in English
Photosynthesis - from regulation of light reactions towards applications to boost bioeconomy (FCoE project)


Eva-Mari Aro, Academician, Professor

e-mail: evaaro [at]
tel. +358 2 333 5931
ARO_CV_Sep2017.pdfCV Aro Eva-Mari



Eva-Mari Aro, Academician, Professor, Project Leader
Natalia Battchikova, PhD, Senior Research Fellow
Duncan Fitzpatrick, PhD, Postdoctoral Researcher
Caterina Gerotto, PhD, Postdoctoral Researcher
Peter Gollan, PhD, Postdoctoral Researcher
Martina Jokel, PhD, Postdoctoral Researcher
Pauli Kallio​, PhD, Postdoctoral Researcher
Henna Mustila, PhD, Postdoctoral Researcher
Dorota Muth-Pawlak, PhD, Postdoctoral Researcher
Csaba Nagy, PhD, Postdoctoral Researcher
Pekka Patrikainen PhD, Postdoctoral Researcher
Arjun Tiwari, PhD, Postdoctoral Researcher
Mikko Tikkanen, PhD, Postdoctoral Researcher
Andrea Trotta, PhD, Postdoctoral Researcher
Martina Angeleri, MSc, Doctoral Student 
Veronica Carbonell, MSc, Doctoral Student
Hariharan Dandapani, MSc, Doctoral Student
Tuomas Huokko, MSc, Doctoral Student
Janne Isojärvi, MSc, Doctoral Student
Sanna Kreula MSc, Doctoral Student
Jari Kämäräinen, MSc, Doctoral Student
Pasi Paananen, MSc, Doctoral Student
Marjaana Rantala, MSc, Doctoral Student
Sanna Rantala, MSc, Doctoral Student
Kati Thiel​, MSc, Doctoral Student
Linda Vuorijoki MSc, Doctoral Student 
Eerika Vuorio, MSc, Doctoral Student
Julia Walter, MSc, Doctoral Student
Maija Lespinasse, MSc, Coordinator, Project Researcher


Photosynthesis harvests solar energy and converts it to chemical form to support all life on Earth. Oxygenic photosynthesis, first evolved in cyanobacteria, has directed the evolution of life as we experience it today. Photosynthesis research is needed to guarantee the basic requirements of life with increasing human population, in order to enhance the production of food and feed, biomass-based energy, direct biosolarfuels and various commodity as well as high-value chemicals. Cyanobacteria-based cell factories are designed and constructed for production of direct biosolarfuels and various fossil fuel-based chemicals, without relying on biomass accumulation. A deep understanding of the structure, function and regulation of the photosynthetic energy capture and storage is of fundamental importance to efficiently exploit the photosynthetic reactions for the wellbeing of the future generations.


Our basic research focuses on regulation of photosynthesis, which is intimately linked with plant stress responses and signalling cascades to regulate nuclear gene expression. Particular emphasis is also put on evolutionary aspects of regulation of thylakoid light harvesting and energy conversion systems. A diverse set of photoautotrophic oxygen evolving organisms, from cyanobacteria, green algae Chlamydomonas reinhardtii and diatoms to a moss Physcomitrella patens and ferns as well as to the model higher plants Arabidopsis thaliana and spruce, are used in our research (Figure 1). We aim at understanding (i) the mechanisms and dynamic regulation of the rapid photophysical and photochemical processes of the light energy capture and the photosynthetic electron transfer (Figure 2), (ii) how the structure and organisation of the thylakoid protein complexes dynamically respond to environmental changes (Figure 3, Figure 4 and Figure 5) and (iii) how the signals originating from the photosynthetic apparatus relay information to the nuclear genome for regulation of gene expression and subsequent acclimation of the photosynthetic organism to changing environmental conditions (Figure 6). Likewise (iv) the evolution of thylakoid regulatory mechanisms, with special emphasis on the replacement of the cyanobacterial flavodiiron proteins with other regulatory mechanisms in higher plants, is investigated to understand how it was possible for photosynthetic organisms to move from oceans to the land (Figure 7). 
Our applied research focuses on the production of carbon–neutral biosolarfuels and chemicals using cyanobacteria as a production chassis (Figure 8). In the synthetic biology program, we aim at production of fuels and chemicals with high photon conversion efficiency, significantly higher compared to natural pathways. The main goal is economically viable conversion of solar energy to a fuel using inexhaustible raw materials: sunlight, water and CO2. Applied research is intimately integrated with our basic research and comprises the designing, engineering and optimisation of light harvesting, electron transfer and metabolic pathways.


In press

Järvi S, Rantala M, Aro EM. 2017. Oxygenic Photosynthesis - Light Reactions within the Frame of Thylakoid Architecture and Evolution. In: Barber J. and Ruban AV (eds.) Photosynthesis and Bioenergetics (in press)
Rantala M, Tikkanen M, Aro EM (2017) Proteomic characterization of hierarchical megacomplex formation in Arabidopsis thylakoid membrane. Plant J.  (in press) (Epub 2017 Oct 4) open access
Tikkanen M, Grebe S (2017) Switching off photoprotection of photosystem I - a novel tool for gradual PSI photoinhibition (in press) (Epub 2017 Aug 16) open access


