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Electrical Power by Micro
Electrical Power by Micro
12
Electricity-producing bacterial
communities in microbial fuel cells
Bruce E. Logan and John M. Regan
Department of Civil and Environmental Engineering, Penn State University, University Park, PA 16802, USA
Microbial fuel cells (MFCs) are not yet commercialized generated to power subsurface devices [5]. It now seems
but they show great promise as a method of water that electricity can be generated from any biodegradable
treatment and as power sources for environmental sen- material, ranging from pure compounds such as acetate,
sors. The power produced by these systems is currently glucose, cysteine, bovine serum albumin and ethanol [6–
limited, primarily by high internal (ohmic) resistance. 11] to complex mixtures of organic matter including domes-
However, improvements in the system architecture will tic (human), animal, food-processing and meat-packing
soon result in power generation that is dependent on the wastewaters [10,12–14].
capabilities of the microorganisms. The bacterial com- With slight modifications to the MFC architecture and
munities that develop in these systems show great operation, hydrogen can be produced instead of electricity
diversity, ranging from primarily d-Proteobacteria that in relatively high yields. If oxygen is removed from the
predominate in sediment MFCs to communities com- cathode and a small additional voltage is applied to the
posed of a-, b-, g- or d-Proteobacteria, Firmicutes and circuit, hydrogen gas is evolved from the cathode. This
uncharacterized clones in other types of MFCs. Much process has been referred to as a bio-electrochemically
remains to be discovered about the physiology of these assisted microbial reactor (BEAMR) [15,16] or simply as
bacteria capable of exocellular electron transfer, collec- the bacterial electrolysis of organic matter [17] because the
tively defined as a community of ‘exoelectrogens’. Here, protons and electrons are derived from the organic matter
we review the microbial communities found in MFCs and and not water. Water electrolysis is a highly endothermic
the prospects for this emerging bioenergy technology. reaction requiring, in practice, the application of 1.8 V.
However, bacterial electrolysis of organic matter is
Microbial fuel cells make it possible to generate exothermic, providing energy for the bacteria and produ-
electricity using bacteria cing an anode potential of approximately 0.3 V (versus a
It has been known for almost one hundred years that normal hydrogen electrode). At neutral pH, this is insuffi-
bacteria could generate electricity [1], but only in the past cient to generate hydrogen gas from the protons and
few years has this capability become more than a labora- electrons produced in this system. By adding 0.25 V
tory novelty. The reasons for this recent interest in using (0.11 V in theory), it is possible to produce hydrogen gas
bacteria to generate electricity are a combination of the at the cathode. Yields of hydrogen production from acetate
need for new sources of energy, discoveries about microbial have reached 2.9 moles of hydrogen per mole of acetate
physiology related to electron transport, and the advance- (versus a theoretical maximum of 4 mol/mol) for an energy
ment of fuel-cell technologies. In a microbial fuel cell input equivalent to 0.5 moles of hydrogen. This represents
(MFC), bacteria are separated from a terminal electron a net energy gain by a factor of 5.8 in terms of the electricity
acceptor at the cathode so that the only means for respira- needed for biohydrogen production by the BEAMR process,
tion is to transfer electrons to the anode (Figure 1). The versus a net loss of energy needed for water electrolysis.
electrons flow to the cathode as a result of the electroche- The net energy input for biohydrogen is, therefore, sus-
mical potential between the respiratory enzyme and the tained by the organic matter degraded by the bacteria.
