Enrichment of Hydrogen Oxidizing Bacteria from High Temperature and Salinity Environments.

There is an urgent need for sustainable protein supply routes with low environmental footprint. Recently, the use of hydrogen oxidizing bacteria (HOB) as a platform for high quality microbial protein (MP) production has regained interest. This study aims to investigate the added value of using conditions such as salt and temperature to steer HOB communities to lower diversities, while maintaining a high protein content and a high quality amino acid profile. Pressure drop and hydrogen consumption were measured for 56 days to evaluate autotrophy of a total of six communities in serum flasks. Of the six communities, four were enriched under saline (0.0, 0.25, 0.5 and 1.0 mol NaCl l-1) and two under thermophilic conditions (65°C). Five communities enriched for HOB were subsequently cultivated in continuously stirred reactors under the same conditions to evaluate their potential as microbial protein producers. The protein percentages ranged from 41 to 80%. The highest protein content was obtained for the thermophilic enrichments. Amino acid profiles were comparable to protein sources commonly used for feed purposes. Members of the genus Achromobacter were found to dominate the saline enrichments while members of the genus Hydrogenibacillus were found to dominate the thermophilic enrichments. Here we show that enriching for HOB while steering the community toward low diversity and maintaining a high quality protein content can be successfully achieved, both in saline and thermophilic conditions.IMPORTANCE Alternative feed and food supply chains are required to decrease water and land use. HOB offer a promising substitute for traditional agricultural practice to produce microbial protein (MP) from residual materials and renewable energy. To safeguard product stability, the composition of the HOB community should be controlled. Defining strategies to maintain the stability of the communities is therefore key for optimization purposes. In this study, we use salt and temperature as independent conditions to stabilize the composition of the HOB communities. Based on the results presented, we conclude that HOB communities can be steered to have low diversity using the presented conditions while producing a desirable protein content with a valuable amino acid profile.

Hydrogen Gas Recycling for Energy Efficient Ammonia Recovery in Electrochemical Systems.

Recycling of hydrogen gas (H2) produced at the cathode to the anode in an electrochemical system allows for energy efficient TAN (Total Ammonia Nitrogen) recovery. Using a H2 recycling electrochemical system (HRES) we achieved high TAN transport rates at low energy input. At a current density of 20 A m-2, TAN removal rate from the influent was 151 gN m-2 d-1 at an energy demand of 26.1 kJ gN-1. The maximum TAN transport rate of 335 gN m-2 d-1 was achieved at a current density of 50 A m-2 and an energy demand of 56.3 kJ gN-1. High TAN removal efficiency (73-82%) and recovery (60-73%) were reached in all experiments. Therefore, our HRES is a promising alternative for electrochemical and bioelectrochemical TAN recovery. Advantages are the lower energy input and lower risk of chloride oxidation compared to electrochemical technologies and high rates and independence of organic matter compared to bioelectrochemical systems.

Prototype of a scaled-up microbial fuel cell for copper recovery.

Background: Bioelectrochemical systems (BESs) enable recovery of electrical energy through oxidation of a wide range of substrates at an anode and simultaneous recovery of metals at a cathode. Scale-up of BESs from the laboratory to pilot scale is a challenging step in the development of the process, and there are only a few successful experiences to build on. This paper presents a prototype BES for the recovery of copper. Results: The cell design presented here had removable electrodes, similar to those in electroplating baths. The anode and cathode in this design could be replaced independently. The prototype bioelectrochemical cell consisted of an 835 cm2 bioanode fed with acetate, and a 700 cm2 cathode fed with copper. A current density of 1.2 A/-2 was achieved with 48 mW m-2 of power production. The contribution of each component (anode, electrolytes, cathode and membrane) was evaluated through the analysis of the internal resistance distribution. This revealed that major losses occurred at the anode, and that the design with removable electrodes results in higher internal resistance compared with other systems. To further assess the practical applicability of BES for copper recovery, an economic evaluation was performed. Conclusion: Analysis shows that the internal resistance of several lab-scale BESs is already sufficiently low to make the system economic, while the internal resistance for scaled-up systems still needs to be improved considerably to become economically applicable.© 2017 The Authors. Journal of Chemical Technology & Biotechnology published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Gas diffusion electrodes improve hydrogen gas mass transfer for a hydrogen oxidizing bioanode.

