ORIGINAL ARTICLE
Zinc toxicity and ATP production in Pseudomonas
fluorescens
A. Alhasawi, C. Auger, V.P. Appanna, M. Chahma and V.D. Appanna
Department Chemistry & Biochemistry, Laurentian University, Sudbury, ON, Canada
Keywords
bioremediation, enzymology, metabolism.
Correspondence
Vasu D. Appanna, Department Chemistry &
Biochemistry, Laurentian University, 935 Ram-
sey Lake Rd, Sudbury, ON P3E 2C6, Canada.
E-mail: vappanna@laurentian.ca
2013/2300: received 15 November 2013,
revised 28 February 2014 and accepted 8
March 2014
doi:10.1111/jam.12497
Abstract
Aims: To identify the molecular networks in Pseudomonas fluorescens that
convey resistance to toxic concentrations of Zn, a common pollutant and
hazard to biological systems.
Methods and Results: Pseudomonas fluorescens strain ATCC 13525 was
cultured in growth medium with millimolar concentrations of Zn. Enzymatic
activities and metabolite levels were monitored with the aid of in-gel activity
assays and high-performance liquid chromatography, respectively. As oxidative
phosphorylation was rendered ineffective, the assimilation of citric acid
mediated sequentially by citrate lyase (CL), phosphoenolpyruvate carboxylase
(PEPC) and pyruvate phosphate dikinase (PPDK) appeared to play a key role
in ATP synthesis via substrate-level phosphorylation (SLP). Enzymes generating
the antioxidant, reduced nicotinamide adenine dinucleotide phosphate
(NADPH) were enhanced, while metabolic modules mediating the formation
of the pro-oxidant, reduced nicotinamide adenine dinucleotide (NADH) were
downregulated.
Conclusions: Pseudomonas fluorescens reengineers its metabolic networks to
generate ATP via SLP, a stratagem that allows the microbe to compensate for
an ineffective electron transport chain provoked by excess Zn.
Significance and Impact of the Study: The molecular insights described here
are critical in devising strategies to bioremediate Zn-polluted environments.
Introduction
Zn is an essential micronutrient in most if not all organ-
isms as it is involved in a variety of biochemical pro-
cesses. It plays a pivotal catalytic, structural and
regulatory role (Stefanidou et al. 2006). This metal is an
important cofactor of numerous enzymes that are critical
in metabolism, transcription and cell signalling (Haase
and Rink 2009; Sera 2009; Bong et al. 2010). However,
when present in elevated amounts, Zn interferes with
numerous biological pathways. This metal can interact
strongly with enzymatic imidazole and sulfhydryl groups
and arrest their reactions (Duruibe et al. 2007). Further-
more, it disturbs the redox potential of the cell and helps
to create an oxidative environment. Indeed, oxidative
stress is an important feature of Zn toxicity (Lemire et al.
2008).
Hence, to survive a Zn-polluted environment, a situa-
tion common due to industrialization and anthropogenic
activity, organisms have evolved a plethora of intricate
strategies to combat this divalent metal (Bondarenko
et al. 2008). Its intracellular immobilization in phytochel-
atins, metalothioneins and other cysteine-rich moieties
has been observed (Blindauer et al. 2002; Di Baccio et al.
2005). Microbes living in Zn-polluted environments are
known to precipitate the metal as sulfide- or phosphate-
containing moieties and actively efflux Zn (Radhika et al.
2006). The synthesis of antioxidants, such as glutathione
and NADPH, is also part of the arsenal at the disposal of
bacteria to counter Zn (Lemire et al. 2010a; Poirier et al.
2013). Although some of the molecular pathways that
mediate Zn detoxification have been reported, the
involvement of metabolic networks in the adaptation to
elevated levels of Zn has yet to be fully uncovered.
Journal of Applied Microbiology © 2014 The Society for Applied Microbiology 1
Journal of Applied Microbiology ISSN 1364-5072