Gralnick Lab Publications


Zhao, F., M.S. Chavez, K.L. Naughton, C.M. Niman, J.T. Atkinson, J.A. Gralnick, M.Y. El-Naggar and J.Q. Boedicker. 2022. Light-induced patterning of electroactive bacterial biofilms. ACS Syn Bio. doi: 10.1021/acssynbio.2c00024.

Bhattarai, S., H. Temme, A. Jain, J.P. Badalamenti, JA. Gralnick and P.J. Novak. 2022. The potential for bacteria from carbon-limited deep terrestrial environments to participate in chlorine cycling. FEMS Microbiol Ecol. 98(6):fiac054. doi: 10.1093/femsec/fiac054.

Baker, I.R., B.E. Conley, J.A. Gralnick and P.R. Girguis. 2022. Evidence for horizontal and vertical transmission of Mtr-mediated extracellular electron transfer among the Bacteria. mBio. 13(1):e0290421. doi: 10.1128/mbio.02904-21.

2021 Bird, L.J., L. Su, B.B. Kundu, A.D. Corts, J.A. Gralnick, C.M. Ajo-Franklin and S.M. Glaven. 2021. Engineering Wired Life: Synthetic biology for electroactive bacteria. ACS SynBio. 10(11):2808-2823. doi: 10.1021/acssynbio.1c00335. 

Jain, A., B.M. Bonis and J.A. Gralnick. 2021. Oligo-heterotrophic activity of Marinobacter subterrani creates an indirect Fe(II)-oxidation phenotype in gradient tubes. Appl Environ Microbiol. 87(24):e0136721. doi: 10.1128/AEM.01367-21.

Boedicker, J.Q., M. Gangan, K. Naughton, F. Zhao, J.A. Gralnick and M.Y. El-Naggar. 2021. Engineering biological electron transfer and redox pathways for nanoparticle synthesis. Bioelectricity. 3(2):126-135. doi: 10/1089/bioe.2021.0010. 

Jiménez Otero, F., G.L. Chadwick, M.D. Yates, R.L. Mickol, S.H. Saunders, S.M. Glaven, J.A. Gralnick, D.K. Newman, L.M. Tender, V.J. Orphan and D.R. Bond. 2021. Evidence of streamlined extracellular electron transfer pathway from biofilm structure, metabolic stratification, and long-rang electron transfer parameters. Appl Environ Microbiol. doi: 10.1128/AEM.00706-21.

Kane, A.L., R.E. Szabo and J.A. Gralnick. 2021. Engineering cooperation in an anaerobic co-culture. Appl Environ Microbiol. 87(11):e02852-20. doi: 11.1128/AEM.02852-20.

Kees, E.D, C.E. Levar, S.P. Miller, D.R. Bond, J.A. Gralnick and A.M. Dean. 2021. Survival of the first rather than the fittest in a Shewanella electrode biofilm. Commun Biol. 4(1):536. doi: 10.1038/s42003-021-02040-1.

Sheik, C.S., J.P. Badalamenti, J. Telling, D. Hsu, S.C. Alexander, D.R. Bond, J.A. Gralnick, B.S. Loller and B.M. Toner. 2021. Novel microbial groups drive productivity in an Archean iron formation. Front Microbiol. 12:627595. doi: 10.3389/fmicb.2021.627595.

Starwalt-Lee, R., M.Y. El-Naggar, D.R. Bond and J.A. Gralnick. 2021. Electrolocation? The evidence for redox-mediated taxis in Shewanella oneidensis. Mol Micro. doi: 10.1111/mmi.14647. 

Jain, A., and J.A. Gralnick. 2021. Engineering lithoheterotrophy in an obligate chemolithoautotrophic Fe(II) oxidizing bacterium. Sci Rep. 11(1):2165. doi: 10.1038/s41598-021-81412-3.


Conley, B.E., M.T. Weinstock, D.R. Bond and J.A. Gralnick. 2020. A hybrid extracellular electron transfer pathway enhances survival of Vibrio natriegens. App Environ Microbiol. 86(19):e01253-20. doi: 10.1128/AEM.01253-20.