Background: Bioelectrochemical systems (BESs) are capable of recovery of metals at a cathode through oxidation of organic substrate at an anode. Recently, also hydrogen gas was used as an electron donor for recovery of copper in BESs. Oxidation of hydrogen gas produced a current density of 0.8 A m-2 and combined with Cu2+ reduction at the cathode, produced 0.25 W m-2. The main factor limiting current production was the mass transfer of hydrogen to the biofilm due to the low solubility of hydrogen in the anolyte. Here, the mass transfer of hydrogen gas to the bioanode was improved by use of a gas diffusion electrode (GDE). Results: With the GDE, hydrogen was oxidized to produce a current density of 2.9 A m-2 at an anode potential of -0.2 V. Addition of bicarbonate to the influent led to production of acetate, in addition to current. At a bicarbonate concentration of 50 mmol L-1, current density increased to 10.7 A m-2 at an anode potential of -0.2 V. This increase in current density could be due to oxidation of formed acetate in addition to oxidation of hydrogen, or enhanced growth of hydrogen oxidizing bacteria due to the availability of acetate as carbon source. The effect of mass transfer was further assessed through enhanced mixing and in combination with the addition of bicarbonate (50 mmol L-1) current density increased further to 17.1 A m-2. Conclusion: Hydrogen gas may offer opportunities as electron donor for bioanodes, with acetate as potential intermediate, at locations where excess hydrogen and no organics are available. © 2017 The Authors. Journal of Chemical Technology & Biotechnology published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Competition between Methanogens and Acetogens in Biocathodes: A Comparison between Potentiostatic and Galvanostatic Control.

Microbial electrosynthesis is a useful form of technology for the renewable production of organic commodities from biologically catalyzed reduction of CO2. However, for the technology to become applicable, process selectivity, stability and efficiency need strong improvement. Here we report on the effect of different electrochemical control modes (potentiostatic/galvanostatic) on both the start-up characteristics and steady-state performance of biocathodes using a non-enriched mixed-culture inoculum. Based on our results, it seems that kinetic differences exist between the two dominant functional microbial groups (i.e., homoacetogens and methanogens) and that by applying different current densities, these differences may be exploited to steer product selectivity and reactor performance.

Fluidized capacitive bioanode as a novel reactor concept for the microbial fuel cell.

The use of granular electrodes in Microbial Fuel Cells (MFCs) is attractive because granules provide a cost-effective way to create a high electrode surface area, which is essential to achieve high current and power densities. Here, we show a novel reactor design based on capacitive granules: the fluidized capacitive bioanode. Activated carbon (AC) granules are colonized by electrochemically active microorganisms, which extract electrons from acetate and store the electrons in the granule. Electricity is harvested from the AC granules in an external discharge cell. We show a proof-of-principle of the fluidized capacitive system with a total anode volume of 2 L. After a start-up period of 100 days, the current increased from 0.56 A/m(2) with 100 g AC granules, to 0.99 A/m(2) with 150 g AC granules, to 1.3 A/m(2) with 200 g AC granules. Contact between moving AC granules and current collector was confirmed in a control experiment without biofilm. Contribution of an electro-active biofilm to the current density with recirculation of AC granules was limited. SEM images confirmed that a biofilm was present on the AC granules after operation in the fluidized capacitive system. Although current densities reported here need further improvement, the high surface area of the AC granules in combination with external discharge offers new and promising opportunities for scaling up MFCs.

Mechanisms of extracellular electron transfer in anaerobic methanotrophic archaea.

Anaerobic methanotrophic (ANME) archaea are environmentally important, uncultivated microorganisms that oxidize the potent greenhouse gas methane. During methane oxidation, ANME archaea engage in extracellular electron transfer (EET) with other microbes, metal oxides, and electrodes through unclear mechanisms. Here, we cultivate ANME-2d archaea ('Ca. Methanoperedens') in bioelectrochemical systems and observe strong methane-dependent current (91-93% of total current) associated with high enrichment of 'Ca. Methanoperedens' on the anode (up to 82% of the community), as determined by metagenomics and transmission electron microscopy. Electrochemical and metatranscriptomic analyses suggest that the EET mechanism is similar at various electrode potentials, with the possible involvement of an uncharacterized short-range electron transport protein complex and OmcZ nanowires.