Bersanini L, Allahverdiyeva Y, Battchikova N, Heinz S, Lespinasse M, Ruohisto E, Mustila H, Nickelsen J, Vass I, Aro EM. 2017. Dissecting the photoprotective mechanism encoded by the flv4-2 operon: a distinct contribution of Sll0218 in photosystem II stabilization. Plant Cell Environ. 40(3): 378-38   (Epub 2016 Dec 7) open access
Fristedt R, Trotta A, Suorsa M, Nilsson AK, Croce R, Aro EM, Lundin B. 2017. PSB33 sustains photosystem II D1 protein under fluctuating light conditions. J Exp Bot. 68(15): 4281-4293 (Epub 2017 Jul 10) open access
Georg J, Kostova G, Vuorijoki L, Schön V, Kadowaki T, Huokko T, Baumgartner D, Müller M, Klähn S, Allahverdiyeva Y, Hihara Y, Futschik ME, Aro EM, Hess WR. 2017. Acclimation of oxygenic photosynthesis to iron starvation is controlled by the sRNA IsaR1. Curr Biol. 27: 1-12 (Epub 2017 May 5) open access
Giovanardi M, Poggioli M, Ferroni L, Lespinasse M, Baldisserotto C, Aro EM, Pancaldi S. 2017. Higher packing of thylakoid complexes ensures a preserved Photosystem II activity in mixotrophic Neochloris oleoabundans. Algal research 25: 255-265
Huokko T, Muth-Pawlak D, Battchikova N, Allahverdiyeva Y, Aro EM. 2017. Role of type 2 NAD(P)H dehydrogenase NdbC in regulation of carbon allocation in Synechocystis 6803. Plant Phys 174(3): 1863-1880 (Epub 2017 May 22) open access
Ilík P, Pavlovič A, Kouřil R, Alboresi A, Morosinotto T, Allahverdiyeva Y, Aro EM, Yamamoto H, Shikanai T. 2017. Alternative electron transport mediated by flavodiiron proteins is operational in organisms from cyanobacteria up to gymnosperms. New Phytol. 214(3):967-972 (Epub 2017 Mar 17)
Kämäräinen J, Huokko T, Kreula S, Jones PR, Aro EM, Kallio P. 2016. Pyridine nucleotide transhydrogenase PntAB is essential for optimal growth and photosynthetic integrity under low-light mixotrophic conditions in Synechocystis sp. PCC 6803. New Phytol. 214(1): 194-204
Niemi M, Aro EM. 2017. Koulu-uudistuksen käyntiinpanon unohdetut arkkitehdit. Tieteessä Tapahtuu 35(4):9-16 (Epub 2017 Jun 14)
Patrikainen P, Carbonell V, Thiel K, Aro EM, Kallio P. 2017. Comparison of orthologous cyanobacterial aldehyde deformylating oxygenases in the production of volatile C3-C7 alkanes in engineered E. coli. Metabolic Engineering Communications 5: 9-18 (Epub 2017 May 8) open access
Ramos-León F, Mariscal V, Battchikova N, Aro EM, Flores E. 2017. Septal protein SepJ from the heterocyst-forming cyanobacterium Anabaena forms multimers and interacts with peptidoglycan. FEBS openbio 7(10): 1515-1526 (2017 Aug 30) open access
Thiel K, Vuorio E, Aro EM, Kallio PT. 2017. The effect of enhanced acetate influx on Synechocystis sp. PCC 6803 metabolism. Microbial Cell Factories 16(21): 1-12  (Epub 2017 Feb 2) open access
Tikkanen M, Rantala S, Grieco M, Aro EM. 2017. Comparative analysis of mutant plants impaired in the main regulatory mechanisms of photosynthetic light reactions - From biophysical measurements to molecular mechanisms. Plant Phys Biochem. 112: 290-301 (Epub 2017 Jan 17) open access

Vuorijoki L, Kallio P, Aro EM. 2017. SRM dataset of the proteome of inactivated iron-sulfur cluster biogenesis regulator SufR in Synechocystis sp. PCC 6803. Biochim Biophys Acta. 11: 572-575 (Epub 2017 Feb 16) open access
Vuorijoki L, Tiwari A, Kallio P, Aro EM. 2017. Inactivation of iron-sulfur cluster biogenesis regulator SufR in Synechocystis sp. PCC 6803 induces unique iron-dependent protein-level responses. Biochim Biophys Acta. 1861: 1085-1098 (Epub 2017 Feb 16) open access
Wittenberg G, Järvi S, Hojka M, Tóth SZ, Meyer EH, Aro EM, Schöttler MA, Bock R. 2017. Identification and characterization of a stable intermediate in photosystem I assembly in tobacco. Plant J. 90(3):478-490


Angeleri M, Muth-Pawlak D, Aro EM, Battchikova N. 2016. Study of O-phosphorylation sites in proteins involved in photosynthesis-related processes in Synechocystis sp. strain PCC 6803: Application of the SRM approach. J Proteome Res. 15(12): 4638-4652
Aro EM. 2016. From first generation biofuels to advanced solar biofuels. Ambio 45(1): S24-S31 (Epub 2015 Dec 14) open access
Carbonell V, Vuorio E, Aro EM, Kallio P. 2016. Sequence optimization of efegene from P. syringae is not required for stable ethylene production in recombinant Synechocystis sp. PCC6803. IJIRTS 4(1): 30-35
Colombo M, Suorsa M, Rossi F, Ferrari R, Tadini L, Barbato R, Pesaresi P. 2016.  Photosynthesis Control: An underrated short-term regulatory mechanism essential for plant viability. Plant Signalling & behavior. 11(4): e1165382 pp.1-6
Ermakova M, Huokko T, Richaud P, Bersanini L, Howe JC, Lea-Smith JD, Peltier G and Allahverdiyeva Y. 2016. Distinguishing the roles of thylakoid respiratory terminal oxidases in the cyanobacterium Synechocystis sp. PCC 6803. PLANT PHYSIOLOGY, 171(2):1307-19 (Epub 2016 Apr 18) open access
Ferroni L, Suorsa M, Aro EM, Baldisserotto C, Pancaldi S. 2016. Light acclimation in the lycophyte Selaginella martensii depends on changes in the amount of photosystems and on the flexibility of the light-harvesting complex II antenna association with both photosystems. New Phytol. 211(2): 554-568  
Gao F, Zhao F, Chen L, Battchikova N, Ran Z, Aro EM, Ogawa T, Ma W. 2016. Novel NDH-1L-CpcG2-Phycobilisome-Photosystem I Supercomplex Is Important for Efficient Cyclic Electron Transport in Synechocystis sp. Strain PCC 6803. Plant Physiol. 172(3): 1451-1464
Gerotto C, Alboresi A, Meneghesso A, Jokel M, Suorsa M, Aro EM, Morosinotto T. 2016. Flavodiiron proteins act as safety valve for electrons in Physcomitrella patens. Proc Natl Acad Sci. 113(43): 12322-12327
Grouneva I, Muth-Pawlak D, Battchikova N, Aro EM. 2016. Changes in relative thylakoid protein abundance induced by fluctuating light in the diatom Thalassiosira pseudonana. J Proteome Res. 15(5): 1649-1658
Järvi S, Isojärvi J, Kangasjärvi S, Salojärvi J, Mamedov F, Suorsa M and Aro EM. 2016. Photosystem II repair and plant immunity: Lessons learned from Arabidopsis mutant lacking the THYLAKOID LUMEN PROTEIN 18.3. Front Plant Sci. 7(405): 1-13 (Epub 2016 Mar 31) open access
Järvi S, Suorsa M, Tadini L, Ivanauskaite A, Rantala S, Allahverdiyeva Y, Leister D, Aro EM. 2016. FtsH is required for biosynthesis of photosystem I in Arabidopsis thaliana. Plant Physiol. 171(2):1333-1343 (Epub 2016 May 2) open access
Mustila H, Paananen P, Battchikova N, Santana-Sánchez A, Muth-Pawlak D, Hagemann M, Aro EM, Allahverdiyeva Y. 2016. The Flavodiiron Protein Flv3 functions as a homo-oligomer during stress acclimation and is distinct from the Flv1/Flv3 hetero-oligomer specific to the O2 photoreduction pathway. Plant Cell Physiol. 57(7): 1468-148 
Najafpour M, Renger G, Hołyńska M, Moghaddam AN, Aro EM, Carpentier R, Nishihara H, Eaton-Rye J, Shen JR, Allakhverdiev SI. 2016. Manganese compounds as water-oxidizing catalysts: From the natural water-oxidizing complex to nanosized manganese oxide structures. Chem Rev. 116:2886-2936
Peltier G, Aro EM, Shikanai T. 2016. NDH-1 and NDH-2 plastoquinone reductases in oxygenic photosynthesis. Annu Rev Plant Biol. 67: 55-80
Plöchinger M, Torabi S, Rantala M, Tikkanen M, Suorsa M, Jensen PE, Aro EM, Meurer J. 2016. The low molecular weight protein PsaI stabilizes the light-harvesting complex II docking site of Photosystem I. Plant Physiol. 172(1): 450-463
Raleiras P, Khanna N, Miranda H, Mészáros LS, Krassen H, Ho F, Battchikova N, Aro EM, Magnuson A, Lindblad P, Styring S. 2016. Turning around the electron flow in an uptake hydrogenase. EPR spectroscopy and in vivo activity of a designed mutant in HupSL from Nostoc punctiforme. Energy Environ Sci. 9(2): 581-594
Rantala M, Lehtimäki N, Aro EM, Suorsa M. 2016. Downregulation of TAP38/PPH1 enables LHCII hyperphosphorylation in Arabidopsis mutant lacking LHCII docking site in PSI. FEBS Lett. 590(6):787-794
Shikanai T, Aro EM. 2016. Evolution of Photosynthetic NDH-1: Structure and Physiological Function. In: Cramer WA. and Kallas T (eds.) Advances in Photosynthesis and Respiration. Springer Link. Vol 41 pp 51-70, ISBN 978-94-017-7481-9; 978-94-017-7479-6 
Suorsa M, Rossi F, Tadini L, Labs M, Colombo, M, Jahns P, Kater M, Leister D, Finazzi G, Aro EM, Barbato R, Pesaresi P. 2016. PGR5-PGRL1-dependent cyclic electron transport modulates linear electron transport rate in Arabidopsis thaliana. Mol Plant. 9(2): 271-288
Tiwari A, Mamedov F, Grieco M, Suorsa M, Jajoo A, Styring S, TIkkanen M, Aro EM. 2016. Photoprotection revised: damage of iron-sulphur clusters FA and FB turns Photosystem I to quencher of excitation energy. Nat Plants. 2(16035): 1-9
Trotta A, Suorsa M, Rantala M, Lundin B, Aro EM. 2016. Serine and threonine residues of plant STN7 kinase are differentially phosphorylated upon changing light conditions and specifically influence the activity and stability of the kinase. Plant J. 87(5): 484-494 (Epub 2016 Aug 2) open access
Vuorijoki L, Isojärvi J, Kallio P, Kouvonen P, Aro EM, Corthals G, Jones PR, Muth-Pawlak D. 2016.  Development of a quantitative SRM-based proteomics method to study iron metabolism of Synechocystis sp. PCC 6803. J Proteome Res. 15: 266-279
Walter J, Lynch F, Battchikova N, Aro EM, Gollan PJ. 2016. Calcium impacts carbon and nitrogen balance in the filamentous cyanobacterium Anabaena sp. PCC 7120. J Exp Bot. 67(13): 3997-4008 (Epub 2016 Mar 24) open access
Zhang ZS, Jin LQ, Li YT, Tikkanen M, Li QM, Ai XZ, Gao HY. 2016. Ultraviolet-B radiation (UV-B) relieves chilling-light-induced PSI photoinhibition and accelerates the recovery of CO2 assimilation in cucumber (Cucumis sativus L.) leaves. Scientific reports. 6:(34455): 1-10


