List of Publications


  • Hemschemier A, Posewitz MC, Happe T (2023) The Chlamydomonas Sourcebook (Third Edition), Volume 2 : Organellar and Metabolic Processes, Chapter 10 - Hydrogenases and hydrogen production. Pages 343-367
  • Posewitz MC, Atteia A, Hemschemier A, Happe T, Grossman AR (2023) The Chlamydomonas Sourcebook (Third Edition), Volume 2 : Organellar and Metabolic Processes, Chapter 9 - Metabolic networks during dark anoxia. Pages 317-341
  • Alavi G, Engelbrecht V, Hemschemeier A, Happe T (2023) The Alga Uronema belkae Has Two Structural Types of [FeFe]-Hydrogenases with Different Biochemical Properties. Int. J. Mol. Sci.
  • Duan J, Veliju A, Lampret O, Liu L, Yadav S, Apfel UP, Armstrong FA, Hemschemeier A, Hofmann E (2023) Insights into the Molecular Mechanism of Formaldehyde Inhibition of [FeFe]-Hydrogenases. J. Am. Chem. Soc.
  • Brocks C, Das CK, Duan J, Yadav S, Apfel UP, Ghosh S, Hofmann E, Winkler M, Engelbrecht V, Schäfer LV, Happe T (2023) A Dynamic Water Channel Affects O2 Stability in [FeFe] Hydrogenases. ChemSusChem
  • Haas R, Engelbrecht V, Lampret O, Yadav S, Apfel UP, Leimkühler S, Happe T (2023) The [4Fe-4S]-Cluster of HydF is not Required for the Binding and Transfer of the diiron site of [FeFe]-Hydrogenases. ChemBioChem
  • Böhmer S, Marx C, Goss R, Gilbert M, Sasso S, Happe T, Hemschemeier A (2023) Chlamydomonas reinhardtii mutants deficient for Old Yellow Enzyme 3 exhibit increased photooxidative stress. Plant Direct
  • Rutz A, Das CK, Fasano A, Jaenecke J, Yadav S, Apfel UP, Engelbrecht V, Fourmond V, Léger C, Schäfer LV, Happe T (2023) Increasing the O2 resistance of the [FeFe]-hydrogenase CbA5H through enhanced protein flexibility. ACS Catalysis, DOI: 10.1021/acscatal.2c04031


  • Duan J, Hemschemeier A, Burr DJ, Stripp ST, Hofmann E, Happe T (2022) Cyanide Binding to [FeFe]-Hydrogenase Stabilizes the Alternative Configuration of the Proton Transfer Pathway. Angew Chem Int Ed Engl.: e202216903 doi: 10.1002/anie.202216903
  • Günzel A, Engelbrecht V, Happe T (2022) Changing the tracks: screening for electron transfer proteins to support hydrogen production. J Biol Inorg Chem (7):631-640 doi: 10.1007/s00775-022-01956-1
  • Heghmanns M, Rutz A, Kutin Y, Engelbrecht E, Winkler M, Happe T, Kasan­mascheff M (2022) The Oxygen-Resistant [FeFe]-Hydrogenase CbA5H Harbors an Unknown Radical Signal. Chemical Science 13, 24: 7289–94
  • Published Erratum Chem Sci (2022) Jul 14;13(29):8704. doi: 10.1039/d2sc90140d. eCollection 2022 Jul 29.


  • Heghmanns M, Günzel A, Brandis D, Kutin Y, Engelbrecht V, Winkler M, Happe T, Kasan­mascheff M (2021) Fine-Tuning of FeS Proteins Monitored via Pulsed EPR Redox Potentiometry at Q-Band. Biophysical Reports 1, 2
  • Laun K, Baranova I, Duan J, Kertess L, Wittkamp F, Apfel UP, Happe T, Senger M, Stripp ST (2021) Site-selective protonation of the one-electron reduced cofactor in [FeFe]-hydrogenase. Dalton Trans 50, 10  doi: 10.26434/chemrxiv.9970571.v3
  • Morra S, Duan J, Winkler M, Ash PA, Happe T, Vincent KA (2021) Electrochemical control of [FeFe]-hydrogenase single crystals reveals complex redox populations at the catalytic site. Dalton Trans  doi: 10.1039/D1DT02219
  • Winkler M, Duan J, Rutz A, Felbek C, Scholtysek L, Lampret O, Jaenecke J, Apfel UP, Gilardi G, Valetti F, Fourmond V, Hofmann E, Léger C, Happe T (2021) A safety cap protects hydrogenase from oxygen attack. Nat Commun 12, 756  doi: 10.1038/s41467-020-20861-2


