coupled to
Fe(III) reduction. Environ Sci Technol 2000, 34:2472-2478.
59. Lloyd JR, Yong P, Macaskie LE: Biological reduction and removal of
• Np(V) by two microorganisms. Environ Sci Technol 2000,
34:1297-1301.
237Np, a problematic component of nuclear waste streams was removed
from solution using a combination of two biological processes. Biological
reduction of Np(V) by Shewanella putrefaciens preceded precipitation of
Np(IV) as an insoluble phosphate biomineral by a Citrobacter sp.
60. Daly MJ: Engineering radiation-resistant bacteria for
environmental biotechnology. Curr Opin Biotechnol 2000,
11:280-285.
61. Fredrickson JK, Kostandarithes HM, Li SW, Plymale AE, Daly MJ:
• Reduction of Fe(III), Cr(VI), U(VI), and Tc(VII) by Deinococcus
radiodurans R1. Appl Environ Microbiol 2000, 66:2006-2011.
This study showed that the radiation-resistant bacterium D. radiodurans is
able to reduce toxic and radioactive metals directly (e.g. Cr(VI)) or indirectly
in the presence of an electron shuttle anthraquinone-2,6-disulfonate (e.g.
U(VI) and Tc(VII)). Possible applications in the bioremediation of radioactive
waste are highlighted.
62. Brocklehurst KR, Morby AP: Metal-ion tolerance in Escherichia coli:
•• analysis of transcriptional profiles by gene-array technology.
Microbiol 2000, 146:2277-2282.
E. coli were grown in high concentrations of Zn2+, Cd2+, Co2+ or Ni2+ and
whole genome transcription profiles analysed for alterations in transcript
analysis using gene-array technologies. An increase in transcript levels of
genes involved in transposition of insertion sequence elements (e.g insA)
was noted, and expression of insA-7 in E. coli confirmed that the corre-
sponding protein was able to confer tolerance to Zn2+, possibly by binding
of the metal to two paired cysteine motifs.