electron acceptor at the cathode. Electron transfer from the It is a surprise to many researchers that the most
anode to the cathode must be matched by an equal number significant block to achieving high power densities in
of protons moving between these electrodes so that MFCs is the system architecture, not the composition of
electroneutrality is preserved (Box 1). the bacterial community. Early systems produced very low
From the 1960s until quite recently, it was thought that power densities (<0.1 mW/m2, normalized by the anode
exogenous mediators needed to be added into the fuel cell surface area) [4] but power output by MFCs has been
to generate reasonable amounts of power. However, Kim consistently increasing over time (Figure 2). In our own
and co-workers [2–4] demonstrated that power could be laboratory, improvements in system architecture and
generated by a naturally existing consortium of bacteria in operation have increased power densities from <1 mW/
the absence of exogenous mediators. Others then showed m2 to >1500 mW/m2 using oxygen as the final electron
that by simply placing an anode into anoxic sediment and a acceptor at the cathode [7,18]. Platinum is used as a
cathode in overlying water, sufficient current could be cathode catalyst, although cobalt- and iron-based catalysts
can both produce similar power densities in these systems
Corresponding author: Logan, B.E. (blogan@psu.edu). [19,20]. Power densities as high as 4.31 W/m2 have been
Available online 16 October 2006. achieved, although this has required a non-renewable
www.sciencedirect.com 0966-842X/$ – see front matter ß 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.tim.2006.10.003
Review TRENDS in Microbiology Vol.14 No.12 513
Figure 4. (a) A scanning electron micrograph (SEM) of wild-type Shewanella oneidensis MR-1 grown under electron-acceptor-limited conditions, showing pilus-type
nanowires that connect to other cells. (b) STM image of a single pilus-type nanowire from wild-type MR-1 (lateral diameter of 100 nm, topographic height of 5–10 nm)
showing ridges and troughs running along the long axis of the structures consistent with a bundle of wires. The corresponding conductivity of the pilus as the tip moves
over the indicated surface is shown beneath the STM image. (c) The anode from an MFC colonized by S. oneidensis MR-1. (d) An SEM image of Pelotomaculum
thermopropionicum and Methanothermobacter thermautotrophicus (arrow) in methanogenic co-cultures showing pili connecting the two genera. Subsequent STM
imaging has shown that the pili are conductive. Parts (a), (b) and (d) reproduced, with permission, from Ref. [38]; part (c) was provided by Y. Gorby.
occur in as little as three to four days when the reactor is lines of evidence for cell–cell interactions based on
emptied and re-filled with fresh medium on a daily basis nanowires. For example, consider the symbiotic interactions
[33]. Rapid acclimation might not be so unexpected between fermentative and methanogenic bacteria. It is well
when using sediments that are already relatively enriched established that close proximity is advantageous to both
in iron-reducing bacteria. However, we see rapid acclima- microorganisms to facilitate the exchange of hydrogen,
tion of an MFC when using domestic wastewater samples, allowing for interspecies hydrogen transfer. However, there
with no apparent selective pressure for electrochemically now seems to be evidence for direct electron transfer as well.
active bacteria to be present in this system. It has been Co-cultures studied by Ishii et al. [40] showed the presence of
found by microautoradiography (MAR) using radiolabeled a flagellum-like filament between the propionate fermentor
acetate, inhibition of sulfate reducers and methanogens, (Pelotomaculum thermopropionicum) and a methanogen
and ferric iron as the sole electron acceptor, that iron- (Methanothermobacter thermautotrophicus; Figure 4d). A
reducing bacteria in activated sludge might comprise as closer morphological examination of the filament produced
much as 3% of the bacteria [39]. Interestingly, the combi- by P. thermopropionicum shows that it closely resembles the
nation of MAR with fluorescence in situ hybridization thick pilus-like appendage produced by S. oneidensis MR-1.
showed that all of the MAR-positive cells hybridized with STM measurements established that the appendage of
a bacteria-targeted probe but the Proteobacteria-subclass- strain MR-1 is conductive and can be considered to be a
targeted probes only identified 20% of these iron reducers nanowire [38] as can the thinner filaments produced by
as g-Proteobacteria and 10% as d-Proteobacteria; 70% did Geobacter sulfurreducens [41]. Furthermore, the oxygenic
not hybridize with Proteobacteria-targeted probes. This phototrophic cyanobacterium Synechocystis PCC6803 also
suggests that the diversity of iron-reducing and potentially produces conductive nanowires, although the ecological
relevant microbes for electricity production might extend advantage to that bacterium is not known [38]. The exam-
beyond the commonly studied Shewanella and Geobacter ination of micrographs of Shewanella biofilms on electrodes
species. shows that some of these wires not only go from the cell to the
The finding that iron-reducing bacteria are a measurable electrode but also from one cell to another (Figure 4c). Thus,
and, presumably, integral component of the bacterial it might be that nanowires have a previously unforeseen role
community in wastewater reactors suggests that bacteria in cell–cell electron transfer and microbial community
capable of wired (or wireless) electron transfer succeed in the development.
community in unexplained ways. There is now preliminary The observations of the ability of a wide range of
evidence from a variety of studies for syntrophic relation- bacteria to transfer electrons exocellularly [42] to iron
ships on the basis of interspecies electron transfer. Although and other metal oxides, to carbon electrodes and possibly
such evidence is not yet conclusive, there are compelling to other bacteria suggest that a new term is needed to
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Review TRENDS in Microbiology Vol.14 No.12 517
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