Humboldt-Universität zu Berlin - Institut für Biologie


Prof. Dr. Athina Zouni 




Golub, J. Gätcke, S. Subramanian, A. Kölsch, T. Darwish, J. K. Howard, A. Feoktystov, O. Matsarskaia, A. Martel, L. Porcar, A. Zouni, and J. Pieper*, “Invisible” Detergents Enable a Reliable Determination of Solution Structures of Native Photosystems by Small-Angle Neutron Scattering J. Phys. Chem. B 2022, 126, 15, 2824–2833




Thomas Fransson, Roberto Alonso-Mori, Ruchira Chatterjee, Mun Hon Cheah, Mohamed Ibrahim, Rana Hussein, Miao Zhang, Franklin Fuller, Sheraz Gul, In-Sik Kim, Philipp S. Simon, Isabel Bogacz, Hiroki Makita, Casper de Lichtenberg, Sanghoon Song, Alexander Batyuk, Dimosthenis Sokaras, Ramzi Massad, Margaret Doyle, Alexander Britz, Clemens Weninger, Athina Zouni, Johannes Messinger, Vittal K. Yachandra, Junko Yano, Jan Kern, and Uwe Bergmann, Effects of x-ray free-electron laser pulse intensity on the Mn Kβ1,3 x-ray emission spectrum in photosystem II—A case study for metalloprotein crystals and solutions Structural Dynamics 8, 064302 (2021)


Stephen M. Keable, Adrian Kölsch, Philipp S. Simon, Medhanjali Dasgupta, Ruchira Chatterjee, Senthil Kumar Subramanian, Rana Hussein, Mohamed Ibrahim, In-Sik Kim, Isabel Bogacz, Hiroki Makita, Cindy C. Pham, Franklin D. Fuller, Sheraz Gul, Daniel Paley, Louise Lassalle, Kyle D. Sutherlin, Asmit Bhowmick, Nigel W. Moriarty, Iris D. Young, Johannes P. Blaschke, Casper de Lichtenberg, Petko Chernev, Mun Hon Cheah, Sehan Park, Gisu Park, Jangwoo Kim, Sang Jae Lee, Jaehyun Park, Kensuke Tono, Shigeki Owada, Mark S. Hunter, Alexander Batyuk, Roland Oggenfuss, Mathias Sander, Serhane Zerdane, Dmitry Ozerov, Karol Nass, Henrik Lemke, Roman Mankowsky, Aaron S. Brewster, Johannes Messinger, Nicholas K. Sauter, Vittal K. Yachandra, Junko Yano, Athina Zouni & Jan Kern, Room temperature XFEL crystallography reveals asymmetry in the vicinity of the two phylloquinones in photosystem I Scientific Reports 11, Article number: 21787 (2021)


Nikoloz Nioradze, Dmitri Ciornii, Adrian Kölsch, Gero Göbel, Dimitri E. Khoshtariya, Athina Zouni, Fred Lisdat, Electrospinning for building 3D structured photoactive biohybrid electrodes Bioelectrochemistry 142 (2021) 107945


Mohamed Ibrahim, Nigel W. Moriarty, Jan Kern, James M. Holton, Aaron S. Brewster, Asmit Bhowmick, Uwe Bergmann, Athina Zouni, Johannes Messinger, Vittal K. Yachandra, Junko Yano, Holger Dobbek, Nicholas K. Sauter, and Paul D. Adams, Reply to Wang et al.: Clear evidence of binding of Ox to the oxygen-evolving complex of photosystem II is best observed in the omit map, Proceedings of the National Academy of Sciences 2021, 118 (24)


Sascha Morlock,* Senthil K. Subramanian, Athina Zouni, and Fred Lisdat, Scalable Three-Dimensional Photobioelectrodes Made of Reduced Graphene Oxide Combined with Photosystem I, ACS Applied Materials & Interfaces 2021 13 (9), 11237-11246

DOI: 10.1021/acsami.1c01142


Maksym Golub, Adrian Kölsch, Artem Feoktystov, Prof. Dr. Athina Zouni and Jörg Pieper, Insights into Solution Structures of Photosynthetic Protein Complexes from Small-Angle Scattering Methods, Crystals 11(2), 203 2021




Maksym Golub, Rana Hussein, Mohamed Ibrahim, Max Hecht, Dietmar Christian, Florian Wieland, Anne Martel, Barbara Machado, Athina Zouni, and Jörg Pieper , Solution Structure of the Detergent-Photosystem II Core Complex Investigated by Small Angle Scattering Techniques,  J. Phys. Chem. B, Just Accepted Manuscript,  Publication Date (Web): 20 Aug 2020


Kölsch, A., Radon, C., Golub, M., Baumert, A., Bürger, J., Mielke, T., Lisdat,F., Feoktystov, A., Pieper, J., Zouni, A., Wendler, P., Current limits of structural biology: the transientinteraction between cytochrome c6 and photosystem I, Current Research in Structural Biology,


Ibrahim, M.;  Fransson, T.;  Chatterjee, R.;  Cheah, M. H.;  Hussein, R.;  Lassalle, L.;  Sutherlin, K. D.;  Young, I. D.;  Fuller, F. D.;  Gul, S.;  Kim, I.-S.;  Simon, P. S.;  de Lichtenberg, C.;  Chernev, P.;  Bogacz, I.;  Pham, C. C.;  Orville, A. M.;  Saichek, N.;  Northen, T.;  Batyuk, A.;  Carbajo, S.;  Alonso-Mori, R.;  Tono, K.;  Owada, S.;  Bhowmick, A.;  Bolotovsky, R.;  Mendez, D.;  Moriarty, N. W.;  Holton, J. M.;  Dobbek, H.;  Brewster, A. S.;  Adams, P. D.;  Sauter, N. K.;  Bergmann, U.;  Zouni, A.;  Messinger, J.;  Kern, J.;  Yachandra, V. K.; Yano, J., Untangling the sequence of events during the S<sub>2</sub> → S<sub>3</sub> transition in photosystem II and implications for the water oxidation mechanism. Proceedings of the National Academy of Sciences 2020, 117 (23), 12624-12635.


