Peer reviewed publications, Preprints and PhD theses


Richter A, Hölscher T, Pausch P, Sehrt T, Brockhaus F, Bange G, Kovács ÁT (2018) Hampered motility promotes the evolution of wrinkly phenotype in Bacillus subtilisbioRxiv 288951 ( [in review]

Peer Reviewed Publications

2018 (5)

Dragoš A, Kiesewalter HT, Martin M, Hsu CY, Hartmann R, Wechsler T, Drescher K, Stanley-Wall N, Kümmerli R, Kovács ÁT (2018) Division of labor during biofilm matrix production. Current Biology [accepted]; preprint version available on bioRxiv 237230 (

Hölscher T, Schiklang T, Dragoš A, Dietel AK, Kost C, Kovács ÁT (2018) Impaired competence in flagellar mutants of Bacillus subtilis is connected to the regulatory network governed by DegU. Environmental Microbiology Reports 10(1):23-32 (; preprint version available on bioRxiv 194027 (

Dragoš A, Lakshmanan N, Martin M, Horváth B, Maróti G, Falcón García C, Lieleg O, Kovács ÁT (2018) Evolution of exploitative interactions during diversification in Bacillus subtilis biofilms. FEMS Microbiology Ecology 94(1):fix155 (; preprint version available on bioRxiv173476 (

Raie D, Mhatre E, El-Desouki D, Labena A, El-Ghannam G, Farahat L, Youssef T, Fritzsche W, Kovács ÁT (2018) Effect of novel quercetin titanium dioxide-decorated multi-walled carbon nanotubes nanocomposite on Bacillus subtilis biofilm development. Materials 11(1):157 (

Tauber JP, Gallegos-Monterrosa R, Kovács ÁT, Shelest E, Hoffmeister D (2018) Dissimilar pigment regulation in Serpula lacrymans and Paxillus involutus during inter-kingdom interactions. Microbiology164(1):65-77 ( 

2017 (11)

van den Esker M, Kovács ÁT, Kuipers OP (2017) From cell death to metabolism: holin-antiholin homologues with new functions. mBio 8(6):e01963-17 (

Kovács ÁT, Grau R, Pollitt EJG (2017) Surfing of bacterial droplets: Bacillus subtilis sliding revisited. Proceedings of the National Academy of Sciences USA 114(42):E8802 (

Gallegos-Monterrosa R, Kankel S, Götze S, Barnett R, Stallforth P, Kovács ÁT (2017) Lysinibacillus fusiformis M5 induces increased complexity in Bacillus subtilis 168 colony biofilms via hypoxanthine signaling. Journal of Bacteriology 199(22): e00204-17 (; preprint version available on bioRxiv 118125 (

Dragoš A, Kovács ÁT, Claessen D (2017) The role of functional amyloids in multicellular growth and development of Gram-positive bacteria. Biomolecules 7(3):E60 (

Hölscher T, Kovács ÁT (2017) Sliding on the surface: bacterial spreading without an active motor. Environmental Microbiology 19(7):2537-2545 (

Martin M, Dragoš A, Hölscher T, Maroti G, Balint B, Westermann M, Kovács ÁT (2017) De novo evolved interference competition promotes the spread of biofilm defectors. Nature Communications 8:15127 (

Mhatre E, Sundaram A, Hölscher T, Mühlstädt M, Bossert J, Kovács ÁT (2017) Presence of calcium lowers the expansion of Bacillus subtilis colony biofilms. Microorganisms 5(10):7 (

Dragoš A, Kovács ÁT (2017) The peculiar functions of bacterial extracellular matrix. Trends in Microbiology 25(4):257-266 (

Krawinkel J, Torres-Mapa ML, Mhatre E, Kovács ÁT, Heisterkamp A (2017) Structural damage of Bacillus subtilis biofilms using pulsed laser interaction with gold thin films. Journal of Biophotonics 10(8):1043-1052 (

Raie D, Mhatre E, Thiele M, Labena AE, Al-Ghannam GA, Farahat LA, Youssef T, Fritzsche W, Kovács ÁT (2017) Application of quercetin and its bio-inspired nanoparticles as anti-adhesive agents against Bacillus subtilis attachment to surface. Materials Science and Engineering C 70:753-762 ( /j.msec.2016.09.038)

van den Esker M, Kovács ÁT, Kuipers OP (2017) YsbA and LytST are essential for pyruvate utilization in Bacillus subtilis. Environmental Microbiology 19(1):83-94 (

2016 (8)