  • Engelbrecht V, Liedtke K, Rutz A, Yadav S, Günzel A, Happe T (2020): One isoform for one task? The second hydrogenase of Chlamydomonas reinhardtii prefers hydrogen uptake. Int J Hydrogen Energy  doi: 10.1016/j.ijhydene.2020.11.231
  • Lampret O, Duan J, Hofmann E, Winkler M, Armstrong FA, Happe T (2020): The roles of long-range proton-coupled electron transfer in the directionality and efficiency of [FeFe]-hydrogenases. Proc Natl Acad Sci USA 117(34):20520-20529.  doi: 10.1073/pnas.2007090117
  • Böhmer S, Marx C, Gómez-Baraibar A, Nowaczyk MM, Tischler D, Hemschemeier A, Happe T (2020): Evolutionary diverse Chlamydomonas reinhardtii Old Yellow Enzymes reveal distinctive catalytic properties and potential for whole-cell biotransformations. Algal Research 50: 101970.  doi: 10.1016/j.algal.2020.10197
  • Lorent C, Katz S, Duan J, Kulka CJ, Caserta G, Teutloff C, Yadav S, Apfel UP, Winkler M, Happe T, Horch M, Zebger I (2020): Shedding Light on Proton and Electron Dynamics in [FeFe] Hydrogenases. J Am Chem Soc. 142(12):5493-5497.  doi:10.1021/jacs.9b13075
  • Luo GF, Biniuri Y, Chen WH, Wang J, Neumann E, Marjault HB, Nechushtai R, Winkler M, Happe T, Willner I (2020): Modelling Photosynthesis with ZnII -Protoporphyrin All-DNA G-Quadruplex/Aptamer Scaffolds. Angew Chem Int Ed Engl. 59(23):9163-9170.  doi:10.1002/anie.202002915
  • Adams EM, Lampret O, König B, Happe T, Havenith M (2020): Solvent dynamics play a decisive role in the complex formation of biologically relevant redox proteins. Phys Chem Chem Phys. 22(14):7451-7459.  doi:10.1039/d0cp00267d


  • Sidabras JW, Duan J, Winkler M, Happe T, Hussein R, Zouni A, Suter D, Schnegg A, Lubitz W, Reijerse EJ (2019): Extending electron paramagnetic resonance to nanoliter volume protein single crystals using a self-resonant microhelix. Sci Adv 5(10):eaay1394.  doi: 10.1126/sciadv.aay1394
  • Esselborn J, Kertess L, Apfel UP, Hofmann E, Happe T (2019): Loss of Specific Active-Site Iron Atoms in Oxygen-Exposed [FeFe]-Hydrogenase Determined by Detailed X-ray Structure Analyses. J Am Chem Soc [Epub ahead of print].  doi: 10.1021/jacs.9b07808
  • Senger M, Eichmann V, Laun K, Duan J, Wittkamp F, Knör G, Apfel UP, Happe T, Winkler M, Heberle J, Stripp ST (2019): How [FeFe]-Hydrogenase Facilitates Bidirectional Proton Transfer. J Am Chem Soc 141(43):17394-17403.  doi: 10.1021/jacs.9b09225
  • Duan J, Mebs S, Senger M, Laun K, Wittkamp F, Heberle J, Happe T, Hofmann E, Apfel UP, Winkler M, Haumann M, Stripp ST (2019): The Geometry of the Catalytic Active Site in [FeFe]-hydrogenases is Determined by Hydrogen Bonding and Proton Transfer. ACS Catalysis [Epub ahead of print].  doi: 10.1021/acscatal.9b02203
  • Luo GF, Biniuri Y, Chen WH, Neumann E, Fadeev M, Marjault HB, Bedi A, Gidron O, Nechushtai R, Stone D, Happe T, Willner I (2019): Artificial Photosynthesis with Electron Acceptor/Photosensitizer-Aptamer Conjugates. Nano Lett [Epub ahead of print].  doi: 10.1021/acs.nanolett.9b02880
  • Lampret O, Esselborn J, Haas R, Rutz A, Booth RL, Kertess L, Wittkamp F, Megarity CF, Armstrong FA, Winkler M, Happe T (2019): The final steps of [FeFe]-hydrogenase maturation. Proc Natl Acad Sci U S A 23.pii:201908121.  doi: 10.1073/pnas.1908121116
  • Mebs S, Duan J, Wittkamp F, Stripp ST, Happe T, Apfel UP, Winkler M, Haumann M (2019): Differential Protonation at the Catalytic Six-Iron Cofactor of [FeFe]-Hydrogenases Revealed by 57Fe Nuclear Resonance X-ray Scattering and Quantum Mechanics/Molecular Mechanics Analyses. Inorg Chem 18;58(6):4000-4013.  doi: 10.1021/acs.inorgchem.9b00100
  • Rodríguez-Maciá P, Kertess L, Burnik J, Birrell JA, Hofmann E, Lubitz W, Happe T, Rüdiger O (2019): His-ligation to the [4Fe-4S] sub-cluster tunes the catalytic bias of [FeFe] hydrogenase. J Am Chem Soc 9;141(1):472-481.  doi: 10.1021/jacs.8b11149