Müh, F.; Zouni, A., Structural basis of light‐harvesting in the photosystem II core complex. Protein Science 2020, 29 (5), 1090-1119.


Gerland, L.;  Friedrich, D.;  Hopf, L.;  Donovan, E. J.;  Wallmann, A.;  Erdmann, N.;  Diehl, A.;  Bommer, M.;  Buzar, K.;  Ibrahim, M.;  Schmieder, P.;  Dobbek, H.;  Zouni, A.;  Bondar, A.-N.;  Dau, H.; Oschkinat, H., pH‐Dependent Protonation of Surface Carboxylate Groups in PsbO Enables Local Buffering and Triggers Structural Changes. ChemBioChem 2020.




Cheah, M. H.; Zhang, M.; Shevela, D.; Mamedov, F.; Zouni, A.; Messinger, J., Assessment of the manganese cluster’s oxidation state via photoactivation of photosystem II microcrystals. Proceedings of the National Academy of Sciences 2019, 201915879.


Kemmler, L.; Ibrahim, M.; Dobbek, H.; Zouni, A.; Bondar, A.-N., Dynamic water bridging and proton transfer at a surface carboxylate cluster of photosystem II. Physical Chemistry Chemical Physics 2019, 21 (45), 25449-25466.


Sidabras, J. W.; Duan, J.; Winkler, M.; Happe, T.; Hussein, R.; Zouni, A.; Suter, D.; Schnegg, A.; Lubitz, W.; Reijerse, E. J., Extending electron paramagnetic resonance to nanoliter volume protein single crystals using a self-resonant microhelix. Science advances 2019, 5 (10).

Kerna, J.; Muhb, F.; Zouni, A., Structural studies on tetrapyrrole containing proteins enabled by femtosecond X-ray pulses. Metabolism, Structure and Function of Plant Tetrapyrroles: Control Mechanisms of Chlorophyll Biosynthesis and Analysis of Chlorophyll-Binding Proteins 2019, 33.


Ruocheng Han, Katharina Rempfer, Miao Zhang, Prof. Dr. Holger Dobbek, Prof. Dr. Athina Zouni, Prof. Dr. Holger Dau, Prof. Dr. Sandra Luber Investigating the Structure and Dynamics of Apo‑Photosystem II ChemCatChem, 21 June 2019,


Chatterjee R, Lasalle L, Gul S, Fuller FD, Young ID, Ibrahim M, de Lichtenberg C, Cheah MH, Zouni A, Messinger J, Yachandra VK, Kern J, Yano J Structural isomers of the S2 state in photosystem II: do they exist at room temperature and are they important for function? Physiol Plant. 2019 May; 166(1):60-72. Epub 2019 Mar 15


Ciornii D, Kölsch A, Zouni A, Lisdat F. Exploiting new ways for a more efficient orientation and wiring of PSI to electrodes: A fullerene C70 approach. Electrochimica Acta. Volume 299, 10 March 2019, Pages 531-539.



Riedel M, Wersig J, Ruff A, Schuhmann W, Zouni A, Lisdat F. A Z-Scheme-Inspired Photobioelectrochemical H2 O/O2 Cell with a 1V Open-Circuit Voltage Combining PhotosystemII and PbS Quantum Dots. Angew Chem Int Ed Engl. 2019 Jan 14;58(3):801-805. doi: 10.1002/anie.201811172. Epub 2018 Dec 13.


Kern J, Chatterjee R, Young ID, FD Fuller, Lassalle L, Ibrahim M, Gul S, Fransson T; Brewster AS, Alonso-Mori R, Hussein R, Zhang M, Douthit L, de Lichtenberg C, Cheah MH, Shevela D, Wersig J, Seuffert I, Sokaras D, Pastor E, Weninger Cl, Kroll T, … , Dobbek H, Adams PD, Bergmann U, Sauter NK, Zouni A, Messinger J, Yano J and Yachandra VK. Structures of the intermediates of Kok’s photosynthetic water oxidation clock. Nature volume 563, pages 421–425 (2018).


Feifel SC, Stieger KR, Hejazi M, Wang X, Ilbert M, Zouni A, Lojou E, Lisat F. Dihemic c 4 -type cytochrome acting as a surrogate electron conduit: Artificially interconnecting a photosystem I supercomplex with electrodes. Electrochemistry Communications 91, June 2018 Pages 49-53. DOI: 10.1016/j.elecom.2018.05.006


Kölsch A, Hejazi M, Stieger KR, Feifel SC, Kern JF, Müh F, Lisdat F, Lokstein H,  Zouni A. Insights into the binding behavior of native and non-native cytochromes to Photosystem I from Thermosynechococcus elongates. J Biol Chem. 2018 Jun 8;293(23):9090-9100. doi: 10.1074/jbc.RA117.000953. Epub 2018 Apr 25.