Allahverdiyeva Y, Isojärvi J, Zhang P, Aro EM. 2015. Cyanobacterial oxygenic photosynthesis is protected by flavodiiron proteins. Life 5: 716-743 (Epub 2015 Mar 9) open access
Allahverdiyeva Y, Suorsa M, Tikkanen M, Aro EM. 2015. Photoprotection of photosystems in fluctuating light intensities. J Exp Bot. 66(9): 2427-36 (Epub 2014 Dec 1) open access
Battchikova N, Angeleri M, Aro EM. 2015.  Proteomic approaches in research of cyanobacterial photosynthesis. Photosynth Res. 126(1): 47-70
Chukhutsina V, Bersanini L, Aro EM, van Amerongen H. 2015. Cyanobacterial flv4-2 operon-encoded proteins optimize light harvesting and charge separation in photosystem II. Mol Plant. 8: 747-761 (Epub 2014 Dec 31) open access
Chukhutsina V, Bersanini L, Aro EM, van Amerongen H. 2015. Cyanobacterial light-harvesting phycobilisomes uncouple from photosystem I during dark-to-light transitions. Sci Rep. 5:14193 (Epub 2015 Sep 21) open access
Gollan PJ, Tikkanen M, Aro EM. 2015. Photosynthetic light reactions; integral to chloroplast retrograde signaling. Curr Opin Plant Biol. 27: 180-191 (Epub 2015  Aug 25) open access
Grieco M, Suorsa M, Jajoo A, Tikkanen M, Aro EM. 2015.  Light-harvesting II antenna trimers connect energetically the entire photosynthetic machinery - including both photosystems II and I. Biochim Biophys acta – Bioenerg. 1847: 607-619
Isojärvi J, Shunmugam S, Sivonen K, Allahverdiyeva Y, Aro EM, Battchikova N. 2015. Draft genome sequence of Calothrix strain 336/3, a novel H2-producing cyanobacterium isolated from a Finnish lake. Genome Announcements. 3(1): e01474-14 (Epub 2015 Jan 22) open access
Jokel M, Kosourov S, Battchikova N, Tsygankov AA, Aro EM, Allahverdiyeva Y. 2015.  Chlamydomonas flavodiiron proteins facilitate acclimation to anoxia during hydrogen production. Plant Cell Physiol. 56: 1598-1607 (Epub 2015 Jun 10) open access
Järvi S, Suorsa M, Aro EM. 2015.  Photosystem II repair in plant chloroplasts – regulation, assisting proteins and shared components with photosystem II biogenesis. Biochim Biophys Acta – Bioenerg. 1847: 900-909
Lynch F, Santana-Sanchez A, Jämsä M, Sivonen K, Aro EM, Allahverdiyeva Y. 2015. Screening native isolates of cyanobacteria and a green alga for integrated wastewater treatment, biomass accumulation and neutral lipid production. Algal research 11: 411-420 (Epub 2015 Jun 3) open access
Martinez DE, Borniego ML, Battchikova N, Aro EM, Tyystjärvi E, Guiamét JJ. 2015.  SASP, a Senescence-Associated Subtilisin Protease, is involved in reproductive development and determination of silique number in Arabidopsis. J Exp Bot. 66: 161-174 (Epub 2014 Nov 4) open access
Mekala NR, Suorsa M, Rantala M, Aro EM, Tikkanen M. 2015.  Plants actively avoid state-transitions upon changes in light intensity - role of light-harvesting complex II protein dephosphorylation in high light. Plant Physiol. 168: 721-734 (Epub 2015 Apr 22) open access
Najafpour MM, Fekete M, Sedigh DJ, Aro EM, Carpentier R, Eaton-Rye JJ, Nishihara H, Shen JR, Allakhverdiev S, Spiccia L. 2015. Damage management in water-oxidizing catalysts: From Photosystem II to nano-sized metal oxides. ACS Catalysis. 5: 1499-1512
Suorsa, M. 2015:  Cyclic electron flow provides acclimatory plasticity for the photosynthetic machinery under various environmental conditions and developmental stages. Frontiers in Plant Science 6(800): 1-8 (Epub 2015 Sep 28) open access
Suorsa M, Rantala M, Mamedov F, Lespinasse M, Trotta A, Grieco M, Vuorio E, Tikkanen M, Järvi S, Aro EM. 2015. Light acclimation involves dynamic re-organisation of the pigment-protein megacomplexes in non-appressed thylakoid domains. Plant J. 84(2): 360-373
Teikari J, Österholm J, Kopf M, Battchikova N, Wahlsten M, Aro EM, Hess WR, Sivonen K. 2015. Transcriptomics and proteomics profiling of Anabaena sp. strain 90 under inorganic phosphorus stress. Appl Environ Microbiol. 81: 5212-5222
Tikkanen M, Rantala S, Aro EM. 2015. Electron flow from PSII to PSI under high light is controlled by PGR5 but not by PSBS. Front Plant Sci. 6:521 (Epub 2015 Jul 8) open access