  • Engelbrecht V, Happe T (2018): [FeFe]-hydrogenases from green algae. Methods Enzymol 613:203-230.  doi: 10.1016/bs.mie.2018.10.004
  • Duan J, Senger M, Esselborn J, Engelbrecht V, Wittkamp F, Apfel UP, Hofmann E, Stripp ST, Happe T, Winkler M (2018): Crystallographic and spectroscopic assignment of the proton transfer pathway in [FeFe]-hydrogenases. Nat Commun 9;9(1):4726.  doi: 10.1038/s41467-018-07140-x
  • Li H, Buesen D, Williams R, Henig J, Stapf S, Mukherjee K, Freier E, Lubitz W, Winkler M, Happe T, Plumeré N (2018): Preventing the coffee-ring effect and aggregate sedimentation by in situ gelation of monodisperse materials. Chem Sci 9:7596-7605.  doi: 10.1039/C8SC03302A
  • Hemschemeier A, Happe T (2018): The plasticity of redox cofactors: from metalloenzymes to redox-active DNA. Nat Rev Chem 2:231–243.  doi: 10.1038/s41570-018-0029-3
  • Bai Y, Chen T, Happe T, Lu Y, Sawyer A (2018): Iron-sulphur cluster biogenesis via the SUF pathway. Metallomics 15;10(8):1038-1052.  doi: 10.1039/c8mt00150b
  • Laun K, Mebs S, Duan J, Wittkamp F, Apfel UP, Happe T, Winkler M, Haumann M, Stripp ST (2018): Spectroscopical Investigations on the Redox Chemistry of [FeFe]-Hydrogenases in the Presence of Carbon Monoxide. Molecules 9;23(7). pii: E1669.  doi: 10.3390/molecules23071669
  • Burlacot A, Sawyer A, Cuiné S, Auroy-Tarrago P, Blangy S, Happe T, Peltier G (2018): Flavodiiron-Mediated O2 Photoreduction Links H2 Production with CO2 Fixation during the Anaerobic Induction of Photosynthesis. Plant Physiol 177(4):1639-1649.  doi: 10.1104/pp.18.00721
  • Bothe H, Happe T, Trebst S, Rögner M (2018): In memory of Achim Trebst (1929-2017): a pioneer of photosynthesis research. Photosynth Res. 137(3):341-359.  doi: 10.1007/s11120-018-0516-x
  • Wiegand K, Winkler M, Rumpel S, Kannchen D, Rexroth S, Hase T, Farès C, Happe T, Lubitz W, Rögner M (2018): Rational redesign of the ferredoxin-NADP+-oxido-reductase/ferredoxin-interaction for photosynthesis-dependent H2-production. Biochim Biophys Acta 1859(4):253-262.  doi: 10.1016/j.bbabio.2018.01.006
  • Düner M, Lambertz J, Mügge C, Hemschemeier A (2018): The soluble guanylate cyclase CYG12 is required for the acclimation to hypoxia and trophic regimes in Chlamydomonas reinhardtii. Plant J 93(2):311-337.  doi: 10.1111/tpj.13779
  • Mebs S, Kositzki R, Duan J, Kertess L, Senger M, Wittkamp F, Apfel UP, Happe T, Stripp ST, Winkler M, Haumann M (2018): Hydrogen and oxygen trapping at the H-cluster of [FeFe]-hydrogenase revealed by site-selective spectroscopy and QM/MM calculations. Biochim Biophys Acta 1859:28-41.  doi: 10.1016/j.bbabio.2017.09.003


  • Sawyer A, Esselborn J, Winkler W, Happe T (2017) Chlamydomonas: Hydrogenase and Hydrogen Production. In Hippler M, ed, Chlamydomonas: Molecular Genetics and Physiology. Microbiology Monographs, vol 31. Springer, Cham, pp 21-44.  10.1007/978-3-319-66360-9_2
  • Hemschemeier A (2017) Chlamydomonas: Anoxic Acclimation and Signaling. In Hippler M, ed, Chlamydomonas: Molecular Genetics and Physiology. Microbiology Monographs, vol 31. Springer, Cham, pp 155-199.  10.1007/978-3-319-66365-4_6
  • Lampret O, Adamska-Venkatesh A, Konegger H, Wittkamp F, Apfel UP, Reijerse EJ, Lubitz W, Rüdiger O, Happe T, Winkler M (2017): Interplay between CN– Ligands and the Secondary Coordination Sphere of the H-Cluster in [FeFe]-Hydrogenases. J Am Chem Soc 139(50):18222-18230.  doi: 10.1021/jacs.7b08735
  • Senger M, Laun K, Wittkamp F, Duan J, Haumann M, Happe T, Winkler M, Apfel UP, Stripp ST (2017): Proton-Coupled Reduction of the Catalytic [4Fe-4S] Cluster in [FeFe]-Hydrogenases. Angew Chem Int Ed Engl 56(52):16503-16506.  doi: 10.1002/anie.201709910
  • Kertess L, Adamska-Venkatesh A, Rodríguez-Maciá P, Rüdiger O, Lubitz W, Happe T (2017): Influence of the [4Fe-4S] cluster coordinating cysteines on active site maturation and catalytic properties of C. reinhardtii [FeFe]-hydrogenase. Chem Sci 8:8127-8137.  doi: 10.1039/C7SC03444J
  • Senger M, Mebs S, Duan J, Shulenina O, Laun K, Kertess L, Wittkamp F, Apfel UP, Happe T, Winkler M, Haumann M, Stripp ST (2017): Protonation/reduction dynamics at the [4Fe-4S] cluster of the hydrogen-forming cofactor in [FeFe]-hydrogenases. Phys Chem Chem Phys 20(5):3128-3140.  doi: 10.1039/c7cp04757f
  • Schuth N, Mebs S, Huwald D, Wrzolek P, Schwalbe M, Hemschemeier A, Haumann M (2017): Effective intermediate-spin iron in O2-transporting heme proteins. Proc Natl Acad Sci USA 1859:28-41.  10.1073/pnas.1706527114
  • Mebs S, Senger M, Duan J, Wittkamp F, Apfel UP, Happe T, Winkler M, Stripp ST, Haumann M (2017): Bridging Hydride at Reduced H-Cluster Species in [FeFe]-Hydrogenases Revealed by Infrared Spectroscopy, Isotope Editing, and Quantum Chemistry. J Am Chem Soc 139:12157-12160.  doi: 10.1021/jacs.7b07548
  • Winkler M, Senger M, Duan J, Esselborn J, Wittkamp F, Hofmann E, Apfel UP, Stripp ST, Happe T (2017): Accumulating the hydride state in the catalytic cycle of [FeFe]-hydrogenases. Nat Commun 8:16115.  doi: 10.1038/NCOMMS16115
  • Engelbrecht V, Rodríguez-Maciá P, Esselborn J, Sawyer A, Hemschemeier A, Rüdiger O, Lubitz W, Winkler M, Happe T (2017): The structurally unique photosynthetic Chlorella variabilis NC64A hydrogenase does not interact with plant-type ferredoxins. Biochim Biophys Acta 1858:771-778.  doi: 10.1016/j.bbabio.2017.06.004
  • Sawyer A, Winkler M (2017): Evolution of Chlamydomonas reinhardtii ferredoxins and their interactions with [FeFe]-hydrogenases. Photosynth Res 134(3):307-316.  doi: 10.1007/s11120-017-0409-4
  • Mosebach L, Heilmann C, Mutoh R, Gäbelein P, Steinbeck J, Happe T, Ikegami T, Hanke G, Kurisu G, Hippler M (2017): Association of Ferredoxin:NADP+ oxidoreductase with the photosynthetic apparatus modulates electron transfer in Chlamydomonas reinhardtii. Photosynth Res 134(3):291-306.  doi: 10.1007/s11120-017-0408-5
  • Pandey K, Islam ST, Happe T, Armstrong FA (2017): Frequency and potential dependence of reversible electrocatalytic hydrogen interconversion by [FeFe]-hydrogenases. Proc Natl Acad Sci USA 114(15):3843-3848.  doi: 10.1073/pnas.1619961114
  • Sawyer A, Bai Y, Lu Y, Hemschemeier A, Happe T (2017): Compartmentalisation of [FeFe]-hydrogenase maturation in Chlamydomonas reinhardtii. Plant J 90:1134-1143.  doi: 10.1111/tpj.13535