Stieger K, Feifel SC, Lokstein H, Hejazi M, Zouni A, Lisdat. Biohybride Architekturen für eine effiziente Umwandlung von Licht in elektrische Energie durch Integration von Photosystem I in skalierbare mesoporöse 3D Elektroden. Wiss. Beitr. TH Wildau 22: 17-24. Mar 2018.



Yachandra V, Kern J, Zouni A, Messinger J, Bergmann U, Alonso-Mori R, Wernet P, Sauter N, Yano J. Taking snapshots of photosynthetic water oxidation with an X-ray laser. Acta Crystallographica Section A: Foundations and Advances 73(a2):C14-C14 · December 2017. DOI: 10.1107/S2053273317095560.


Lassalle, Young ID, Ibrahim M, Chatterjee R, Gul S, Fuller F, Brewster A, Douthit L, Pastor E, Sauter N, Zouni A, Kern J, Yachandra V and Yano J. Exploring the dynamic of PSII at room temperature by simultaneous femtosecond X-ray spectroscopy and diffraction. Acta Cryst. (2017). A73, a114.


Ciornii D, Riedel M, Stieger KR, Feifel SC, Hejazi M, Lokstein H, Zouni A, Lisdat F. Bioelectronic Circuit on a 3D Electrode Architecture: Enzymatic Catalysis Interconnected with Photosystem I. J Am Chem Soc. 2017 Nov 22;139(46):16478-16481. doi: 10.1021/jacs.7b10161. Epub 2017 Nov 14.


Hussein R, Mohamed Ibrahim M, Chatterjee R, Coates L, Müh F, Vittal K. Yachandra V.K., Yano J, Jan Kern J, Dobbek H, and Zouni A. Optimizing Crystal Size of Photosystem II by Macroseeding: Toward Neutron Protein Crystallography. Cryst. Growth Des., Article ASAP DOI: 10.1021/acs.cgd.7b00878 Publication Date (Web): November 13, 2017.


Bommer M, Coates L, Dau H, Zouni A, Dobbek H. Protein crystallization and initial neutron diffraction studies of the photosystem II subunit PsbO. Acta Crystallogr F Struct Biol Commun. 2017 Sep 1;73(Pt 9):525-531. doi: 10.1107/S2053230X17012171. Epub 2017 Aug 31.


Zhang M, Bommer M, Chatterjee R, Hussein R, Yano J, Dau H, Kern J, Dobbek H, Zouni A. Structural insights into the light-driven auto-assembly process of the water-oxidizing Mn4CaO5-cluster in photosystem II. Elife. 2017 Jul 18;6. pii: e26933. doi 10.7554/eLife.26933


Skandary S, Müh F, Ashraf I, Ibrahim M, Metzger M, Zouni A, Meixner AJ, and Brecht M. Role of missing carotenoid in reducing the fluorescence of single monomeric photosystem II core complexes. Phys Chem Chem Phys. 2017 May 24;19(20):13189-13194. doi: 10.1039/c6cp07748j.!divAbstract


Dmitri Ciornii, Feifel SC, Hejazi M, Kölsch A, Lokstein H, Zouni A, and Lisdat F. Construction of photobiocathodes using multi-walled carbon nanotubes and photosystem I. PSS. Phys. Status Solidi A 214, No. 9, 1700017 (2017) / DOI 10.1002/pssa.201700017. First published:


Golub M, Hejazi M, Kölsch A, Lokstein H, Wieland D.C.F, Zouni A, and Pieper J. Solution structure of monomeric and trimeric photosystem I of Thermosynechococcus elongatus investigated by small-angle X-ray scattering. Photosynthesis Research September 2017, Volume 133, Issue 1–3, pp 163–173. First Online: 03 March 2017


Fuller F.D, Gul S, Chatterjee R, Burgie E.S, Young I.D, Lebrette H, Srinivas V, Brewster A.S, Michels-Clark T, Clinger J.A, Andi B, Ibrahim M, Pastor E, de Lichtenberg C, Hussein R, Pollock C.J, Zhang M, Stan C.A, Kroll T,…,Zouni A, Johannes Messinger, Uwe Bergmann,…,Orville A.M, Kern J, Yachandra V.K & Yano J. Drop-on-demand sample delivery for studying biocatalysts in action at X-ray free-electron lasers. Nature Methods 14, 443–449 (2017) doi:10.1038/nmeth.4195 Published online: 27 February 2017



Young ID, Ibrahim M, Chatterjee R, Gul S, Koroidov S, Brewster AS, Tran R, Alonso-Mori R, Kroll T, Michels-Clark T, Laksmono H, Sierra RG, Stan CA, Hussein R, Zhang M, Douthit L, Kubin M, de Lichtenberg C, Pham LV, Nilsson H, Cheah MH, Shevela D, Saracini C, Bean MA, Seuffert I, Sokaras D, Weng TC. Pastor E,…, Zouni A, Messinger J, Bergmann U, Sauter NK, Kern J, Yanchandra VK, Junko Yano. Structure of photosystem II and substrate binding at room temperature. Nature 540, 453–457 (15 December 2016) doi:10.1038/nature20161