Allahverdiyeva Y, Aro EM, Kosourov S. 2014. Recent Developments on cyanobacteria and green algae for Biohydrogen Photoproduction and Its Importance in CO2 Reduction. In: Gupta V., Tuohy M., Kubicek C.P., Saddler J. and Xu F. (eds.) Bioenergy Research: Advances and Applications, 1st Edition. Elsevier: 367-387, ISBN 978-0-444-59561-4
Allakhverdiev SI, Aro EM, Tomo T. 2014. Special issue on photosynthesis research for sustainability. – Plant Phys Biochem. 81: 1-2 
Battchikova N, Aro EM. 2014. Proteomics in revealing the composition, acclimation and biogenesis of thylakoid membranes. In: Flores E. and Herrero A. (eds.) The Cell Biology of Cyanobacteria. Caister Academic press Norfolk UK pp: 89-121
Bersanini L, Battchikova N, Jokel M, Rehman A, Vass I, Allahverdiyeva Y, Aro EM. 2014. Flavodiiron protein Flv2/Flv4-related photoprotective mechanism dissipates excitation pressure of photosystem II in co-operation with phycobilisomes in cyanobacteria. Plant Physiol. 164: 805-818
Ermakova M, Battchikova, N, Richaud, P, Leino H., Kosourov S, Isojärvi J, Peltier G, Flores E, Cournac, L, Allahverdiyeva Y, Aro EM. 2014. Heterocyst-specific flavodiiron protein Flv3B enables oxic diazotrophic growth of the filamentous cyanobacterium Anabaena sp. PCC 7120. Proc Natl Acad Sci USA. 111: 11205-11210 (Epub 2014 Jul 7) open access
Fears R, Aro EM, Pais MS, ter Meulen V. 2014.  How should we tackle the global risks to plant health? Trends Plant Sci. 19: 206-208 (Epub 2014 Mar 31)
Ferroni L, Angeleri M, Pantaleoni L, Pagliano C, Longoni P, Marsano F, Aro EM, Suorsa M, Baldisserotto C, Giovanardi M, Cella R, Pancaldi S. 2014.  Light-dependent reversible phosphorylation of the minor photosystem II antenna Lhcb6 (CP24) occurs in lycophytes. Plant J. 77: 893-905
Jada B, Soitamo AJ, Siddiqui SA, Murukesan G, Aro EM, Salakoski T, Lehto K. 2014. Multiple different defense mechanisms are activated in the young transgenic tobacco plants which express the full length genome of the Tobacco mosaic virus and are resistant against this virus. Plos One. 9(9): e107778
Jajoo A, Mekala NR, Tomar RS, Grieco M, Tikkanen M, Aro EM. 2014.  Inhibitory effects of polycyclic aromatic hydrocarbons (PAHs) on photosynthetic performance are not related to their aromaticity. J Photochem Photobiol B. 137: 151-155
Jajoo A, Mekala NR, Tongra T, Tiwari A, Grieco M, Tikkanen M, Aro EM. 2014. Low pH-induced regulation of excitation energy between the two photosystems. FEBS Lett. 588: 970-974
Kangasjärvi S, Tikkanen M, Durian G, Aro EM. 2014. Photosynthetic light reactions – An adjustable hub in basic production and plant immunity signaling. Plant Pyhsiol Biochem. 81: 128-134
Kosourov S, Leino H, Murukesan G, Lynch F, Sivonen K, Tsygankov AA, Aro EM, Allahverdiyeva Y. 2014. Hydrogen photoproduction by immobilized N2-fixing cyanobacteria: understanding the role of the uptake hydrogenase in the long-term process. Appl Environ Microbiol. 80: 5807-17 (Epub 2014 Sep) open access
Leino H, Shunmugam S, Isojärvi J, Oliveira P, Mulo P, Saari L, Battchikova N, Sivonen K, Lindblad P, Aro EM, Allahverdiyeva Y. 2014.  Characterization of ten H2 producing cyanobacteria isolated from the Baltic Sea and Finnish lakes. Int J Hydrog energy. 39: 8983-8991
Mustila H, Allahverdiyeva Y, Isojärvi J, Aro EM, Eisenhut M. 2014. The bacterial-type [4Fe-4S] ferredoxin 7 has a regulatory function under photooxidative stress conditions in the cyanobacterium Synechocystis sp. PCC 6803. Biochim Biophys acta – Bioenerg. 1837: 1293-1304
Pietrzykowska M, Suorsa M, Semchonok DA, Tikkanen M, Boekema EJ, Aro EM, Jansson S. 2014.  The light-harvesting chlorophyll a/b binding proteins Lhcb1 and Lhcb2 play complementary roles during state transitions in Arabidopsis. Plant Cell. 26: 3646-3660 (Epub 2014 Sep 5) open access
Shunmugam S, Jokela J, Wahlsten M, Battchikova N, ur Rehman A, Vass I,  Karonen M, Sinkkonen J, Permi P, Sivonen K, Aro EM, Allahverdiyeva Y. 2014. Secondary metabolite from Nostoc XPORK14A inhibits photosynthesis and growth of Synechocystis PCC 6803. Plant Cell Environ 37: 1371-1381

Suorsa M, Rantala M, Danielsson R, Järvi S, Paakkarinen V, Schröder WP, Styring S, Mamedov F, Aro EM. 2014. Dark-adapted spinach thylakoid protein heterogeneity offers insights into the Photosystem II repair cycle. Biochim Biophys Acta – Bioenerg. 1837:1463-1471
Tikkanen M, Aro EM. 2014. Integrative regulatory network of plant thylakoid energy transduction. Trends Plant Sci. 19: 10-17
Tikkanen M, Gollan PJ, Mekala NR, Isojärvi J, Aro EM. 2014. Light-harvesting mutants show differential gene expression upon shift to high light as a consequence of photosynthetic redox and reactive oxygen species metabolism. Philos Trans R Soc Lond B Biol Sci. 369(1640): 20130229
Tikkanen M, Mekala NR, Aro EM. 2014. Photosystem II photoinhibition-repair cycle protects Photosystem I from irreversible damage. Biochim Biophys Acta – Bioenerg. 1837: 210-215