Shaw D.R., A. Muhammad, K.P. Katuri, J.A. Gralnick, J. Reimann, R. Mesman, L. van Niftrik, M.S.M. Jetten and P.E. Saikaly. 2020. Extracellular electron transfer-dependent anaerobic oxidation of ammonium by anammox bacteria. Nat Commun. 11(1):2058. doi: 10.1038/s41467-020-16016-y.

Jain, A. and J.A. Gralnick. 2020. Evidence for auxiliary anaerobic metabolism in obligately aerobic Zetaproteobacteria. ISME J. doi: 10.1038/s41396-020-0586-6.


Nisbett, L.M., L. Binnenkade, B. Bacon, S. Hossain, N.J. Kotloski, E.D. Brutinel, R. Hartmann, K. Drescher, D.P. Arora, S. Muralidharan, K.M. Thormann, J.A. Gralnick and E.M. Boon. 2019. NosP signaling modulates the NO/H-NOX-mediated multicomponent c-Di-GMP network and biofilm formation in Shewanella oneidensis. Biochemistry. 58(48):4827-4841. doi: 10/1021/acs.biochem.9b00706.

Chadwick, G.L., F. Otero Jiménez, J.A. Gralnick, D.R. Bond and V.J. Orphan. 2019. NanoSIMS imaging reveals metabolic stratification within current-producing biofilms. Proc Natl Acad Sci USA. pil: 21912498. doi: 10.1073/pnas.1912498116.

Kees, E.D., A.R. Pendleton, C.M. Paquete, M.B. Arriola, A.L. Kane, N.J. Kotloski, P.J. Intile and J.A. Gralnick. 2019. Secreted flavin cofactors for anaerobic respiration of fumarate and urocanate by Shewanella oneidensis: cost and role. Appl Environ Microbiol. pil: AEM.00852-19. doi: 10.112/AEM.00852-19.

Corts, A.D., L.C. Thomason, R.T. Gill and J.A. Gralnick. 2019. Efficient and precise genome editing in Shewanella with recombineering and CRISPR/Cas9-mediated counter-selection. ACS Synth Biol. 8(8):1877-1889. doi: 10.1021/acssynbio.9b00188.

Bennett, B.D. and J.A. Gralnick. 2019. Mechanisms of toxicity by and resistance to ferrous iron in anaerobic systems. Free Radic Biol Med. pil: S0891-5849(18)32165-8. doi: 10.1016/j.freeradbiomed.2019.06.027. 

Kaplan, M., P. Subramanian, D. Ghosal, C.M. Oikonomou, S. Pirbadian, R. Starwalt-Lee, S.K. Mageswaran, D.R. Ortega, J.A. Gralnick, M.Y. El-Naggar and G.J. Jensen. 2019. In situ imaging of the bacterial flagellar motor disassembly and assembly process. EMBO J. pil: e100957. doi: 10.15252/embj.2018100957.

Corts, A.D., L.C. Thomason, R.T. Gill and J.A. Gralnick. 2019. A new recombineering system for precise genome-editing in Shewanella oneidensis strain MR-1 using single-stranded oligonucleotides. Sci Rep. 9(1):39. doi: 10.1038/s41598-018-37025-4.

Wang, Q., A.A. Jones III, J.A. Gralnick, L. Lin, C.R. Buie. 2019. Microfluidic dielectrophoresis illuminates the relationship between microbial cell envelope polarizability and electrochemical activity. Sci Adv. 5(1):eaat5664. doi: 10.1126/sciadv.aat5664.


Conley, B.E., P.J. Intile, D.R. Bond and J.A. Gralnick. 2018. Divergent Nrf-family proteins and MtrCAB homologs facilitate extracellular electron transfer in Aeromonas hydrophila. Appl Environ Microbiol. doi: 10.1128/AEM.02134-18.

Philips J., N. Van den Driessche, K. De Paepe, A. Prévoteau, J.A. Gralnick, J.B.A. Arends and K. Rabaey. 2018. A novel Shewanella isolate enhances corrosion by using metallic iron as the electron donor with fumarate as the electron acceptor. Appl Environ Microbiol. 84(20):e00154-18 doi: 10.1128/AEM.01154-18.