Stieger KR, Feifel SC, Lokstein H, Hejazi M, Zouni A, Lisdat F. Biohybrid architectures for efficient light-to-current conversion based on photosystem I within scalable 3D mesoporous electrodes. J. Mater. Chem. A, 2016, 4, 17009-17017 DOI: 10.1039/C6TA07141D!divAbstract


Alonso-Mori R, Asa K, Bergmann U, Brewster AS, Chatterjee R, Cooper JK, Frei HM, Fuller FD, Goggins E, Gul S, Fukuzawa H, Iablonskyi D, Ibrahim M, Katayama T, Kroll T, Kumagai Y, McClure BA, Messinger J, Motomura K, Nagaya K, Nishiyama T, Saracini C, Sato Y, Sauter NK, Sokaras D, Takanashi T, Togashi T, Ueda K, Weare WW, Weng TC, Yabashi M, Yachandra VK, Young ID, Zouni A, Kern JF, Yano J. Towards characterization of photo-excited electron transfer and catalysis in natural and artificial systems using XFELs. Faraday Discuss. 2016 Dec 16;194:621-638. doi: 10.1039/c6fd00084c.


Bommer M, Bondar AN, Zouni A, Dobbek H, Dau H. Crystallographic and Computational Analysis of the Barrel Part of the PsbO Protein of Photosystem II: Carboxylate–Water Clusters as Putative Proton Transfer Relays and Structural Switches. Biochemistry201655 (33), pp 4626–4635 DOI: 10.1021/acs.biochem.6b00441


Sauter NK, Echols N, Adams PD, Zwart PH, Kern J, Brewster AS, Koroidov S, Alonso-Mori R, Zouni A, Messinger J, Bergmann U, Yano J, Yachandra V, No observable conformational changes in PSII. Nature 533, E1–E2(19 May 2016) doi:10.1038/nature17983


Stieger KR, Ciornii DKölsch AHejazi MLokstein HFeifel SCZouni ALisdat F.Engineering of supramolecular photoactive protein architectures: the defined co-assembly of photosystem I and cytochrome c using a nanoscaled DNA-matrix. Nanoscale. 2016 May 19;8(20):10695-705. doi: 10.1039/c6nr00097e.!divAbstract


Roessler CG, Agarwal R, Allaire M, Alonso-Mori R, Andi B, Bachega JF, Bommer M, Brewster AS, Browne MC, Chatterjee R, Cho E, Cohen AE, Cowan M, Datwani S, Davidson VL, Defever J, Eaton B, Ellson R, Feng Y, Ghislain LP, Glownia JM, Han G, Hattne J, Hellmich J, Héroux A, Ibrahim M, Kern J, Kuczewski A, Lemke HT, Liu P, Majlof L, McClintock WM, Myers S, Nelsen S, Olechno J, Orville AM, Sauter NK, Soares AS, Soltis SM, Song H, Stearns RG, Tran R, Tsai Y, Uervirojnangkoorn M, Wilmot CM, Yachandra V, Yano J, Yukl ET, Zhu D, Zouni A. Acoustic Injectors for Drop-On-Demand Serial Femtosecond Crystallography. Structure. 2016 Apr 5;24(4):631-40. doi: 10.1016/j.str.2016.02.007. Epub 2016 Mar 17.


Baxter E.L, Aquila L, Alonso-Mori R, Barnes C.O, Bonagura C.A, Brehmer W, Brunger A.T, Calero G, Caradoc-Davies T.T, Chatterjee R, Degrado W.F, Fraser J.S, Ibrahim M, Kern J, Kobilka B.K, Kruse A.C, Larsson K.M, Lemke, H.T, Lyubimov A.Y, Manglik A, McPhillips S.E, Norgren E, Pang S.S, Soltis S.M, Song J, Thomaston J, Tsai Y, Weis W.I, Woldeyes R.A, Yachandra V, Yano J, Zouni A, and Cohen A.E, High-density grids for efficient data collection from multiple crystals, Acta Crystallographica Section D Structural Biology 01/2016; 72(1). DOI: 10.1107/S2059798315020847


Sierra R.G, Gati C, Laksmono H, Han Dao E, Gul S, Fuller F, Kern J, Chatterjee R, Ibrahim M, Brewster A.S, Young I.D, Michels-Clark T, Aquila A, Liang M, Hunter M.S, Koglin J.E, Boutet S, Junco E.A, Hayes B, Bogan M.J, Hampton C.Y, Puglisi E.V, Sauter N.K, Stan C.A, Zouni A, Yano J, Yachandra V.K, Soltis S.M, Puglisi J.D, and DeMirci H, Concentric-flow electrokinetic injector enables serial crystallography of ribosome and photosystem II, Nature Methods.11/2015; DOI: 10.1038/nmeth.3667



Ibrahim M, Chatterjee R, Hellmich J, Tran R, Bommer M, Yachandra V.K, Yano J, Kern J and Zouni A, Improvements in serial femtosecond crystallography of photosystem II by optimizing crystal uniformity using microseeding procedures, Structural Dynamics 07/2015; 2(4):041705. DOI: 10.1063/1.4919741


Müh F, DiFiore D, and Zouni A, Influence of poly(ethylene glycol) on the micelle formation of alkyl maltosides used in membrane protein crystallization, Physical Chemistry Chemical Physics 03/2015; 17(17). DOI: 10.1039/C5CP00431D