Allahverdiyeva Y, Suorsa M, Rossi F, Pavesi A, Kater M, Antonacci A, Tadini L, Pribil M, Schneider A, Wanner G, Leister D, Aro EM, Barbato R, Pesaresi P. 2013. Arabidopsis plants devoid of PsbQ and PsbR subunits of the oxygen-evolving complex show altered PSII supercomplex organization and short-term adaptive mechanisms. Plant J. 75: 671-684
Allahverdiyeva Y, Mustila H, Ermakova M, Bersanini L, Richaud P, Ajlani G, Battchikova N, Cournac L, Aro EM. 2013. Flavodiiron proteins Flv1 and Flv3 enable cyanobacterial growth and photosynthesis under fluctuating light. Proc Natl Acad Sci USA. 110: 4111-4116 
Carmel D, Dahlström KM, Holmström M, Allahverdiyeva Y, Battchikova N, Aro EM, Salminen TA, Mulo P. 2013. Structural model, physiology and regulation of Slr0006 in Synechocystis PCC 6803. Arch Microbiol. 195: 727-736
Ermakova M, Battchikova N, Allahverdiyeva Y, Aro EM. 2013. Novel heterocyst-specific flavodiiron proteins in Anabaena sp. PCC 7120. FEBS Lett. 587: 82-87
Ferroni L, Pantaleoni L, Baldisserotto C, Aro EM, Pancaldi S. 2013. Low photosynthetic activity is linked to changes in the organization of photosytem II in the fruit Arum italicum. Plant Physiol Biochem. 63: 140-150
Grouneva I, Gollan PJ, Kangasjärvi S, Suorsa M, Tikkanen M, Aro EM. 2013. Phylogenetic viewpoints on regulation of light harvesting and electron transport in eukaryotic photosynthetic organisms. Planta. 237: 399-412
Järvi S, Gollan P, Aro EM. 2013. Understanding the roles of the thylakoid lumen in photosynthesis regulation. Front Plant Sci 4(434): 1-14
Leoni C, Pietrzykowska M, Kiss A, Suorsa M, Ceci L, Aro EM, Jansson S. 2013. Very rapid phosphorylation kinetics suggest a unique role for Lhcb2 during state transitions in Arabidopsis. Plant J. 72: 236-246 
Nath K, Jajoo A, Poudyala RS, Timilsina R, Park YS, Aro EM, Nam HG, Lee CH. 2013. Towards critical understanding of photosystem II repair mechanisms and its regulation during stress conditions. FEBS Lett. 587: 3372-3381
Shunmugam S, Hinttala R, Lehtimäki N, Miettinen M, Uusimaa J, Majamaa K, Sivonen K, Aro EM, Mulo P. 2013. Nodularia spumigena extract induces upregulation of mitochondrial respiratory chain complexes in spinach (Spinacia oleracea L.). Acta Physiol Plant. 35: 969-974
Sirpiö S, Kanervo E, Aro EM. 2013. Assembly and Turnover of the D1 Protein. In: Lennarz, WJ. and Lane MD. (eds.). Encyclopedia of Biological Chemistry 2nd Edition. Elsevier Inc Academic Press. 508-511. ISBN 978-0123786302
Suorsa M, Grieco M, Järvi S, Gollan PJ, Kangasjärvi S, Tikkanen M, Aro EM. 2013. PGR5 ensures photosynthetic control to safeguard photosystem I under fluctuating light conditions. Plant Signal Behav. 8: 1-6.


Eisenhut M, Georg J, Klähn S, Sakurai I, Silen H, Zhang P, Hess RW, Aro EM. 2012. The antisense RNA As1_flv4 in the cyanobacterium Synechocystis sp. PCC6803 prevents premature expression of the flv4-2 operon upon shift in inorganic carbon supply. J Biol Chem. 287: 33153-33162
Gollan PJ, Bhave M, Aro EM. 2012. The FKBP families of higher plants; exploring the structures and functions of protein interaction specialists. FEBS Lett. 586: 3539-3547
Grieco M, Tikkanen M, Paakkarinen V, Kangasjärvi S, Aro EM. 2012. Steady-state phosphorylation of light-harvesting complex II proteins preserves Photosystem I under fluctuating white light. Plant Physiol. 160: 1896-1910
Hebbelmann I, Selinski J, Wehmeyer C, Goss T, Voss I, Mulo P, Kangasjärvi S, Aro EM, Oelze ML, Dietz KJ, Nunes-Nesi A, Do PT, Fernie AR, Talla SK, Raghavendra AS, Linke V, Scheibe R. 2012. Multiple strategies to prevent oxidative stress in Arabidopsis plants lacking the malate valve enzyme NADP-malate dehydrogenase. J Exp Bot. 63: 1445-1459
Kangasjärvi S, Neukerman J, Aro EM, Noctor G. 2012.  Photosynthesis, photorespiration, and light signalling in defence responses. J Exp Bot. 63: 1619-1636
Kobayashi K, Baba S, Obayashi T, Sato M, Toyooka K, Keränen M, Aro EM, Fukaki H, Ohta H, Sugimoto K, Masuda T. 2012. Regulation of Root Greening by Light and Auxin/Cytokinin Signaling in Arabidopsis. Plant Cell. 24: 1081-1095
Kämäräinen J, Knoop H, Stanford NJ, Guerrero F, Akhtar MK, Aro EM, Steuer R, Jones PR. 2012. Physiological tolerance and stoichiometric potential of cyanobacteria for hydrocarbon fuel production. J Biotechnol. 162: 67-74
Leino H, Kosourov S, Saari S, Sivonen K, Tsygankov A, Aro EM, Allahverdiyeva Y. 2012.  Extended H2 photoproduction by N2-fixing cyanobacteria immobilized in thin alginate films. Int J Hydrog Energy. 37: 151-161
Lintala M, Lehtimäki M, Benz JP, Jungfer A, Soll J, Aro EM, Bölter B, Mulo P. 2012. Depletion of the leaf-type ferredoxin-NADP+ oxidoreductase results in permanent induction of photoprotective mechanisms in Arabidopsis chloroplasts. Plant J. 70: 809-817
Mulo P, Sakurai I, Aro EM. 2012. Strategies for psbA gene expression in cyanobacteria, algae and higher plants. From transcription to PSII repair. Biochim Biophys Acta – Bioenerg. 1817: 247-257
Najafpour MM, Moghaddam AN, Yang YN, Aro EM, Carpentier R, Eaton-Rye JJ, Lee CH, Allakhverdiev SI. 2012. Biological water oxidizing complex: a nano-sized manganese-calcium oxide in a protein environment. Photos Res 114: 1-13
Najafpour MM, Rahimi F, Aro EM, Lee CH, Allakhverdiev SI. 2012. Nano-sized manganese oxides as biomimetic catalysts for water oxidation in artificial photosynthesis: a review. J. R. Soc. Interface. 9: 2383-2395
Sakurai I, Stazic D, Eisenhut M, Vuorio E, Steglich C, Hess RW, Aro EM. 2012.  Positive regulation of psbA gene expression by cis encoded antisense RNAs in Synechocystis sp. PCC 6803. Plant Physiol. 160: 1000-1010
Suorsa M, Järvi S, Grieco M, Pietrzykowska M, Tikkanen M, Rantala M, Paakkarinen V, Jansson S, Aro EM. 2012. PGR5 is essential for proper acclimation of Arabidopsis photosystem I to naturally and artificially fluctuating light conditions. Plant Cell. 24: 2934-2948
Tikkanen M, Aro EM. 2012. Thylakoid protein phosphorylation in dynamic regulation of photosystem II in higher plants. Biochim Biophys Acta – Bioenerg. 1817: 232-238
Tikkanen M, Gollan PJ, Suorsa M, Kangasjärvi S, Aro EM. 2012. STN7 operates in retrograde signalling through controlling redox balance in the electron transfer chain. Front Plant Sci. 3: 277
Tikkanen M, Grieco M, Nurmi M, Rantala M, Suorsa M, Aro EM. 2012. Regulation of the photosynthetic apparatus under fluctuating growth light. Philos Trans R Soc Lond B Biol Sci. 367: 3486-3493
Tikkanen M, Suorsa M, Gollan P, Aro EM. 2012. Post-genomic insight into thylakoid membrane lateral heterogeneity and redox balance. FEBS Lett. 586: 2911-2916
Zhang P,  Eisenhut M, Brandt AM, Carmel D, Silén HM, Vass I, Allahverdiyeva Y, Salminen TA, Aro EM. 2012. Operon flv4-flv2 provides cyanobacterial photosystem II with flexibility of electron transfer.  Plant Cell. 24: 1952-1971