Comparing theoretical and practical biomass yields calls for revisiting thermodynamic growth models for electroactive microorganisms.

Research on electroactive microorganisms (EAM) often focuses either on their physiology and the underlying mechanisms of extracellular electron transfer or on their application in microbial electrochemical technologies (MET). Thermodynamic understanding of energy conversions related to growth and activity of EAM has received only a little attention. In this study, we aimed to prove the hypothesized restricted energy harvest of EAM by determining biomass yields by monitoring growth of acetate-fed biofilms presumably enriched in Geobacter, using optical coherence tomography, at three anode potentials and four acetate concentrations. Experiments were concurrently simulated using a refined thermodynamic model for EAM. Neither clear correlations were observed between biomass yield and anode potential nor acetate concentration, albeit the statistical significances are limited, mainly due to the observed experimental variances. The experimental biomass yield based on acetate consumption (YX/ac = 37 ± 9 mgCODbiomass gCODac-1) was higher than estimated by modeling, indicating limitations of existing growth models to predict yields of EAM. In contrast, the modeled biomass yield based on catabolic energy harvest was higher than the biomass yield from experimental data (YX/cat = 25.9 ± 6.8 mgCODbiomass kJ-1), supporting restricted energy harvest of EAM and indicating a role of not considered energy sinks. This calls for an adjusted growth model for EAM, including, e.g., the microbial electrochemical Peltier heat to improve the understanding and modeling of their energy metabolism. Furthermore, the reported biomass yields are important parameters to design strategies for influencing the interactions between EAM and other microorganisms and allowing more realistic feasibility assessments of MET.

Starvation combined with constant anode potential triggers intracellular electron storage in electro-active biofilms.

The accumulation of electrons in the form of Extracellular Polymeric Substances (EPS) and poly-hydroxyalkanoates (PHA) has been studied in anaerobic processes by adjusting the access of microorganisms to the electron donor and final electron acceptor. In Bio-electrochemical systems (BESs), intermittent anode potential regimes have also recently been used to study electron storage in anodic electro-active biofilms (EABfs), but the effect of electron donor feeding mode on electron storage has not been explored. Therefore, in this study, the accumulation of electrons in the form of EPS and PHA was studied as a function of the operating conditions. EABfs were grown under both constant and intermittent anode potential regimes and fed with acetate (electron donor) continuously or in batch. Confocal Laser Scanning Microscopy (CLSM) and Fourier-Transform Infrared Spectroscopy (FTIR) were used to assess electron storage. The range of Coulombic efficiencies, from 25 to 82%, and the biomass yields, between 10 and 20%, indicate that storage could have been an alternative electron consuming process. From image processing, a 0.92 pixel ratio of poly-hydroxybutyrate (PHB) and amount of cells was found in the batch fed EABf grown under a constant anode potential. This storage was linked to the presence of living Geobacter and shows that energy gain and carbon source starvation were the triggers for intracellular electron storage. The highest EPS content (extracellular storage) was observed in the continuously fed EABf under an intermittent anode potential, showing that constant access to electron donor and intermittent access to the electron acceptor leads to the formation of EPS from the excess energy gained. Tailoring operating conditions can thus steer the microbial community and result in a trained EABf to perform a desired biological conversion, which can be beneficial for a more efficient and optimized BES.

Capacitive bioanodes enable renewable energy storage in microbial fuel cells.

We developed an integrated system for storage of renewable electricity in a microbial fuel cell (MFC). The system contained a capacitive electrode that was inserted into the anodic compartment of an MFC to form a capacitive bioanode. This capacitive bioanode was compared with a noncapacitive bioanode on the basis of performance and storage capacity. The performance and storage capacity were investigated during polarization curves and charge-discharge experiments. During polarization curves the capacitive electrode reached a maximum current density of 1.02 ± 0.04 A/m(2), whereas the noncapacitive electrode reached a current density output of only 0.79 ± 0.03 A/m(2). During the charge-discharge experiment with 5 min of charging and 20 min of discharging, the capacitive electrode was able to store a total of 22,831 C/m(2), whereas the noncapacitive electrode was only able to store 12,195 C/m(2). Regarding the charge recovery of each electrode, the capacitive electrode was able to recover 52.9% more charge during each charge-discharge experiment compared with the noncapacitive electrode. The capacitive electrode outperformed the noncapacitive electrode throughout each charge-discharge experiment. With a capacitive electrode it is possible to use the MFC simultaneously for production and storage of renewable electricity.