Bennett B.D., K.E. Redford and J.A. Gralnick. 2018. Survival of anaerobic Fe2+ stress requires the ClpXP protease. J Bacteriol. 200(8):e00671-17 doi: 10.1128/JB.00671-17.

Bennett, B.D., K.E. Redford and J.A. Gralnick. 2018. The MgtE homolog FicI acts as a secondary ferrous iron importer in Shewanella oneidensis strain MR-1. Appl Environ Microbiol. 84(6):e01245-17 doi: 10.1128/AEM0.1245-17.

Rowe, A.R., P. Rajeev, A. Jain, S. Pirbadian, A. Okamoto, J.A. Gralnick, M.Y. El-Naggar and K.H. Nealson. 2018. Tracking electron uptake from a cathode into Shewanella cells: implications for energy acquisition from solid-substrate electron donors. mBio. 9(1):e02203-17. doi: 10.1128/mBio.02203-17.


West, E.A., A. Jain and J.A. Gralnick. 2017. Engineering a native inducible expression system in Shewanella oneidensis to control extracellular electron transfer. ACS Synth Biol. doi: 10.1021/acssynbio.6b00349

Bursac, T., J.A. Gralnick and J. Gescher. 2017. Acetoin production via unbalanced fermentation in Shewanella oneidensis. Biotechnol Bioeng. 114(6):1283-1289. doi: 10.1002/bit.26243


Badalamenti, J.P., Z.M. Summers, C.H. Chan, J.A. Gralnick and D.R. Bond. 2016. Isolation and genomic characterization of ‘Desulfuromonas soudanensis WTL’, a metal- and electrode-respiring bacterium from anoxic deep subsurface brine. Front Microbiol. 7:913. doi:10.3389/fmicb.2016.00913

Hajimorad, M. and J.A. Gralnick. 2016. Towards enabling engineered microbial electronic systems RK2-based conjugal transfer system for Shewanella synthetic biology. Electron Lett. 52(6):426-428.

Kane, A.L., E.D. Brutinel, H. Joo, R. Maysonet Sanchez, C.M. VanDrisse, N.J. Kotloski and J.A. Gralnick. 2016. Formate metabolism in Shewanella oneidensis generates proton motive force and prevents growth without an electron acceptor. J Bacteriol. 198(8):1337-1346.

Kane, A.L., B. Al-Shayeb, P.V. Holec, S. Rajan, N.E. Le Mieux, S.C. Heinsch, S. Psarska, K.G. Aukema, C.A. Sarkar, E.A. Nater and J.A. Gralnick. 2016. Toward bioremediation of methylmercury using silica encapsulated Escherichia coli harboring the mer operon. PLoS ONE 11(1):e0147036.


Bennett, B.D., E.D. Brutinel and J.A. Gralnick. 2015. A ferrous iron exporter mediates iron resistance in Shewanella oneidensis MR-1. Appl Environ Microbiol. 81(22):7938-44.

Bonis, B.M. and J.A. Gralnick. 2015. Marinobacter subterrani, a genetically tractable neutrophillic Fe(II)-oxidizing strain isolated from the Soudan Iron Mine. Front Microbiol. 6:719.


Yang, H., E.W. Krumholz, E.D. Brutinel, N.P. Palani, M.J. Sadowsky, A.M. Odlyzko, J.A. Gralnick and I.G. Libourel. 2014. Genome-scale metabolic network validation of Shewanella oneidensis using transposon insertion frequency analysis. PLoS Comput Biol. 10(9):e1003848.

Webster, D.P., M.A. TerAvest, D.F. Doud, A. Chakravorty, E.C. Holmes, C.M. Radens, S. Sureka, J.A. Gralnick and L.T. Angenent. 2014. An arsenic-specific biosensor with genetically engineered Shewanella oneidensis in a bioelectrochemical system. Biosens Bioelectron. 62:320-4.

TerAvest, M.A., M.A. Rosenbaum, N.J. Kotloski, J.A. Gralnick and L.T. Angenent. 2014. Oxygen allows Shewanella oneidensis MR-1 to overcome mediator washout in a continuously-fed bioelectrochemical system. Biotechnol Bioeng. 111(4):692-9.