Allahverdiyeva  Y, Aro EM. 2011. Photosynthetic responses of plants to excess light: Mechanisms and conditions for photoinhibition, excess energy dissipation and repair. In: Eaton-Rye JJ., Tripathy BC. and Sharkey TD. (eds.) Photosynthesis: A comprehensive Treatise Physiology, Biochemistry, Biophysics and Molecular Biology, Advances in Photosynthesis and Respiration series. Springer, Netherlands. Vol 33 pp. 275-297, ISBN 978-94-007-1578-3.
Allahverdiyeva Y, Ermakova M, Eisenhut M, Zhang P, Richaud P, Hagemann M, Cournac L, Aro EM. 2011. Interplay between flavodiiron proteins and photorespiration in Synechocystis sp. PCC 6803. J Biol Chem. 286: 24007-24014
Alperovitch-Lavy A, Sharon I, Rohwer F, Aro EM, Glaser F, Milo R, Nelson N, Béjà O. 2011.  Reconstructing a puzzle: existence of cyanophages containing both photosystem-I and photosystem-II gene suites inferred from oceanic metagenomic datasets. Environ Microbiol. 13: 24-32
Battchikova N, Aro EM, Nixon PJ. 2011. Structure and physiological function of NDH-I complexes in cyanobacteria. In: Peschek GA., Obinger C. and Renger G. (eds.) Bioenergetic Processes of Cyanobacteria - From Evolutionary Singularity to Ecological Diversity. Berlin, Heidelberg, New York: Springer Verlag, pp. 445-467, ISBN: 978-94-007-0352-0.
Battchikova N, Eisenhut M, Aro EM. 2011. Cyanobacterial NDH-1 complexes: novel insights and remaining puzzles. Biochim Biophys Acta – Bioenerg. 1807: 935-944
Battchikova N, Wei L, Du L, Bersanini L, Aro EM, Ma W. 2011. Identification of a novel Ssl0352 Protein (NdhS), essential for efficient operation of cyclic electron transport around photosystem I, in NADPH:Plastoquinone oxidoreductase (NDH-1) complexes of Synechocystis sp. PCC 6803. J Biol Chem. 286: 36992-37001
Carmel D, Mulo P, Battchikova N, Aro EM. 2011. Membrane attachment of Slr0006 in Synechocystis sp. PCC 6803 is determined by divalent ions. Photos Res. 108: 241-245
Chen G, Allahverdiyeva Y, Aro EM, Styring S, Mamedov F. 2011. Electron paramagnetic resonance study of the electron transfer reactions in photosystem II membrane preparations from Arabidopsis thaliana. Biochim Biophys Acta – Bioenerg, 1807: 205-215
Chen KM, Piippo M, Holmström M, Nurmi M, Pakula E, Suorsa M, Aro EM. 2011. A chloroplast-targeted DnaJ protein AtJ8 is negatively regulated by light and has rapid turnover in darkness. J Plant Physiol. 168: 1780-1783
Grouneva I, Rokka A, Aro EM. 2011.  The thylakoid membrane proteome of two marine diatoms outlines both diatom-specific and species-specific features of the photosynthetic machinery. J Proteome Res. 10: 5338-5353
Ifuku K, Endo T, Shikanai T, Aro EM. 2011.  Structure of the chloroplast NADH dehydrogenase-like complex: nomenclature for nuclear-encoded subunits. Plant Cell Physiol. 52: 1560-8
Järvi S, Suorsa M, Paakkarinen V, Aro EM. 2011.  Optimized native gel systems for separation of thylakoid protein complexes: novel super- and megacomplexes. Biochem J. 439: 207-214
Lehtimäki N, Shunmugam S, Jokela J, Wahlsten M, Carmel D, Keränen M, Sivonen K, Aro EM, Allahverdiyeva Y, Mulo P. 2011. Nodularin uptake and induction of oxidative stress in spinach (Spinachia oleracea). J Plant Physiol. 168: 594-600
Philosof A, Battchikova N, Aro EM, Béjà O. 2011. Marine cyanophages: tinkering with the electron transport chain. Isme J. 5: 1568-1570
Rodina E, Vorobieva N, Kurilova S, Mikulovich J, Vainonen J, Aro EM, Nazarova T. 2011. Identification of new protein complexes of Escherichia coli inorganic pyrophosphatase using pull-down assay. Biochimie. 93: 1576-1583
Rokka A, Aro EM, Vener AV. 2011. Thylakoid phosphoproteins: identification of phosphorylation sites. In: Carpentier R (ed.) Photosynthesis Research Protocols. Methods in Molecular Biology 684:171-86. Humana Press, NY, Springer.  ISBN 978-1-60761-924-6
Sharon I, Battchikova N, Aro EM, Giglione C, Meinnel T, Glaser F, Pinter RY, Breitbart M, Rohwer F, Béjà O. 2011. Comparative metagenomics of microbial traits within oceanic viral communities. Isme J. 5: 1178-1190
Sirpiö S, Suorsa M, Aro EM. 2011. Analysis of thylakoid protein complexes by two dimensional electrophoretic systems. In: Jarvis RP. (ed.) Chloroplast Research in Arabidopsis, Volume II: Methods and Protocols. Methods in Molecular Biology 775: 19-30.  Humana Press, NY, USA. ISBN 978-1-61779-236-6
Tikkanen M, Grieco M, Aro EM. 2011. Novel insights into light-harvesting complex II phosphorylation and "state-transitions". Trends Plant Sci. 16: 126-31
Trotta A, Konert G, Rahikainen M, Aro EM, Kangasjärvi S. 2011.  Knock-down of protein phosphatase 2A subunit B'γ promotes phosphorylation of CALRETICULIN 1 in Arabidopsis thaliana. Plant Signal Behav. 6: 1665-1668
Trotta A, Wrzaczek M, Scharte J, Tikkanen M, Konert G, Rahikainen M, Holmström M, Hiltunen HM, Rips S, Sipari N, Mulo P, Weis E, von Schaewen A, Aro EM, Kangasjärvi S. 2011. Regulatory subunit B’γ of protein phosphatase 2A prevents unnecessary defense reactions under low light in Arabidopsis thaliana. Plant Physiol. 156: 1464-1480