Opportunities for visual techniques to determine characteristics and limitations of electro-active biofilms.

Optimization of bio-electrochemical systems (BESs) relies on a better understanding of electro-active biofilms (EABfs). These microbial communities are studied with a range of techniques, including electrochemical, visual and chemical techniques. Even though each of these techniques provides very valuable and wide-ranging information about EABfs, such as performance, morphology and biofilm composition, they are often destructive. Therefore, the information obtained from EABfs development and characterization studies are limited to a single characterization of EABfs and often limited to one time point that determines the end of the experiment. Despite being scarcer and not as commonly reported as destructive techniques, non-destructive visual techniques can be used to supplement EABfs characterization by adding in-situ information of EABfs functioning and its development throughout time. This opens the door to EABfs monitoring studies that can complement the information obtained with destructive techniques. In this review, we provide an overview of visual techniques and discuss the opportunities for combination with the established electrochemical techniques to study EABfs. By providing an overview of suitable visual techniques and discussing practical examples of combination of visual with electrochemical methods, this review aims at serving as a source of inspiration for future studies in the field of BESs.

Maximum thickness of non-buffer limited electro-active biofilms decreases at higher anode potentials.

The accumulation of protons in electro-active biofilms (EABfs) has been reported as a critical parameter determining produced currents at the anode since the very beginning of the studies on Bio-electrochemical systems (BESs). Even though the knowledge gained on the influence of this parameter on the produced currents, its influence on EABfs growth is frequently overlooked. In this study, we quantified EABfs thicknesses in real-time and related them to the produced current at three buffer concentrations, two anode potentials and two acetate concentrations. The thickest EABfs (80 µm) and higher produced currents (2.5 A.m-2) were measured when a 50 mM buffer concentration was used. By combining the measured EABfs thicknesses with the pH in the anolyte, a simple model was developed to identify buffer limitations. Buffer limited EABfs with thicknesses of 15 and 42 µm were identified at -0.3 V vs Ag/AgCl when 10 and 50 mM buffer concentrations were used, respectively. At -0.2 V vs Ag/AgCl, the thicknesses of buffer limited EABfs decreased to 13 and 20 µm, respectively. The model also estimated buffer and acetate diffusion rates in EABfs and allowed to determine the boundary between a buffer and acetate limited EABfs. The diffusion rates reported in this study and the definition of the boundary between buffer and acetate limited EABfs provide a powerful tool to avoid limitations, leading to higher produced currents at the anode.

Ammonia recovery from anaerobic digester centrate using onsite pilot scale bipolar membrane electrodialysis coupled to membrane stripping.

Ammonia recovery from centrate of an anaerobic digester was investigated using an onsite bipolar-electrodialysis (BP-ED) pilot scale plant coupled to two liquid/liquid membrane contactor (LLMC) modules. To investigate the process performance and robustness, the pilot plant was operated at varying current densities, load ratio (current to nitrogen loading), and in continuous and intermittent current (Donnan) mode. A higher load ratio led to higher total ammonium nitrogen (TAN, sum of ammonia and ammonium) removal efficiency, whereas the increase in the applied current did not have a significant impact the TAN removal efficiency. Continuous current application resulted in the higher TAN removal compared with the Donnan dialysis mode. The lowest specific energy consumption of 6.3 kWh kgN-1 was recorded in the Donnan mode, with the load ratio of 1.4, at 200 L h-1 flowrate and current density of 75 A m-2. Lower energy demand observed in the Donnan mode was likely due to the lower scaling and fouling of the ion exchange membranes. Nevertheless, scaling and fouling limited the operation of the BP-ED stack in all operational modes, which had to be interrupted by the daily cleaning procedures. The LLMC module enabled a highly selective recovery of ammonia as ammonium sulfate ((NH4)2SO4), with the concentration of ammonia ranging from 19 to 33 gN L-1. However, the analysis of per- and polyfluoroalkyl substances (PFASs) in the obtained (NH4)2SO4 product revealed the presence of 212-253 ng L-1 of 6:2 fluorotelomer sulfonate (FTS), a common substitute of legacy PFAS. Given the very low concentrations of 6:2 FTS (i.e., < 2 ng L-1) encountered in the concentrated stream, 6:2 FTS was likely released from the Teflon-based components in the sulfuric acid dosage line. Thus, careful selection of the pilot plant tubing, pumps and other components is required to avoid any risks associated with the PFAS presence and ensure safe use of the final product as fertilizer.