Gallegos-Monterrosa R, Maróti G, Bálint B, Kovács ÁT (2016) Draft genome sequence of the soil isolate Lysinibacillus fusiformis M5, a potential hypoxanthine producer. Genome Announcements 4(6):e01272-16 ( 10.1128/genomeA.01272-16)

Gallegos-Monterrosa R, Mhatre E, Kovács ÁT (2016) Specific Bacillus subtilis 168 variants do form biofilms on nutrient rich medium. Microbiology 162(11):1922-1932 (

Seccareccia I, Kovács ÁT, Gallegos-Monterrosa R, Nett M (2016) Unraveling the predator-prey relationship of Cupriavidus necator and Bacillus subtilis. Microbiological Research 192:231-238 ( /j.micres.2016.07.007)

Hölscher T, Dragoš A, Gallegos-Monterrosa R, Martin M, Mhatre E, Richter A, Kovács ÁT (2016) Monitoring spatial segregation in surface colonizing microbial populations. Journal of Visualized Experiments 116: e54752, (

Martin M, Hölscher T, Dragoš A, Cooper VS, Kovács ÁT (2016) Laboratory evolution of microbial interactions in bacterial biofilms. Journal of Bacteriology 198(19):2564-2571 (

Mhatre E, Troszok A,  Gallegos-Monterrosa R, Lindstädt S, Hölscher T, Kuipers OP, Kovács ÁT (2016) The impact of manganese on biofilm development of Bacillus subtilis. Microbiology 162(8):1468-1478 (

Kovács ÁT (2016) The global regulator CodY is required for the fitness of Bacillus cereus in various laboratory media and certain beverages. FEMS Microbiology Letters 363(13):fnw126 (

Kovács ÁT (2016) Bacterial differentiation via gradual activation of global regulators. Current Genetics 62(1):125-128 (

2015 (5)

Hölscher T, Bartels B, Lin Y-C, Gallegos-Monterrosa R, Price-Whelan A, Kolter R, Dietrich LEP, Kovács ÁT (2015) Motility, chemotaxis and aerotaxis contribute to competitiveness during bacterial pellicle biofilm development. Journal of Molecular Biology 427(23):3695-3708. (
Article recommended in F1000Prime (

Grau RG, de Oña P, Kunert M, Leñini C, Gallegos-Monterrosa R, Mhatre E, Vileta D, Donato V, Hölscher T, Boland W, Kuipers OP, Kovács ÁT (2015) A duo of potassium-responsive histidine kinases govern the multicellular destiny of Bacillus subtilis. mBio 6(3):e00581-15. (

Benoit I, van den Esker MH; Patyshakuliyeva A, Mattern D, Blei F, Zhou M, Dijksterhuis J, Brakhage AA, Kuipers OP, de Vries RP, Kovács ÁT (2015) Bacillus subtilis attachment to Aspergillus niger hyphae results in mutually altered metabolism. Environmental Microbiology 17(6): 2099-2113 (

Schneider J, Klein T, Mielich-Süss B, Koch G, Franke C, Kuipers OP, Kovács ÁT, Sauer M, Lopez D (2015) Spatio-temporal remodeling of functional membrane microdomains organizes the signaling networks of a bacterium. PLoS Genetics 11(4): e1005140 (

Detert Oude Weme R, Kovács ÁT, de Jong S, Veening JW, Siebring J, Kuipers OP (2015) Single cell FRET analysis for the identification of optimal FRET-pairs in Bacillus subtilis using a prototype MEM-FLIM system. PLoS One 10(4): e0123239 (

2014 (7)

Kovács ÁT (2014) Impact of spatial distribution on the development of mutualism in microbes. Frontiers in Microbiology 5: 649. (

van Gestel J, Weissing FJ, Kuipers OP, Kovács ÁT (2014) Density of founder cells affects spatial pattern formation and cooperation in Bacillus subtilis biofilms. ISME Journal 8(10): 2069–2079. (

Mhatre E, Gallegos Monterrosa R, Kovács ÁT (2014) From environmental signals to regulators: modulation of biofilm development in Gram-positive bacteria. Journal of Basic Microbiology 54(7): 616-632. (

Puentes-Téllez PE, Kovács ÁT, Kuipers OP, van Elsas JD (2014) Comparative genomics and transcriptomics analysis of experimentally evolved Escherichia coli MC1000 in complex environments. Environmental Microbiology 16(3): 856-870. 