Allahverdiyeva Y, Leino H, Saari L, Fewer DP, Shunmugam S, Sivonen K, Aro EM. 2010. Screening for biohydrogen production by cyanobacteria isolated from the Baltic Sea and Finnish lakes. Int J Hydrog Energy. 35: 1111-1127
Battchikova N, Vainonen J, Vorontsova N, Keränen M, Carmel D, Aro EM. 2010. Dynamic changes in the proteome of Synechocystis 6803 in response to CO2 limitation revealed by quantitative proteomics. J Proteome Res. 9: 5896-5912
Birungi M, Folea M, Battchikova N, Xu M, Mi H, Ogawa T, Aro EM, Boekema EJ. 2010. Possibilities of subunit localization with fluorescent protein tags and electron microscopy examplified by a cyanobacterial NDH-1 study. Biochim Biophys Acta – Bioenerg. 1797: 1681-1686
Chen KM, Holmström M, Raksajit W, Suorsa M, Piippo M, Aro EM. 2010.  Small chloroplast-targeted DnaJ proteins are involved in optimization of photosynthetic reactions in Arabidopsis thaliana. BMC Plant Biol. 10: 43
Kubota H, Sakurai I, Katayama K, Mizusawa N, Ohashi S, Kobayashi M, Zhang P, Aro EM, Wada H. 2010. Purification and characterization of photosystem I complex from Synechocystis sp. PCC 6803 by expressing histidine-tagged subunits. Biochim Biophys Acta – Bioenerg. 1797: 98-105
Lehtimäki N, Lintala M, Allahverdiyeva Y, Aro EM, Mulo P. 2010. Drought stress -induced upregulation of components involved in ferredoxin-dependent cyclic electron transfer. J Plant Physiol 167: 1018-22
Suorsa M, Sirpiö S, Paakkarinen V, Kumari N, Holmström M, Aro EM. 2010. Two proteins homologous to PsbQ are novel subunits of the chloroplast NAD(P)H dehydrogenase. Plant Cell Physiol. 51: 877-83
Tikkanen M, Grieco M, Kangasjärvi S, Aro EM. 2010. Thylakoid protein phosphorylation in higher plant chloroplasts optimises electron transfer under fluctuating light. Plant Physiol. 152: 723-735
Yin L, Lundin B, Bertrand M, Nurmi M, Solymosi K, Kangasjärvi S, Aro EM, Schoefs B, Spetea C. 2010.  Role of thylakoid ATP/ADP carrier in photoinhibition and photoprotection of photosystem II in Arabidopsis. Plant Physiol. 153: 666-77


Allahverdiyeva Y, Mamedov F, Holmström M, Nurmi M, Lundin B, Styring S, Spetea C, Aro EM. 2009.  Comparison of the electron transport properties of the psbo1 and psbo2 mutants of Arabidopsis thaliana. Biochim Biophys Acta – Bioenerg. 1787: 1230-1237
Boehm M, Nield J, Zhang P, Aro EM, Komenda J, Nixon PJ. 2009. Structural and mutational analysis of band 7 proteins in the cyanobacterium Synechocystis sp. strain PCC 6803. J Bacteriol. 1797(5): 566-75
Cardona T, Battchikova N, Zhang PP, Stensjö K, Aro EM, Lindblad P, Magnuson A. 2009. Electron transfer protein complexes in the thylakoid membranes of heterocysts from the cyanobacterium Nostoc punctiforme. Biochim Biophys Acta – Bioenerg. 1787: 252-263
Fristedt R, Carlberg I, Zygadlo A, Piippo M, Nurmi M, Aro EM, Scheller HV, Vener A. 2009. Intrinsically unstructured phosphoprotein TSP9 regulates light harvesting in Arabidopsis thaliana. Biochemistry 48: 499-509
Kangasjärvi S, Nurmi M, Tikkanen M, Aro EM. 2009. Cell-specific mechanisms and systemic signalling as emerging themes in light acclimation of C3 plants. Plant Cell Environ. 32: 1230-1240
Keränen M, Aro EM, Nevalainen O, Tyystjärvi E. 2009. Toxic and non-toxic Nodularia strains can be distinguished from each other and from eukaryotic algae with chlorophyll fluorescence fingerprinting. Harmful Algae 8: 817-822
Lintala M, Allahverdiyeva Y, Kangasjärvi S, Lehtimäki N, Keränen M, Rintamäki E, Aro EM, Mulo P. 2009. Comparative analysis of leaf-type ferredoxin-NADP+-oxidoreductase isoforms in Arabidopsis thaliana. Plant J. 57: 1103-1115
Mulo P, Sicora C, Aro EM. 2009. Cyanobacterial psbA gene family: optimization of oxygenic photosynthesis. Cell Mol Life Sci. 66: 3697-710
Pantaleoni L, Ferroni L, Baldisserotto C, Aro EM, Pancaldi S. 2009.  Photosystem II organisation in chloroplasts of Arum italicum leaf depends on tissue location. Planta 230: 1019-1031
Sicora CI, Ho F, Salminen T, Styring S, Aro EM. 2009. Transcription of a "silent" cyanobacterial psbA gene is induced by microaerobic conditions. Biochim Biophys Acta – Bioenerg. 1787: 105-112
Sirpiö S, Allahverdiyeva Y, Holmström M, Khrouchtchova A, Haldrup A, Battchikova N, Aro EM. 2009.  Novel nuclear-encoded subunits of the chloroplast NAD(P)H dehydrogenase complex. J Biol Chem. 284: 905-912
Sirpiö S, Holmström M, Battchikova N, Aro EM. 2009. AtCYP20-2 is an auxiliary protein of the chloroplast NAD(P)H dehydrogenase complex. FEBS Lett. 583: 2355-2358
Suorsa M, Sirpiö S, Aro EM. 2009. Towards Characterization of the Chloroplast NAD(P)H Dehydrogenase Complex. Mol Plant. 2: 1127-1140
Tikkanen M, Suorsa M, Aro EM. 2009. The flow of solar energy to biofuel feedstock via photosynthesis. Int Sugar J. 111:156-163
Vainonen JP, Vener AV, Aro EM. 2009. Determination of in vivo Protein Phosphorylation in Photosynthetic Membranes. In: Pflannschmidt T. (ed.) Plant signal transduction - Methods in Molecular Biology 479: 133-146, Humana Press, NY.
Zhang P, Allahverdiyeva Y, Eisenhut M, Aro EM. 2009.  Flavodiiron proteins in oxygenic photosynthetic organisms: Photoprotection of photosystem II by Flv2 and Flv4 in Synechocystis sp. PCC 6803. PLoS ONE. 2009;4(4):e5331