Skandary S, Hussels M, Konrad A, Renger T, Müh F, Bommer M, Zouni A, Meixner A.J, and Brecht M, Variation of Exciton-Vibrational Coupling in Photosystem II Core Complexes from Thermosynechococcus elongatus As Revealed by Single-Molecule Spectroscopy, The Journal of Physical Chemistry B 02/2015; 119(11). DOI:10.1021/jp510631x


Müh F., Zouni A. (2015) Cytochrome b559 in photosystem II. In: Cramer WA, Kallas T (eds) Cytochrome Complexes: Evolution, Structures, Energy Transduction, and Signaling. Springer, Dordrecht



Kern, J., Tran, R., Alonso-Mori, R., Koroidov, S., Echols, N., Hattne, J., Ibrahim M., Gul, S, Laksmono, H., Sierra, R. G., Gildea, R. J., Han, G., Hellmich, J., Lassalle-Kaiser, B., Chatterjee, R., Brewster, A. S., Stan, C. A., Glöckner, C., Lampe, A., DiFiore, D.,    Milathianaki, D., Fry, A. R., Seibert, M. M., Koglin, J. E., Gallo, E., Uhlig, J., Sokaras, D.,   Weng, T.-C., Zwart, P. H., Skinner, D. E., Bogan, M. J., Messerschmidt, M., Glatzel, P., Williams, G. J., Boutet, S., Adams, P. D., Zouni, A., Messinger, J., Sauter, N. K., Bergmann, U., Yano, J., and Yachandra, V. K., Taking snapshots of photosynthetic water oxidation using femtosecond X-ray diffraction and spectroscopy Nat. Commun. 2014, 5, DOI: 10.1038/ncomms5371


Hellmich J, Bommer M, Burkhardt A, Ibrahim M, Kern J, Meents A, Müh F, Dobbek H, Zouni A, Native-like Photosystem II Superstructure at 2.44 Å Resolution through Detergent Extraction from the Protein Crystal, Structure 2014, 22, 1607-1615


Tran R., Kern J., Hattne J., Kroidov S., Hellmich J., Alonso-Mori R., Sauter N.K., Bergmann U., Messinger J., Zouni A., Yano J., Yachandra V.K., The Mn4Ca photosynthetic water-oxidation catalyst studied by simultaneous X-ray spectroscopy and crystallography using an X-ray free-electron laser, Phil. Trans. R. Soc. B (2014): 2014369 20130324, DOI: 10.1098/rstb.2013.0324


Brecht M, Skandary S., Hellmich J., Glöckner C., Konrad A., Hussels M., Meixner A. J., Zouni A., Schlodder E., Spectroscopic properties of photosystem II core complexes from Thermosynechococcus elongatus revealed by single-molecule experiments, Biochim Biophys Acta 2014, 1837, 773-781


Hattne J, Ecols N., Tran R., Kern J., Gildea R. J., Zouni A., Yano J., Bergmann U., Yachandra V. K., Adams P. D., Sater N. K.,  Accurate macromolecular structures using minimal measurements from X-ray free-electron lasers, Nature Methods 2014, 11, No 5, 545-548.



Glöckner C, Kern J, Broser M, Zouni A, Yachandra V, Junko Y, Structural Changes of the Oygen-evolving Complex in Photosystem II during the Catalytic Cycle, J. Biol. Chem. 2013, 288, 22607-22620


Kern J, Alonso-Mori R, Tran R, Hattne J, Gildea R.G., Echols N, Glöckner C, Hellmich J, Laksmono H, Sierra R G, Lassalle-Kaiser B, Koroidov S, Lampe A, Han G, Gul S,  DiFiore D, Milathianaki D, Fry A R, Miahnari A, Schafer D W, Messerschmidt M, Seibert M M, Jason Koglin E, Sokaras D, Weng T C, Sellberg J, Latimer M J, Grosse-Kunstleve R W, Zwart P H, White W H, Glatzel P, Adams P D, Bogan M J, Williams G j, Boutet S, Messinger J, Zouni A, Sauter N K, Yachandra V K, Bergmann U, Yano J,  Simultaneous Femtosecond X-Ray Spectroscopy and Diffraction of Photosystem II at Room Temperature. Science 2013, 340, 491-495


Müh F., Zouni A., The nonheme iron in photosystem II, Photosynth Res (2013) 116: 295-414, DOI 10.1007/s11120-013-9926-y.


Brose K., Zouni A., Müh F., Mroginski M.A., Maultzsch J., Simulations of the polarization-dependent Raman intensity of β-carotene in photosystem II crystals, Chemical Physics 2013, 418, 65-73



J. Kern, R. Alonso-Mori, J. Hellmich, R. Tran, J. Hattne, H. Laksmono, C. Glöckner, N. Echols, R.G. Sierra, J. Sellberg, B. Lassalle-Kaiser, R. J. Gildea, P. Glatzel, R. W. Grosse-Kunstleve, M. J. Latimer, T. A. McQueen, D. DiFiore, A. R. Fry, M. Messerschmidt, A. Miahnahri, D. W. Schafer, M. M., Seibert, D. Sokaras, T. C. Weng, P. H. Zwart, W. E. White, P. D. Adams, M. J. Bogan, S. Boutet, G. J. Williams, J. Messinger, N. K. Sauter, A. Zouni, U. Bergmann, J. Yano, V. K. Yachandra, Room Temperature Femtosecond X-ray Diffraction of Photosystem II Microcrystals, Proc. Natl. Acad. Sci. U. S. A. 2012, 109, 9721-9726