  • Adam D, Bösche L, Castaneda-Losada L, Winkler M, Apfel UP, Happe T (2016): Sunlight dependent hydrogen production by photosensitizer/hydrogenase systems. ChemSusChem 10(5):894-902.  doi: 10.1002/cssc.201601523
  • Katz S, Noth J, Horch M, Shafaat HS, Happe T, Hildebrandt P, Zebger I (2016): Vibrational spectroscopy reveals the initial steps of biological hydrogen evolution. Chem Sci 7:6746-6752.  doi: 10.1039/c6sc01098a
  • Megarity CF, Esselborn J, Hexter SV, Wittkamp F, Apfel UP, Happe T, Armstrong FA (2016): Electrochemical investigations of the mechanism of assembly of the active-site H-cluster of [FeFe]-hydrogenases. J Am Chem Soc 138(46):15227-15233.  doi: 10.1021/jacs.6b09366
  • Noth J, Esselborn J, Güldenhaupt J, Brünje A, Sawyer A, Apfel UP, Gerwert K, Hofmann E, Winkler M, Happe T (2016): [FeFe]-Hydrogenase with chalcogenide substitutions at the H-cluster maintains full H2 evolution activity. Angew Chem Int Ed Engl 55:8396-8400.  doi: 10.1002/anie.201511896
  • Esselborn J, Muraki N, Klein K, Engelbrecht V, Metzler-Nolte N, Apfel UP, Hofmann E, Kurisu G, Happe T (2016): A structural view of synthetic cofactor integration into [FeFe]-hydrogenases. Chem Sci 7:959-968.  doi: 10.1039/C5SC03397G


  • Noth J, Kositzki R, Klein K, Winkler M, Haumann M, Happe T (2015): Lyophilization protects [FeFe]-hydrogenases against O2-induced H-cluster degradation. Sci Rep 5:13978.  doi: 10.1038/srep13978
  • Oughli AA, Conzuelo F, Winkler M, Happe T, Lubitz W, Schuhmann W, Rüdiger O, Plumeré N (2015): A Redox Hydrogel Protects the O2 -Sensitive [FeFe]-Hydrogenase from Chlamydomonas reinhardtii from Oxidative Damage. Angew Chem Int Ed Engl 54(42):12329-12333.  doi: 10.1002/anie.201502776
  • Huwald D, Schrapers P, Kositzki R, Haumann M, Hemschemeier A (2015): Characterization of unusual truncated hemoglobins of Chlamydomonas reinhardtii suggests specialized functions. Planta 242(1):167-185.  doi:10.1007/s00425-015-2294-4
  • Bachmeier A, Esselborn J, Hexter SV, Krämer T, Klein K, Happe T, McGrady JE, Myers WK, Armstrong FA (2015): How Formaldehyde Inhibits Hydrogen Evolution by [FeFe]-Hydrogenases: Determination by (13)C ENDOR of Direct Fe-C Coordination and Order of Electron and Proton Transfers. J Am Chem Soc 137(16):5381-9.  doi:10.1021/ja513074m
  • Winkler M, Happe T (2015): Catalytic properties and maturation of [FeFe]-hydrogenases. Biohydrogen (Rögner M eds) De Gruyter, In press: 41-60  doi:10.1515/9783110336733.41
  • Hemschemeier A, Happe T (2015): Eukaryotic microalgae in biotechnological applications. Biotechnology. (Frankenberg-Dinkel N, Kück U eds) De Gruyter, In press: 165-196  doi:10.1515/9783110342635-009