Irigül-Sönmez Ö, Köroğlu TE, Öztürk B, Kovács ÁT, Kuipers OP, Yazgan-Karataş A (2014) In Bacillus subtilis LutR is part of the global complex regulatory network governing the adaptation to the transition from exponential growth to stationary phase. Microbiology 160(2): 243-260.

Diethmaier C, Newman JA, Kovács ÁT, Kaever V, Herzberg C, Rodrigues C, Boonstra M,  Kuipers OP, Lewis RJ, Stülke J (2014) The YmdB phosphodiesterase is a global regulator of late adaptive responses in Bacillus subtilis. Journal of Bacteriology 196(2): 265-275.

Siebring J, Elema MJH, Drubi Vega F, Kovács ÁT, Haccou P, Kuipers OP (2014) Repeated triggering of sporulation in Bacillus subtilis selects against a protein that affects timing of cell division. ISME Journal 8(1): 77-87. 

2013 (3)

Overkamp W, Beilharz K, Detert Oude Weme R, Solopova A, Karsens H, Kovács ÁT, Kok J, Kuipers OP, Veening JW (2013) Benchmarking various GFP variants in Bacillus subtilis, Streptococcus pneumoniae and Lactococcus lactis for live cell imaging. Applied and Environmental Microbiology 79(20): 6481-6490.

Lehnik-Habrink M, Rempeters L, Kovács ÁT, Wrede C, Baierlein C, Krebber H, Kuipers OP, Stülke J (2013) The DEAD-box RNA helicases in Bacillus subtilis have multiple functions and act independent from each other. Journal of Bacteriology 195(3): 534-544.

Kovács ÁT, Eckhardt TH, van Kranenburg R, Kuipers OP (2013) Functional analysis of the ComK protein of Bacillus coagulans. PLOS One 8(1): e53471.

2012 (3)

Kovács ÁT, van Gestel J, Kuipers OP (2012) The protective layer of biofilm: a repellent function for a new class of amphiphilic proteins. Molecular Microbiology 85(1):8-11.

Fibriansah G, Kovács ÁT, Pool TJ, Boonstra M, Kuipers OP, Thunnissen AMWH (2012) Crystal structures of two transcriptional regulators from Bacillus cereus define the conserved structural features of a PadR subfamily. PLOS One 7(11):e48015

Lindbäck T, Mols M, Basset C, Granum PE, Kuipers OP, Kovács ÁT (2012) CodY, a pleiotropic regulator, influences multicellular behavior and efficient production of virulence factors in Bacillus cereus. Environmental Microbiology 14(8):2233-2246.

2011 (4)

Mironczuk AM, Manu A, Kuipers OP, Kovács ÁT (2011) Distinct roles of ComK1 and ComK2 in gene regulation in Bacillus cereus. PLOS One 6(7):e21859

Mellegård H, Kovács ÁT, Lindbäck T, Christensen BE, Kuipers OP, Granum PE (2011) Transcriptional responses of Bacillus cereus towards challenges with the polysaccharide chitosan. PLOS One 6(9):e24304

Kovács ÁT, Kuipers OP (2011) Rok regulates yuaB expression during architecturally complex colony development of Bacillus subtilis 168. Journal of Bacteriology 193(4): 998-1002

Abee T, Kovács ÁT, Kuipers OP, van der Veen S (2011) Biofilm formation and dispersal in Gram-positive bacteria. Current Opinion in Biotechnology 22(2): 172-179

2010 (1)

Kovács ÁT, van Hartskamp M, Kuipers OP, van Kranenburg R (2010) Genetic tool development for a new host for biotechnology, the thermotolerant bacterium Bacillus coagulans. Applied and Environmental Microbiology 76(12): 4085-4088

2009 (2)

Kovács ÁT, Smits WK, Mironczuk AM, Kuipers OP (2009) Ubiquitous late competence genes in Bacillus species indicate the presence of functional DNA uptake machineries. Environmental Microbiology 11(8): 1911-1922

Grande Burgos MJ*, Kovács ÁT*, Mironczuk AM, Abriouel H, Galvez A, Kuipers OP (2009) Response of Bacillus cereus ATCC 14579 to challenges with sublethal concentrations of enterocin AS-48. BMC Microbiology 9:227 *contributed equally

2008 (1)

Mironczuk AM, Kovács ÁT, Kuipers OP (2008) Induction of natural competence in Bacillus cereus ATCC14579. Microbial Biotechnology 1(3): 226-235. 