Burkhardt A., Warmer M., Pannneerselvam S., Wagner, A., Zouni A., Glöckner C., Reimer R., Hohenberg H, Meents A., Fast high-pressure freezing of protein crystals in their mother liquor, Acta Cryst 2012, F68, 495-500


Müh F., Glöckner C., Hellmich J., Zouni A., Light-induced quinone reduction in photosystem II, Biochim Biophys Acta 2012, 1817, 44-65 


Alonso-Mori R., Kern J., Gildea R. J., Sokaras D., Wenig T.-C., Lassalle-Kaiser B., Tran R., Hattne J., Laksmono H., Hellmich J., Glöckner C., Echols N., Sierra R. G., Schafer D. W., Sellberg J., Kenney C., Herbst R., Pines J., Hart P., Herrmann S., Grosse-Kunstleve R. W., Latimer M. J., Fry A. R., Messerschmidt M. M., Miahnahri A., Seibert M. M., Zwart P. H., White W. E., Adams P. D., Bogan M. J., Boutet S., Williams G. J., Zouni A., Messinger J., Glatzel P., Sauter N. K., Yachandra V. K., Yano J., Bergmann U, Energy-dispersive X-ray emission spectroscopy using an X-ray free-electron laser in a shot-by-shot mode, Proc. Natl. Acad. Sci. U. S. A. 2012, 109, 19103-19107


Sierra R. G.,Laksmono H, Kern,J., Tran R., Hattne J.,  Alonso-Mori,R., Lassalle-Kaiser B.,  Glöckner C., Hellmich J., Schafer D. W., Echols N., Gildea R. J., ,b Grosse-Kunstleve R. W., Sellberg J., McQueen T. A., Fry A. R., Messerschmidt M. M.,Miahnahri A., Seibert M. M., Hampton C. Y., Starodub D., Loh N. D., Sokaras D., Wenig T. C., Zwart P.H., Glatzel P, Milathianaki D., White W. E., Adams P. D., Williams G. J., Boutet S., Zouni A., Messinger J.,  Sauter N. K., Bergmann U., Yano J., Yachandra V. K.,  Bogan M., (2012),  Nanoflow electrospinning serial femtosecond crystallography, Acta Cryst., D68, 1584-1587



Broser M, Glöckner C, Gabdulkhakov A, Guskov A, Buchta J, Kern J, Müh F, Dau H, Saenger W, Zouni A, Structural Basis of Cyanobacterial Photosystem II Inhibition by the Herbicide Terbutryn, J. Biol. Chem. 2011, 286, 15964-15972 


Müh, F., Zouni, A. (2011) Light-induced water oxidation in photosystem II, Frontiers in Bioscience-Landmark, 16, 3072-3132.



Broser, M., Gabdulkhakov, A., Kern, J., Guskov, A., Müh, F., Saenger, W.,  Zouni, A. (2010) Crystal Structure of Monomeric Photosystem II from Thermosynechococcus elongatus at 3.6 Å Resolution. J. Biol. Chem. 285, 26255-26262.


Guskov, A., Gabdulkhakov, A., Broser, M.,  Glöckner, C., Hellmich, J., Kern, J.,  Frank, J.,  Müh, F., Saenger, W., Zouni, A. (2010) Recent Progress in the Crystallographic Studies of Photosystem II . Chem Phys Chem , 11, 1160-1171.


Keßen, S., Teutloff, C.,  Kern, J.,  Zouni, A.,  Bittl, R. (2010) High-Field 2H-Mims-ENDOR spectroscopy on PSII single crystals: The Hydrogen Bonding of Y.D Chem Phys Chem, 11, 1275-1282


Brose K, Tschirner N., Zouni A., Müh F., Yanagi K., Kataura H., Thomson C., Maultzsch J. (2010) Polarised Raman measurements of β-carotene encapsulated in SWNTs. Phys. Status Solid B, 247, Nos. 11-12, 2871-2874.



Kern, J., Zouni, A.,  Guskov, A., Krauss, N. (2009) Lipids in the Structure of Photosystem I, Photosystem II and the Cytochrome b6f Complex in H. Wada, N. Murata and Govindjee (eds.), Lipids in Photosynthesis: Essential and Regulatory Functions, PP. 203-241 Springer Science + Business Media B. V. 2009


Gabdulkhakov, A.,  Guskov, A. , Broser, M., Kern, J., Müh, F., Saenger, W., Zouni, A. (2009). Probing the Accessibilty of the Mn4Ca Cluster in Photosystem II: Channels Calculation, Noble Gas Derivatization, and Cocrystallization with DMSO. Structure, 17, 1223-1234.


Teutloff, C., Pudollek, S., Kessen, S.,  Broser, M. , Zouni, A., Bittl, R. (2009). Electronic structure of the tyrosine D radical and the water-splitting complex from pulsed ENDOR spectroscopy on photosystem II single crystals.  Phys Chem Chem Phys, 11, 6715-6726.