  • Applegate AM, Lubner CE, Knörzer P, Happe T, Golbeck JH (2014): Quantum yield measurements of light-induced H2 generation in a photosystem I-[FeFe]-H2ase nanoconstruct. Photosynth Res 127(1):5-11.  doi:10.1007/s11120-014-0064-y
  • Chernev P, Lambertz C, Brünje A, Leidel N, Sigfridsson KG, Kositzki R, Hsieh CH, Yao S, Schiwon R, Driess M, Limberg C, Happe T, Haumann M (2014): Hydride binding to the active site of [FeFe]-hydrogenase. Inorg Chem 53(22):12164-77.  doi:10.1021/ic502047q
  • Adamska-Venkatesh A, Krawietz D, Siebel J, Weber K, Happe T, Reijerse E, Lubitz W (2014): New redox states observed in [FeFe] hydrogenases reveal redox coupling within the H-cluster. J Am Chem Soc 136(32):11339-46.  doi:10.1021/ja503390c
  • Rumpel S, Siebel JF , Farès C, Duan J, Reijerse E, Happe T, Lubitz W, Winkler M (2014): Enhancing hydrogen production of microalgae by redirecting electrons from photosystem I to Hydrogenase. Energy Environ Sci 7:3296-3301.  doi:10.1039/C4EE01444h
  • Happe T, Hemschemeier A (2014): Metalloprotein mimics - old tools in a new light. Trends Biotechnol 32(4):170-6.  doi:10.1016/j.tibtech.2014.02.004


  • Hemschemeier A (2013): Photo-bleaching of Chlamydomonas reinhardtii after dark-anoxic incubation. Plant Signal Behav 8(11):e27263.  doi:10.4161/psb.27263
  • Lambertz C, Chernev P, Klingan K, Leidel N, Sigfridsson K G. V., Happe T, Haumann M (2013): Electronic and molecular structures of the active-site H-cluster in [FeFe]-hydrogenase determined by site-selective X-ray spectroscopy and quantum chemical calculations. Chem Sci 5(3):1187-1203.  doi:10.1039/C3SC52703D
  • Hemschemeier A, Casero D, Liu B, Benning C, Pellegrini M, Happe T, Merchant SS (2013): COPPER RESPONSE REGULATOR1-Dependent and -Independent Responses of the Chlamydomonas reinhardtii Transcriptome to Dark Anoxia. Plant Cell 25(9):3186-211.  doi:10.1105/tpc.113.115741
  • Esselborn J, Lambertz C, Adamska-Venkatesh A, Simmons T, Berggren G, Noth J, Siebel J, Hemschemeier A, Artero V, Reijerse E, Fontecave M, Lubitz W, Happe T (2013): Spontaneous activation of [FeFe]-hydrogenases by an inorganic [2Fe] active site mimic. Nature Chem Biol 9(10):607-9.  doi:10.1038/nchembio.1311
  • Berggren G, Adamska A, Lambertz C, Simmons TR, Esselborn J, Atta M, Gambarelli S, Mouesca JM, Reijerse E, Lubitz W, Happe T, Artero V, Fontecave M (2013): Biomimetic assembly and activation of [FeFe]-hydrogenases. Nature 499(7456):66-9.  doi:10.1038/nature12239
  • Hemschemeier A, Düner M, Casero D, Merchant SS, Winkler M, Happe T (2013): Hypoxic survival requires a 2-on-2 hemoglobin in a process involving nitric oxide. Proc Natl Acad Sci U S A 110(26):10854-9.  doi:10.1073/pnas.1302592110
  • Li-Beisson Y, Peltier G, Knörzer P, Happe T, Hemschemeier A (2013): Hydrogen and biofuel production in the chloroplast. Plastid biology in the series Advances in Plant Biology. (Theg S, Wollman FA (eds)) Springer, In press: 559-585.  doi:10.1007/978-1-4939-1136-3_19
  • Winkler M, Esselborn J, Happe T (2013): Molecular basis of [FeFe]-hydrogenase function: An insight into the complex interplay between protein and catalytic cofactor. Biochim Biophys Acta 1827(8-9):974-85.  doi:10.1016/j.bbabio.2013.03.004
  • Noth J, Krawietz D, Hemschemeier A, Happe T (2013): Pyruvate:ferredoxin oxidoreductase is coupled to light-independent hydrogen production in Chlamydomonas reinhardtii. J Biol Chem 288(6):4368-77.  doi:10.1074/jbc.M112.429985