Brutinel, E.D., A.M. Dean and J.A. Gralnick. 2013. Description of a riboflavin biosynthetic gene variant prevalent in the phylum Proteobacteria. J Bacteriol. 195(24):5479-86.

Kane, A.L., D.R. Bond and J.A. Gralnick. 2013. Electrochemical analysis of Shewanella oneidensis engineered to bind gold electrodes. ACS Synth Biol. 2(2):93-101.

Summers, Z.M., J.A. Gralnick and D.R. Bond. 2013. Cultivation of an obligate Fe(II)-oxidizing lithoautotrophic bacterium using electrodes. mBio. 4(1) e00420-12.

Kotloski, N.J. and J.A. Gralnick. 2013. Flavin electron shuttles dominate extracellular electron transfer by Shewanella oneidensis. mBio. 4(1) e00553-12. 


Gralnick, J.A. 2012. On conducting electron traffic across the periplasm. Biochem Soc Trans. 40(6):1178-80.

Brutinel, E.D. and J.A. Gralnick. 2012. Preferential utilization of D-lactate by Shewanella oneidensis. Appl Environ Microbiol. 78(23)8474-6.

Brutinel, E.D. and J.A. Gralnick. 2012. Anomalies of the anaerobic TCA cycle in Shewanella oneidensis revealed by Tn-seq. Mol Microbiol. 86(2):273-83.

Coursolle, D. and J.A. Gralnick. 2012. Reconstruction of extracellular respiratory pathways for iron(III) reduction in Shewanella oneidensis strain MR-1. Front Microbiol. 3:56.

Liu, N., Y. Xu, S. Hossain, N. Huang, D. Coursolle, J.A. Gralnick and E.M. Boon. 2012. Nitric oxide regulation of cyclic di-GMP synthesis and hydrolysis in Shewanella woodyi. Biochemistry. 51(10):2087-99.

Brutinel, E.D. and J.A. Gralnick. 2012. Shuttling Happens: soluble flavin mediators of extracellular electron transfer in Shewanella. Appl Microbiol Biotechnol. 93(1):41-8.

Flynn, C.M., K.A. Hunt, J.A. Gralnick and F. Srienc. 2012. Construction and elementary model analysis of a metabolic model for Shewanella oneidensis MR-1. Biosystems. 107(2):120-8.





Pinzon, N.M., K.G. Aukema, J.A. Gralnick and L.P. Wackett. 2011. Nile red detection of bacterial hydrocarbons and ketones in a high-throughput format. mBio. 2(4):e00109-11.

Sundararajan, A., J. Kurowski, T. Yan, D.M. Klingeman, M.P. Joachimiak, J. Zhou, B. Naranjo, J.A. Gralnick and M.W. Fields. 2011. Shewanella oneidensis MR-1 sensory box protein involved in aerobic and anoxic growth. Appl Environ Microbiol. 77(13):4647-56.

Jiao, Y., F. Qian, Y. Li, G. Wang, C.W. Saltikov and J.A. Gralnick. 2011. Deciphering the electron transport pathway for graphene oxide reduction by Shewanella oneidensis MR-1. J Bacteriol. 193(14):3662-5.

Cruz-Garcia, C., A.E. Murray, J.L. Rodrigues, J.A. Gralnick, L.A. McCue, M.F. Romine, F.E. Löffler and J.M. Tiedje. 2011. Fnr (EtrA) acts as a fine-tuning regulator of anaerobic metabolism in Shewanella oneidensis MR-1. BMC Microbiol. Mar;11:64.

Ross D.E., J.M. Flynn, D.B. Baron, J.A. Gralnick and D.R. Bond. 2011. Towards electrosynthesis in Shewanella: energetics of reversing the Mtr pathway for reductive metabolism. PLoS One. Feb; 6(2):e16649.