Balaban, T.S., Braun ,P., Hättig, C., Hellweg, A., Kern, J., Saenger, W., Zouni, A. (2009) Preferential pathways for light-trapping involving b-ligated chlorophylls. Biochim Biophys Acta, 1787, 1254-1265


Guskov,A., Kern,J., Gabdulkhakov,A, Broser,M., Zouni,A., Saenger,W. (2009). Cyanobacterial photosystem II at 2.9-Å resolution and the role of quinones, lipids, channels and chloride. Nat Struct Mol Biol, 16 (3), 334-342.



Loll, B., Broser, M., Kos, P. B., Kern, J., Biesiadka, J.,  Vass, I., Saenger, W., Zouni, A. (2008). Modeling of variant copies of subunit D1 in the structure of photosystem II from Thermosynechococcus elongatus. Biol. Chem., 389, 609-617.


Müh, F., Renger, T, Zouni, A. (2008). Crystal structure of cyanobacterial photosystem II at 3.0 Å resolution: A closer look at the antenna system and the small membrane-intrinsic subunits, Plant Physiol. Biochem. 46, 238-264.


Müh, F., Zouni, A. (2008). Micelle formation in the presence of photosystem I, Biochim. Biophys. Acta 1778, 2298-2307.


Zouni, A. (2008). From cell growth to the 3.0 Å resolution crystal structure of cyanobacterial photosystem II, in G. Renger (ed.) Primary processes of photosynthesis: Basic principles and apparatus, Vol. II, Royal Society Chemistry, Cambridge, pp 193-236.


Yano, J., Kern, J., Pushkar, Y., Sauer, K., Glatzel, P., Bergmann, U., Messinger, J., Zouni, A., Yachandra, V. (2008). High-resolution structure of the photosynthetic Mn4Ca catalyst from X-ray spectroscopy. Philos. Trans. R. Soc. Lond. B Biol. Sci. 363, 1139-1147.


Zein, S., Kulik, L.V., Yano, J., Kern, J., Pushkar, Y., Zouni, A., Yachandra, V.K., Lubitz, W., Neese, F., Messinger, J. (2008). Focusing the view on nature’s water-splitting catalyst. Phil. Trans. R. Soc. Lond. B Biol Sci., 363, 1167-1177.



Ishikita, H., Loll, B., Biesiadka, J., Kern, J., Irrgang, K.-D., Zouni, A., Saenger, W., Knapp, E.-W. (2007). Function of two beta-carotenes near the D1 and D2 proteins in photosystem II dimers. Biochim. Biophys. Acta 1767, 79-87.


Kern, J., Biesiadka, J., Loll, B., Saenger, W., Zouni, A. (2007). Structure of the Mn4Ca cluster as derived from X-ray diffraction. Photosynth. Res. 92, 389-405


Krivanek, R., Kern, J., Zouni, A., Dau, H., Haumann, M. (2007). Spare quinones in the QB cavity of crystallized photosystem II from Thermosynechococcus elongatus. Biochim. Biophys. Acta 1767, 520-527.

Loll, B., Kern, J., W. Saenger, W., Zouni, A., Biesiadka, J. (2007). Lipids in Photosystem II: Interactions with protein and cofactors. Biochim. Biophys. Acta 1767,  509-519.



Teutloff, C., Keßen, S., Kern, J., Zouni, A., Bittl, R. (2006). High-field (94-GHz) EPR spectroscopy on the S2 multiline signal of photosystem II. FEBS Lett. 580, 3605-3609.

Yano, J, Kern, J., Sauer, K., Latimer, M. J., Pushkar, Y., Biesiadka, J., Loll, B., Saenger, W., Messinger, J., Zouni, A., Yahandra, V. K. (2006). Where water is oxidized to dioxygen: Structure of the photosynthetic Mn4Ca cluster. Science 314, 821-825.


Zimmermann, K, Heck, M., Frank, J., Kern, J., Vass, I., Zouni, A. (2006). Herbicide binding and thermal stability of photosystem II isolated from Thermosynechococcus elongatus. Biochim. Biophys. Acta 1757, 106-114.



Kern, J., Loll, B., Lüneberg, C., DiFiore, D., Biesiadka, J., Irrgang, K-D., Zouni, A. (2005). Purification, characterisation and crystallisation of photosystem II from Thermosynechococcus elongatus cultivated in a new type of photobioreactor. Biochim. Biophys. Acta 1706, 147-157


Kern, J., Loll, B., Zouni, A., Saenger, W., Irrgang, K.-D., Biesiadka, J. (2005). Cyanobacterial photosystem II at 3.2 Å resolution - the plastoquinone binding pockets. Photosynth. Res. 84, 153-159.


Loll, B., Kern, J., Saenger, W. Zouni, A., Biesiadka, J. (2005). Towards complete cofactor arrangement in the 3.0 Å resolution structure of photosystem II. Nature 438, 1040-1044.


Loll, B., Kern, J., Zouni, A., Saenger, W., Biesiadka, J., Irrgang, K.-D. (2005). The antenna system of Photosystem II from Thermosynechococcus elongatus at 3.2 Å resolution. Photosynth. Res. 86, 175-184.


Müh, F. Zouni, A. (2005). Extinction coefficients and critical solubilisation concentrations of photosystems I and II from Thermosynechococcus elongatus. Biochim. Biophys. Acta 1708, 219-228.