  • Adamska A, Silakov A, Lambertz C, Rüdiger O, Happe T, Reijerse E, Lubitz W (2012): Identification and Characterization of the "Super-Reduced" State of the H-Cluster in [FeFe] Hydrogenase: A New Building Block for the Catalytic Cycle? Angew Chem Int Ed Engl 51(46):11458–11462.  doi:10.1002/anie.201204800
  • Hexter SV, Grey F, Happe T, Climent V, Armstrong FA (2012): Electrocatalytic mechanism of reversible hydrogen cycling by enzymes and distinctions between the major classes of hydrogenases. Proc Natl Acad Sci U S A 109(29):11516-21.  doi:10.1073/pnas.1204770109
  • Pape M, Lambertz C, Happe T, Hemschemeier A (2012): Differential Expression of the Chlamydomonas [FeFe]-Hydrogenase-Encoding HYDA1 Gene Is Regulated by the COPPER RESPONSE REGULATOR1. Plant Physiol 159(4):1700-12.  doi:10.1104/pp.112.200162
  • Happe T, Lambertz C, Kwon JH, Rexroth S, Rögner M (2012): Hydrogen Production by natural and semiartificial systems; in: "Microalgal Biotechnology: Integration and economy", Posten C, Walter C (eds), De Gruyter Berlin 2012:111-128.  doi:10.1515/9783110298321.111
  • Foster CE, Krämer T, Wait AF, Parkin A, Jennings DP, Happe T, McGrady JE, Armstrong FA (2012): Inhibition of [FeFe]-Hydrogenases by Formaldehyde and Wider Mechanistic Implications for Biohydrogen Activation. J Am Chem Soc 134(17):7553-7.  doi:10.1021/ja302096r
  • Knörzer P, Silakov A, Foster CE, Armstrong FA, Lubitz W, Happe T (2012): Importance of the Protein Framework for Catalytic Activity of [FeFe]-Hydrogenases. J Biol Chem 287(2):1489-99.  doi:10.1074/jbc.M111.305797
  • Philipps G, Happe T, Hemschemeier A (2012): Nitrogen deprivation results in photosynthetic hydrogen production in Chlamydomonas reinhardtii. Planta 235(4):729-45.  doi:10.1007/s00425-011-1537-2


  • Lubner CE, Applegate AM, Knörzer P, Ganago A, Bryant DA, Happe T, Golbeck JH (2011): Solar hydrogen-producing bionanodevice outperforms natural photosynthesis. Proc Natl Acad Sci U S A 108(52):20988-91.  doi:10.1073/pnas.1114660108
  • Lambertz C, Leidel N, Havelius KG, Noth J, Chernev P, Winkler M, Happe T, Haumann M (2011): O2-reactions at the six-iron active site (H-cluster) in [FeFe]-hydrogenase. J Biol Chem 286(47):40614-23.  doi:10.1074/jbc.M111.283648
  • Tolleter D, Ghysels B, Alric J, Petroutsos D, Tolstygina I, Krawietz D, Happe T, Auroy P, Adriano JM, Beyly A, Cuine S, Plet J, Reiter IM, Genty B, Cournac L, Hippler M, Peltier G (2011): Control of Hydrogen Photoproduction by the Proton Gradient Generated by Cyclic Electron Flow in Chlamydomonas reinhardtii. Plant Cell 23(7):2619-2630.  doi:10.1105/tpc.111.086876
  • Scoma A, Krawietz D, Faraloni C, Giannelli L, Happe T, Torzillo G (2011): Sustained H(2) production in a Chlamydomonas reinhardtii D1 protein mutant. J Biotechnol 157(4):613-619.  doi:10.1016/j.jbiotec.2011.06.019
  • Winkler M, Kawelke S, Happe T (2011): Light driven hydrogen production in protein based semi-artificial systems. Bioresur Technol 102(18):8493-8500.  doi:10.1016/j.biortech.2011.05.019
  • Hemschemeier A, Happe T (2011): Alternative photosynthetic electron transport pathways during anaerobiosis in the green alga Chlamydomonas reinhardtii. Biochim Biophys Acta 1807(8):919-926.  doi:10.1016/j.bbabio.2011.02.010
  • Krassen H, Stripp ST, Böhm N, Berkessel A, Happe T, Ataka K, Heberle J (2011): Tailor-Made Modification of a Gold Surface for the Chemical Binding of a High-Activity [FeFe] Hydrogenase. Eur J Inorg Chem 7:1138-1146.  doi:10.1002/ejic.201001190
  • Philipps G, Krawietz D, Hemschemeier A, Happe T (2011): A pyruvate formate lyase-deficient Chlamydomonas reinhardtii strain provides evidence for a link between fermentation and hydrogen production in green algae. Plant J 66(2):330-340.  doi:10.1111/j.1365-313X.2011.04494.x
  • Wait AF, Brandmayr C, Stripp S, Cavazza C, Fontecilla-Camps JC, Happe T, Armstrong FA (2011): Formaldehyde-A Rapid and Reversible Inhibitor of Hydrogen Production by [FeFe]-Hydrogenases. J Am Chem Soc 133(5):1282-1285.  doi:10.1021/ja110103p