Effects of Current on the Membrane and Boundary Layer Selectivity in Electrochemical Systems Designed for Nutrient Recovery.

During electrochemical nutrient recovery, current and ion exchange membranes (IEM) are used to extract an ionic species of interest (e.g., ion) from a mixture of multiple ions. The species of interest (ion 1) has an opposing charge to the IEM. When ion 1 is extracted from the solution, the species fractions at the membrane and the adjunct boundary layers are affected. Hence, the species transport through the electrochemical system (ES) can no longer be described as electrodialysis-like. A dynamic state is observed in the compartments, where the ionic species are recovered. When the boundary layer-membrane interface is depleted, the IEM is at maximum current. If the ES is operated at a current higher than the maximum current, the fluxes of both ion 1 and other competing ions, with the same charge (ion 2), occur. This means, for example, ion 1 will be recovered, and the concentration of ion 2 will build up in time. Therefore, a steady state is never reached. Ideally, to prevent the effect of limiting current at the boundary layer-membrane interface, ES for nutrient recovery should be operated at low currents.

Methane-Dependent Extracellular Electron Transfer at the Bioanode by the Anaerobic Archaeal Methanotroph "Candidatus Methanoperedens".

Anaerobic methanotrophic (ANME) archaea have recently been reported to be capable of using insoluble extracellular electron acceptors via extracellular electron transfer (EET). In this study, we investigated EET by a microbial community dominated by "Candidatus Methanoperedens" archaea at the anode of a bioelectrochemical system (BES) poised at 0 V vs. standard hydrogen electrode (SHE), in this way measuring current as a direct proxy of EET by this community. After inoculation of the BES, the maximum current density was 274 mA m-2 (stable current up to 39 mA m-2). Concomitant conversion of 13CH4 into 13CO2 demonstrated that current production was methane-dependent, with 38% of the current attributed directly to methane supply. Based on the current production and methane uptake in a closed system, the Coulombic efficiency was about 17%. Polarization curves demonstrated that the current was limited by microbial activity at potentials above 0 V. The metatranscriptome of the inoculum was mined for the expression of c-type cytochromes potentially used for EET, which led to the identification of several multiheme c-type cytochrome-encoding genes among the most abundant transcripts in "Ca. Methanoperedens." Our study provides strong indications of EET in ANME archaea and describes a system in which ANME-mediated EET can be investigated under laboratory conditions, which provides new research opportunities for mechanistic studies and possibly the generation of axenic ANME cultures.

The effect of intermittent anode potential regimes on the morphology and extracellular matrix composition of electro-active bacteria.

Electro-active bacteria (EAB) can form biofilms on an anode (so-called bioanodes), and use the electrode as electron acceptor for oxidation of organics in wastewater. So far, bioanodes have mainly been investigated under a continuous anode potential, but intermittent anode potential has resulted in higher currents and different biofilm morphologies. However, little is known about how intermittent potential influences the electron balance in the anode compartment. In this study, we investigated electron balances of bioanodes at intermittent anode potential regimes. We used a transparent non-capacitive electrode that also allowed for in-situ quantification of the EAB using optical coherence tomography (OCT). We observed comparable current densities between continuous and intermittent bioanodes, and stored charge was similar for all the applied intermittent times (5 mC). Electron balances were further investigated by quantifying Extracellular Polymeric Substances (EPS), by analyzing the elemental composition of biomass, and by quantifying biofilm and planktonic cells. For all tested conditions, a charge balance of the anode compartment showed that more electrons were diverted to planktonic cells than biofilm. Besides, 27-43% of the total charge was detected as soluble EPS in intermittent bioanodes, whereas only 15% was found as soluble EPS in continuous bioanodes. The amount of proteins in the EPS of biofilms was higher for intermittent operated bioanodes (0.21 mg COD proteins mg COD biofilm-1) than for continuous operated bioanodes (0.05 mg COD proteins mg COD biofilm-1). OCT revealed patchy morphologies for biofilms under intermittent anode potential. Overall, this study helped understanding that the use of a non-capacitive electrode and intermittent anode potential deviated electrons to other processes other than electric current at the electrode by identifying electron sinks in the anolyte and quantifying the accumulation of electrons in the form of EPS.