Yano, J., Kern, J., Irrgang, K.-D., Latimer, M., Bergmann, U., Glatzel, P., Pushkar, Y., Biesiadka, J., Loll, B., Sauer K., Messinger, J., Zouni, A., Yachandra, V. K. (2005). X-ray damage to the Mn4Ca complex in single crystals of photosystem II: A case study for metalloprotein crystallography. Proc. Natl. Acad Sci. U. S. A. 102, 12047-12052.


Zouni, A., Kern, J., Frank, J., Hellweg, T., Behlke, J., Saenger, W., Irrgang, K.-D. (2005). Size determination of cyanobacterial and higher plant photosystem II by gel permeation chromatography, light scattering and ultracentrifugation. Biochemistry 44, 4572-4581.



Biesiadka, J., Loll, B., Kern, J., Irrgang, K-D., Zouni, A. (2004). Crystal structure of cyanobacterial photosystem II at 3.2 Å resolution: a closer look at the Mn-cluster. Phys. Chem. Chem. Phys. 6, 4733-4736.


Kern, J., Biesiadka, J., Loll, B., Zouni, A., Irrgang, K.-D., Saenger, W. (2004). Crystal structure of cyanobacterial photosystem II at 3.2 Å resolution. In Photosynthesis: Fundamental Aspects to Global Perspectives, A. van der Est and D. Bruce, eds (Allen Press, Lawrence (KS), USA), pp. 230-234.


Yano, J., Kern, J., Sauer, K., Messinger, J., Zouni, A., Yachandra, V. K. (2004). Mn4Ca Cluster of the water-oxidation enzyme studied by polarized X-ray spectroscopy of PS II single crystals. In Photosynthesis: Fundamental Aspects to Global Perspectives, A. van der Est and D. Bruce, eds (Allen Press, Lawrence (KS), USA), pp. 234-236.


Zouni, A., Kern, J., Loll, B., Biesiadka, J., Saenger, W., Irrgang, K.-D. (2004). Biochemical and functional analysis of Photosystem II crystals from Thermosynechococcus elongatus diffracting to 3.2 Å. In Photosynthesis: Fundamental Aspects to Global Perspectives, A. van der Est and D. Bruce, eds (Allen Press, Lawrence (KS), USA), pp. 259-261.



Sarrou, J., Isgandarova, S., Kern, J., Zouni, A., Renger, G., Lubitz, W., and Messinger, J. (2003). Nitric oxide-induced formation of the S-(2) state in the oxygen-evolving complex of photosystem II from Synechococcus elongatus. Biochemistry 42, 1016-1023.



Fromme, P., Kern, J., Loll, B., Biesiadka, J., Saenger, W., Witt, H.T., Krauss, N., and Zouni, A. (2002). Functional implications on the mechanism of the function of photosystem II including water oxidation based on the structure of photosystem II. Philos. Trans. R. Soc. Lond. B Biol. Sci. 357, 1337-1344



Hofbauer, W., Zouni, A., Bittl, R., Kern, J., Orth, P., Lendzian, F., Fromme, P., Witt, H.-T., and Lubitz, W. (2001). Photosystem II single crystals studied by EPR spectroscopy at 94 GHz: the tyrosine radical Y(D)(*). Proc. Natl. Acad. Sci. U. S. A. 98, 6623-6628.


Irrgang, K.-D., Georgalis, Y., Franke, P., Behlke, J., Saenger, W., and Zouni, A. (2001). Structural comparison of oxygen-evolving photosystem II core complexes from Synechococcus elongatus and Spinacia oleracea. In 12th International Conference on Photosynthesis (Brisbane, Australia: CSIRO Publishers, Melbourne, Australia), pp. S5-028.


Vasil'ev, S., Orth, P., Zouni, A., Owens, T.G., and Bruce, D. (2001). Excited-state dynamics in photosystem II: insights from the x-ray crystal structure. Proc. Natl. Acad. Sci. U. S. A. 98, 8602-8607.


Witt, H.T., Zouni, A., Kern, J., Fromme, P., Krauss, N., Saenger, W., and Orth, P. (2001). Crystal structure of Photosystem II and aspects of its function. In 12th International congress on photosynthesis (Brisbane: CSIRO Publishers, Melbourne, Australia), pp. PL01.


Zouni, A., Witt, H.T., Kern, J., Fromme, P., Krauss, N., Saenger, W., and Orth, P. (2001). Crystal structure of photosystem II from Synechococcus elongatus at 3.8 A resolution. Nature 409, 739-743.


Zouni, A., Kern, J., Loll, B., Fromme, P., Witt, H.T., Orth, P., Krauss, N., Saenger, W., and Biesiadka, J. (2001). Biochemical characterization and crystal structure of water oxidizing photosystem II from Synechococcus elongatus. In 12th International congress on photosynthesis (Brisbane: CSIRO Publishers, Melbourne, Australia), pp. S05-003.

vor 2001

Zouni, A., Lüneberg, C., Fromme, P., Schubert, W.D., Saenger, W., and Witt, H.T. (1998). Characterization of single crystals of Photosystem II from the thermophilic cyanobacterium Synechococcus elongatus. In Photosynthesis: Mechanisms and Effects, G. Garab, ed (Dordrecht: Kluwer Academic), pp. 925-928.


Zouni, A., Jordan, R., Schlodder, E., Fromme, P., and Witt, H.T. (2000). First photosystem II crystals capable of water oxidation. Biochim. Biophys. Acta 1457, 103-105.