  • Hemschemeier A, Müllner K, Rühle T, Happe T (2010): Hydrogen generation by microbial cultures. Biomass to biofuels. (Vertes A, Qureshi N, Blaschek H, Yukawa H (eds)), Wiley and Sons, Ltd, West Sussex, UK, 359-385.  doi:10.1002/9780470750025.ch18
  • Czech I, Stripp S, Sanganas O, Leidel N, Happe T, Haumann M (2010): The [FeFe]-hydrogenase maturation protein HydF contains a H-cluster like [4Fe4S]-2Fe site. FEBS Lett 585(1):225-230.  doi:10.1016/j.febslet.2010.11.052
  • Lubner CE, Knörzer P, Silva PJ, Vincent KA, Happe T, Bryant DA, Golbeck JH (2010): Wiring an [FeFe]-Hydrogenase with Photosystem I for Light-Induced Hydrogen Production. Biochemistry 49(48):10264-10266.  doi:10.1021/bi1016167
  • Lambertz C, Hemschemeier A, Happe T (2010): Anaerobic expression of the ferredoxin-encoding FDX5 gene of Chlamydomonas reinhardtii is regulated by the Crr1 transcription factor. Eukaryot Cell 11:1747-1754.  doi:10.1128/EC.00127-10
  • Winkler M, Hemschemeier A, Jacobs J, Stripp S, Happe T (2010): Multiple ferredoxin isoforms in Chlamydomonas reinhardtii - Their role under stress conditions and biotechnological implications. Eur J Cell Biol 12:998-1004.  doi:10.1016/j.ejcb.2010.06.018
  • Calusinska M, Happe T, Joris B, Wilmotte A (2010): The surprising diversity of clostridial hydrogenases: a comparative genomic perspective. Microbiology 156:1575-1588.  doi:10.1099/mic.0.032771-0
  • Czech I, Silakov A, Lubitz W, Happe T (2010): The[FeFe]-hydrogenase maturase HydF from Clostridium acetobutylicum contains a CO and CN-ligated iron cofactor. FEBS lett 584(3):638-642.  doi:10.1016/j.febslet.2009.12.016


  • Winkler M, Kuhlgert S, Hippler M, Happe T (2009): Characterization of the key step for light-driven hydrogen evolution in green algae. J Biol Chem 284(52):36620-36627.  doi:10.1074/jbc.M109.053496
  • Stripp S, Happe T (2009) How Algae produce Hydrogen – News from the photosynthetic hydrogenase. Dalton Trans 7(45):9960-9969.  doi:10.1039/b916246a
  • Stripp S, Goldet G, Brandmayr C, Sanganas O, Vincent KA, Haumann M, Armstrong FA, Happe T(2009): How oxygen attacks [FeFe] hydrogenases from photosynthetic organisms. PNAS 106(41):17331-17336.  doi:10.1073/pnas.0905343106
  • Goldet G, Brandmayr C, Stripp S, Happe T, Cavazza C, Fontecilla-Camps J, Armstrong FA (2009): Electrochemical Kinetic Investigations of the Reactions of [FeFe]-hydrogenases with Carbon Monoxide and Oxygen: Comparing the Importance of Gas Tunnels and Active-Site Electronic/Redox Effects. J Am Chem Soc 131(41):14979-14989.  doi:10.1021/ja905388j
  • Silakov A, Kamp S, Reijerse E, Happe T, Lubitz W (2009) Spectroelectrochemical characterization of the active site of the [FeFe] hydrogenase HydA1 from Chlamydomonas reinhardtii. Biochemistry 48:7780-7786.  doi:10.1021/bi9009105
  • Hemschemeier A, Melis A, Happe T(2009): Analytical approaches to photobiological hydrogen production in unicellular green algae. Photosynth Res 2009 Mar 17.  doi:10.1007/s11120-009-9415-5
  • Stripp S, Sanganas O, Happe T, Haumann M (2009): The structure of the active site H-cluster of [FeFe] hydrogenase from the green alga Chlamydomonas reinhardtii studied by X-ray absorption spectroscopy. Biochemistry 48:5042-5049.  doi:10.1021/bi900010b
  • Krassen H, Stripp S, von Abendroth G, Ataka K, Happe T, Heberle J (2009): Immobilization of the [FeFe]-hydrogenase CrHydA1 on a gold electrode: design of a catalytic surface for the production of molecular hydrogen. J Biotechnol 142:3-9.  doi:10.1016/j.jbiotec.2009.01.018
  • Jacobs J, Pudollek S, Hemschemeier A, Happe T (2009): A novel, anaerobically induced ferredoxin in Chlamydomonas reinhardtii. FEBS Lett 583:325-329.  doi:10.1016/j.febslet.2008.12.018


  • Rühle T, Hemschemeier A, Melis A, Happe T (2008): A novel screening protocol for the isolation of hydrogen producing Chlamydomonas reinhardtii strains. BMC Plant Biology 8:107-112.  doi:10.1186/1471-2229-8-107
  • von Abendroth G, Stripp S, Silakov A, Croux C, Soucaille P, Girbal L, Happe T (2008): Optimized over-expression of [FeFe] hydrogenases with high specific activity in Clostridium acetobutylicum. Int J Hydrogen Energy 33:6076-6081.  doi:10.1016/j.ijhydene.2008.07.122
  • Hemschemeier A, Fouchard S, Cournac L, Peltier G, Happe T (2008): Hydrogen production by Chlamydomonas reinhardtii: an elaborate interplay of electron sources and sinks. Planta 227:397-407.  doi:10.1007/s00425-007-0626-8
  • Kamp C, Silakov A, Winkler M, Reijerse EJ, Lubitz W, Happe T (2008): Isolation and first EPR characterization of the [FeFe]-hydrogenases from green algae. Biochim Biophys Acta 1777:410-416.  doi:10.1016/j.bbabio.2008.02.002
  • Hemschemeier A, Jacobs J, Happe T (2008): Biochemical and physiological characterization of the pyruvate formate-lyase Pfl1 of Chlamydomonas reinhardtii, a typically bacterial enzyme in a eukaryotic alga. Eukaryot Cell 7:518-526. doi:10.1128/EC.00368-07