Flynn J.M., D.E. Ross, K.A. Hunt, D.R. Bond and J.A. Gralnick. 2010. Engineering unbalanced fermentations by using engineered electrode-interfaced bacteria. mBio. Nov;1(5):e00190-10.
mBiosphere Comment

Covington, E.D., C.B., Gelbmann, N.J. Kotloski and J.A. Gralnick. 2010. An essential role for UshA in processing of extracellular flavin electron shuttles by Shewanella oneidensis. Mol Microbiol. Oct; 78(2):519-32.

Coursolle, D. and J.A. Gralnick. 2010. Modularity of the Mtr respiratory pathway of Shewanella oneidensis strain MR-1. Mol Microbiol. Aug; 77(4):995-1008.

Johnson, E.T., D.B. Baron, B. Naranjo, D.R. Bond, C. Schmidt-Dannert and J.A. Gralnick. 2010. Light-driven proton pumping in an engineered bacterium enhances survival and electricity production. Appl Environ Microbiol. Jul; 76(13):4123-9.
Celebrating Ethan’s Life

Sukovich, D.J., J.L Seffernick, J.E. Richman, K.A. Hunt, J.A. Gralnick and L.P. Wackett. 2010. Head-to-head hydrocarbon biosynthetic genes in Shewanella oneidensis strain MR-1. Appl Environ Microbiol. Jun; 76(12):3842-9.

Sukovich, D.J., J.L. Seffernick, J.E. Richman, J.A. Gralnick and L.P. Wackett. 2010. Widespread head-to-head hydrocarbon biosynthesis in bacteria. Appl Environ Microbiol. Jun; 76(12):3850-62.

Hunt, K.A., J.M. Flynn, B. Naranjo, I.D. Shikhare and J.A. Gralnick. 2010. Substrate-level phosphorylation is the primary source of energy conservation during anaerobic respiraton of Shewanella oneidensis strain MR-1. J Bacteriol. Jul; 192(13):3345-51.

Coursolle, D., D.B. Baron, D.R. Bond and J.A. Gralnick. 2010. The Mtr respiratory pathway is essential for reducing flavins and electrodes in Shewanella oneidensis. J Bacteriol. Jan; 192(2):467-74.


Boal A.K., J.C. Genereux, P.A. Sontz, J.A. Gralnick, D.K. Newman and J.K. Barton. Redox signaling between DNA repair proteins for efficient lesion detection. 2009. Proc Natl Acad Sci U S A. Aug 31.

Baron, D.B., E. Labelle, D. Coursolle, J.A. Gralnick and D.R. Bond. 2009. Electrochemical measurement of electron transfer kinetics by Shewanella oneidensis MR-1. J Biol Chem. Oct 16; 284(42):28865-73.


Hau, H.H., A. Gilbert, D. Coursolle and J.A. Gralnick. 2008. Mechanism and Consequences of anaerobic respiration of cobalt by Shewanella oneidensis strain MR-1. Appl Environ Mircobiol. Nov;74(22):6880-6.

Marsili, E., D.B. Baron, I.D. Shikhare, D. Coursolle, J.A. Gralnick and D.R. Bond. 2008. Shewanella secretes flavins that mediate extracellular electron transfer. Proc Natl Acad Sci U S A. Mar 11; 105(10): 3968-3973.


Hau, H.H. and J.A. Gralnick. 2007. Ecology and biotechnology of the genus Shewanella. Ann Rev Microbiol. Jun 18; 61: 237 – 258.

Gralnick, J.A. and D.K. Newman. 2007. Extracellular Respiration. Mol Microbiol. July; 65(1): 1-11.


Gralnick, J.A., H. Vali, D.L. Lies and D.K. Newman. 2006. Extracellular respiration of dimethyl sulfoxide by Shewanella oneidensis strain MR-1. Proc Natl Acad Sci U S A. Mar 21; 103(12): 4669-4674.


Newman, D.K. and J.A. Gralnick. 2005. What genetics offers geobiology. Reviews in Mineralogy and Geochemistry. V. 59; p. 9-26.

Kappler A., Emerson D., Edwards K., Amend J.P., Gralnick J.A., Grathwohl P., Hoehler T., Straub K.L., 2005. Microbial activity in biogeochemical gradients - new aspects of research. Geobiology, 3, 229-233.