Effective orthophosphate removal from surface water using hydrogen-oxidizing bacteria: Moving towards applicability.

Effective orthophosphate removal strategies are needed to counteract eutrophication and guarantee water quality. Previously, we established that hydrogen-oxidizing bacteria (HOB) have the ability to remove orthophosphate from artificial surface water. In the present study, we expand the application of the HOB orthophosphate removal strategy (1) to treat artificial surface water with low initial orthophosphate concentrations, (2) to treat real surface water and real wastewater effluent, and (3) to remove orthophosphate continuously. For synthetic surface water, irrespective of the initial concentration of 0.7, 0.5, 0.3, and 0.1 mg PO43--P/L, ultra-low concentrations (0.0058 ± 0.0028 mg PO43--P/L) were obtained. When artificial surface water was replaced by real surface water, without added nutrients or other chemicals, it was shown that over 90% orthophosphate could be removed within 30 min of operation in a batch configuration (0.031 ± 0.023 mg PO43--P/L). In continuous operation, orthophosphate removal from surface water left an average concentration of 0.040 ± 0.036 for 60 days, and the lowest orthophosphate concentration measured was 0.013 mg PO43-/L. Simultaneously, nitrate was continuously removed for 60 days below 0.1 mg/L. The ability to remove orthophosphate even under nitrogen limiting conditions might be related to the ability of HOB to fix nitrogen. This study brings valuable insights into the potential use of HOB biofilms for nutrient remediation and recovery.

Donnan Dialysis for scaling mitigation during electrochemical ammonium recovery from complex wastewater.

Inorganic scaling is often an obstacle for implementing electrodialysis systems in general and for nutrient recovery from wastewater specifically. In this work, Donnan dialysis was explored, to prevent scaling and to prolong operation of an electrochemical system for TAN (total ammonia nitrogen) recovery. An electrochemical system was operated with and without an additional Donnan dialysis cell, while being supplied with synthetic influent and real digested black water. For the same Load Ratio (nitrogen load vs applied current) while treating digested black water, the system operated for a period three times longer when combined with a Donnan cell. Furthermore, the amount of nitrogen recovered was higher. System performance was evaluated in terms of both TAN recovery and energy efficiency, at different Load Ratios. At a Load Ratio 1.3 and current density of 10 A m-2, a TAN recovery of 83% was achieved while consuming 9.7 kWh kgN-1.

Reduction of pH buffer requirement in bioelectrochemical systems.

Low pH buffer capacity of waste streams limits further development of bioelectrochemical systems (BES) because accumulation of protons potentially leads to acidification of the anodic biofilm. Here we introduce a system that makes it possible to recover alkalinity in an extra recovery compartment. The system consisted of this extra compartment which was located between anode and cathode compartment. The compartment was separated from the anode by a cation exchange membrane and from the cathode by an anion exchange membrane, which made clean hydrogen production possible. To compensate for the charge movement as a result of the flow of electrons, both cations and hydroxyl ions moved into the new recovery compartment. When a synthetic waste stream was fed through this recovery compartment, both pH and conductivity increased. When this stream is then fed to the anode of the BES, no additional buffer was required to produce the same current (3.5 A/m(2)) at an applied voltage of 1 V.

New applications and performance of bioelectrochemical systems.

Bioelectrochemical systems (BESs) are emerging technologies which use microorganisms to catalyze the reactions at the anode and/or cathode. BES research is advancing rapidly, and a whole range of applications using different electron donors and acceptors has already been developed. In this mini review, we focus on technological aspects of the expanding application of BESs. We will analyze the anode and cathode half-reactions in terms of their standard and actual potential and report the overpotentials of these half-reactions by comparing the reported potentials with their theoretical potentials. When combining anodes with cathodes in a BES, new bottlenecks and opportunities arise. For application of BESs, it is crucial to lower the internal energy losses and increase productivity at the same time. Membranes are a crucial element to obtain high efficiencies and pure products but increase the internal resistance of BESs. The comparison between production of fuels and chemicals in BESs and in present production processes should gain more attention in future BES research. By making this comparison, it will become clear if the scope of BESs can and should be further developed into the field of biorefineries.