  • Happe T, Müllner K (2007) Biofuel from algae - photobiological hydrogen production and CO2-fixation. Int J of Energy Technology and Policy 5:290-295.  doi:10.1504/IJETP.2007.014735


  • Fouchard S, Hemschemeier A, Caruana A, Pruvost J, Legrand J, Happe T, Peltier G, Cournac L (2005): Autotrophic and mixotrophic hydrogen photoproduction in sulphur-deprived Chlamydomonas cells. Appl Env Microbiol 71:6199-6205.  10.1128/AEM.71.10.6199-6205.2005
  • Girbal L, von Abendroth G, Winkler M, Benton P, Meynial-Salles I, Croux C, Peters J, Happe T, Soucaille P (2005): Homologous and Heterologous Overexpression in Clostridium acetobutylicum and Characterization of Purified Clostridial and Algal Fe-Only Hydrogenases with High Specific Activity. Appl Env Microbiol 71:2777-2781.  doi:10.1128/AEM.71.5.2777-2781.2005
  • Hemschemeier A, Happe T (2005): The exceptional photofermentative hydrogen metabolism of the green alga Chlamydomonas reinhardtii. Biochem Soc Trans 33:39-41.  doi:10.1042/BST0330039


  • Winkler M, Maeurer C, Hemschemeier A, Happe T (2004): The isolation of green algal strains with outstanding H2-productivity. Biohydrogen III, (Miyake J, Igarashi Y, Rögner M, eds), Elsevier Science, Oxford, 103-115.  doi:10.1016/B978-008044356-0/50009-9
  • Stirnberg M, Happe T (2004): Identification of a cis-acting element controlling anaerobic expression of the hydA gene from Chlamydomonas reinhardtii. Biohydrogen III, (Miyake J, Igarashi Y, Rögner M, eds), Elsevier Science, Oxford, 117-127.  doi:10.1016/B978-008044356-0/50010-5
  • Melis A, Seibert M, Happe T (2004): Genomics of green algal hydrogen research. Photosyn Res 82:277-288.  doi:10.1007/s11120-004-2050-2
  • Melis A, Happe T (2004): Trails of green alga hydrogen research - from Hans Gaffron to new frontiers. Photosyn Res 80:401-409.  doi:10.1023/B:PRES.0000030421.31730.cb
  • Schütz K, Happe T, Troshina O, Lindblad P, Leitao E, Oliveira P, Tamagnini P (2004): Cyanobacterial H(2) production - a comparative analysis. Planta 218:350-359.  doi:10.1007/s00425-003-1113-5


  • Forestier M, King P, Zhang L, Posewitz M, Schwarzer S, Happe T, Ghirardi ML, Seibert M (2003): Expression of two [Fe]-hydrogenases in Chlamydomonas reinhardtii under anaerobic conditions. Eur J Biochem 270:2750-2758.  doi:10.1046/j.1432-1033.2003.03656


  • Winkler M, Hemschemeier A, Gotor C, Melis A, Happe T (2002): [Fe]-hydrogenases in green algae: photo-fermentation and hydrogen evolution under sulfur deprivation. Int J Hydrogen Energy 27:1431-1439.  doi:10.1016/S0360-3199(02)00095-2
  • Schmitz O, Boison G, Salzmann H, Bothe H, Schütz K, Wang SH, Happe T (2002): HoxE - a subunit specific for the pentameric bidirectional hydrogenase complex (HoxEFUYH) of cyanobacteria. Biochim Biophys Acta 1554:66-74.  doi:10.1016/S0005-2728(02)00214-1
  • Lindberg P, Schütz K, Happe T, Lindblad P (2002): A hydrogen-producing, hydrogenase-free mutant strain of Nostoc punctiforme ATCC 29133. Int J Hydrogen 27:1291-1296.  doi:10.1016/S0360-3199(02)00121-0
  • Schiefer W, Schütz, K, Hachtel W, Happe T (2002): Molecular cloning and characterization of hetR genes from filamentous cyanobacteria. Biochim Biophys Acta 1577:139-143.  doi:10.1016/S0167-4781(02)00399-8
  • Winkler M, Heil, B, Heil B, Happe T (2002): Isolation and molecular characterization of the [Fe]-hydrogenase from the unicellular green alga Chlorella fusca. BBA 1576:330-334.  doi:10.1016/S0167-4781(02)00239-7
  • Happe T, Hemschemeier A, Winkler M, Kaminski A (2002): Hydrogenases in green algae: do they save the algae´s life and solve our energy problems? Trends Plant Sci 7:246-250.  doi:10.1016/S1360-1385(02)02274-4
  • Horner D, Heil B, Happe T, Embley M (2002): Iron hydrogenases - ancient enzymes in modern eukaryotes. Trends Biochem Sci 27:148-153.  doi:10.1016/S0968-0004(01)02053-9
  • Happe T, Kaminski A (2002): Differential regulation of the [Fe]-hydrogenase during anaerobic adaptation in the green alga Chlamydomonas reinhardtii. Eur J Biochem 269:1022-1032.  doi:10.1046/j.0014-2956.2001.02743.x