2005 (6)

Kovács ÁT, Rákhely G, Balogh J, Maróti G, Cournac L, Carrier P, Mészáros LS, Peltier G, Kovács KL (2005) Hydrogen independent expression of hupSL genes in Thiocapsa roseopersicina. FEBS Journal 272(18): 4807-4816.

Kovács ÁT, Rákhely G, Kovács KL (2005) The PpsR regulator family. (Mini-review) Research in Microbiology 156(5-6): 619-625.

Kovács ÁT, Rákhely G, Browning DF, Fülöp A, Maróti G, Busby SJW, Kovács KL (2005) An FNR-type regulator controls the anaerobic expression of Hyn hydrogenase in Thiocapsa roseopersicina. Journal of Bacteriology 187(8): 2618-2627.

Kovács ÁT, Rákhely G, Balogh J, Maróti G, Fülöp A, Kovács KL (2005) Anaerobic regulation of hydrogenase transcription in different bacteria. (Mini-review) Biochemical Society Transactions 33(1): 36-38.

Kovács KL, Kovács ÁT, Maróti G, Mészáros LS, Balogh J, Latinovics D, Fülöp A, Dávid R, Dorogházi E, Rákhely G (2005) The hydrogenases of Thiocapsa roseopersicina. Biochemical Society Transactions 33(1): 61-63.

Kovács KL, Kovács ÁT, Maróti G, Bagi Z, Csanádi G, Perei K, Bálint B, Balogh J, Fülöp A, Mészáros LS, Tóth A, Dávid R, Latinovics D, Varga A, Rákhely G (2005) Improvement of biohydrogen production and intensification of biogas formation. Reviews in Environmental Science and Biotechnology 3(4): 321-330.

2004 (2)

Rákhely G, Kovács ÁT, Maróti G, Fodor BD, Csanádi G, Latinovics D, Kovács KL (2004) A cyanobacterial type, heteropentameric NAD+ reducing [NiFe] hydrogenase in the purple sulfur photosynthetic bacterium, Thiocapsa roseopersicina. Applied and Environmental Microbiology 64(2): 722-728. (IF 3.8)

Fodor BD, Kovács ÁT, Csáki R, Hunyadi-Gulyás E, Klement E, Maróti G, Mészáros LS, Medzihradszky KF, Rákhely G, Kovács KL (2004) Modular broad-host-range expression vectors for single protein and protein complex purification. Applied and Environmental Microbiology 64(2): 712-721.

2003 (2)

Kovács ÁT, Rákhely G, Kovács KL (2003) Genes involved in the biosynthesis of photosynthetic pigments in the purple sulfur photosynthetic bacterium Thiocapsa roseopersicina. Applied and Environmental Microbiology 63(6): 3093-3102.

Maróti G, Fodor BD, Rákhely G, Kovács ÁT, Arvani S, Kovács KL (2003) Accessory proteins functioning selectively and pleiotropically in the biosynthesis of [NiFe] hydrogenases in Thiocapsa roseopersicina. European Journal of Biochemistry 270(10):2218-2227.

2002 (1)

Kovács KL, Fodor B, Kovács ÁT, Csanádi G, Maróti G, Balogh J, Arvani S, Rákhely G (2002) Hydrogenases, accessory genes and the regulation of [NiFe]  hydrogenase biosynthesis in Thiocapsa roseopersicina. International Journal of Hydrogen Energy 27:1463-1469.

2001 (1)

Fodor B, Rákhely G, Kovács ÁT, Kovács KL (2001) Transposon mutagenesis in purple sulfur photosynthetic bacteria: identification of hypF, encoding a protein capable of processing [NiFe] hydrogenases in alpha, beta, and gamma subdivisions of the proteobacteria. Applied and Environmental Microbiology 61(6): 2476-2483.

1999 (1)

Dahl C, Rákhely G, Pott AS, Fodor B, Takács M, Tóth A, Kraeling M, Győrfi K, Kovács Á, Tusz J, Kovács KL (1999) Genes involved in hydrogen and sulfur metabolism in phototrophic sulfur bacteria. FEMS Microbiology Letters 180(2):317-324.


PhD Theses

Theresa Hölscher (thesis submitted) Social interactions and regulatory pathways influencing Bacillus subtilis biofilm formation and motility

Ramses Gallegos Monterrosa (2018) Cellular interactions in single- and mixed-species biofilms of Bacillus subtilis

Eisha Mhatre (2017) Extrinsic and intrinsic factors governing bacterial biofilms


Ákos T. Kovács
DTU Bioengineering
+45 45 25 25 27

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