Allahverdiyeva Y, Sairanen I, Stensjö K, Lindblad P, Aro EM. 2008. Photosynthetic electron transport properties in Nostoc punctiforme PCC 73102. In: Allen JF., Gantt E., Goldbeck JH. and Osmond B. (eds.) Photosynthesis. Energy from the sun. 14th International Gongress of Photosynthesis. Springer, Heidelberg, pp. 3-5, ISBN: 978-1-4020-6707-5
Folea IM, Zhang P, Nowaczyk MM, Ogawa T, Aro EM, Boekema EJ. 2008. Single particle analysis of thylakoid proteins from Thermosynechococcus elongatus and Synechocystis 6803: Localization of the CupA subunit of NDH-1. FEBS Lett. 582: 249-254
Folea IM, Zhang P, Nowaczyk MM, Ogawa T, Aro EM, Boekema EJ. 2008.  Domain organization of photosystem II in membranes of the cyanobacterium Synechocystis PCC6803 investigated by electron microscopy. FEBS Lett. 582:1749-54
Kanervo E, Singh M, Suorsa M, Paakkarinen V, Aro E, Battchikova N, Aro EM. 2008. Expression of protein complexes and individual proteins upon transition of etioplasts to chloroplasts in pea (Pisum sativum). Plant Cell Physiol. 49: 396-410
Kangasjärvi S, Lepistö A, Hänninen K, Aro EM, Rintamäki E. 2008. Diverse roles for chloroplast stromal and thylakoid-bound ascorbate peroxidases in plant stress responses. Biochem J. 412: 275-85
Lintala M, Allahverdiyeva Y, Kangasjärvi S, Lehtimäki N, Keränen M, Rintamäki E, Aro EM, Mulo P. 2008. Comparative analysis of leaf-type ferredoxin-NADP+-oxidoreductase isoforms in Arabidopsis thaliana. Plant J. 57: 1103-1115
Lintala M, Allahverdiyeva Y, Lehtimäki N, Aro EM, Mulo P. (2008) Structural and Functional Characterization of Leaf-Type Ferredoxin-NADP+-Oxidoreductase Isoforms in Arabidopsis thaliana. In: Allen JF., Gantt E., Golbeck JH., Osmond B. (eds.) Photosynthesis. Energy from the Sun. 14th International Gongress of Photosynthesis. Springer, Dordrecht, pp 933-936. ISBN: 978-1-4020-6707-5
Lundin B, Nurmi M, Rojas-Stuetz M, Aro EM, Adamska I, Spetea C. 2008. Towards understanding the functional difference between the two PsbO isoforms in Arabidopsis thaliana-insights from phenotypic analyses of psbo knockout mutants. Photos Res. 98: 405-14
Mulo P, Sirpiö S, Suorsa M, Aro EM. 2008. Auxiliary proteins involved in the assembly and sustenance of photosystem II. Photos Res. 98: 489-501
Sicora CI, Aro EM. 2008. Differential Expression of hoxY Gene, Encoding the Small Subunit of Bidirectional Hydrogenase, Under Ar-Induced Microaerobic Conditions in Synechocystis sp. PCC6803 and Anabaena sp. PCC7120. In: Allen JF., Gantt E., Goldbeck JH. and Osmond B. (eds.) Photosynthesis. Energy from the sun. 14th International Gongress of Photosynthesis. Springer, Heidelberg, ISBN: 978-1-4020-6707-5, pp. 19-22. ISBN: 978-1-4020-6707-5
Singh M, Satoh K, Yamamoto Y, Kanervo E, Aro EM. 2008. In vivo quality control of photosystem II in cyanobacteria Synechocystis sp. PCC 6803: D1 protein degradation and repair under the influence of light, heat and darkness. Indian J Biochem Biophys. 45:237-243
Sirpiö S, Khrouchtchova A, Allahverdiyeva Y, Hansson M, Fristedt R, Vener A, Scheller HV, Jensen P, Haldrup A, Aro EM. 2008. AtCYP38 ensures early biogenesis, correct assembly and sustenance of photosystem II. Plant J. 55: 639-51
Soitamo A, Piippo M, Allahverdiyeva Y, Battchikova N, Aro EM. 2008. Light has a specific role in modulating Arabidopsis gene expression at low temperature. BMC Plant Biology. 8: 13
Suorsa M, Tikkanen M, Aro EM. 2008. Mikrobien valjastaminen energiantuottoon -fotosynteesitutkimuksen uusia ulottuvuuksia. Luonnontutkija 4: 112-119
Tikkanen M, Nurmi MJ, Kangasjärvi S, Aro EM. 2008. Core protein phosphorylation facilitates the repair of photodamaged Photosystem II at High Light. Biochim Biophys Acta – Bioenerg. 1777: 1432-1437
Tikkanen M, Nurmi M, Suorsa M, Danielsson R, Mamedov F, Styring S, Aro EM. 2008. Phosphorylation-dependent regulation of excitation energy distribution between the two photosystems in higher plants. Biochim Biophys Acta – Bioenerg. 1777: 425-432
Vainonen, J., *Sakuragi, Y., *Stael, S., Tikkanen, M., Allahverdiyeva, Y., Paakkarinen, V., Aro, E., Suorsa, M., *Scheller, HV., *Vener, A., Aro, EM. 2008: Light regulation of CaS, a novel phosphoprotein in the thylakoid membrane of Arabidopsis thaliana. - FEBS Journal . 275: 1767?1777. ( doi:10.1111/j.1742-4658.2008.06335.x)
Vass I, Aro EM. 2008. Photoinhibition of Photosystem II electron transport. In: Regner G. (ed.) Primary Processes of Photosynthesis: Basic Principles and Apparatus, Comprehensive Series in Photochemical and Photobiological Sciences. Publ Roy Soc Chem, Cambridge. Volume 8 pp. 393-411 ISBN 978-0-85404-364-9 
Zhang PP, Sicora CI, Vorontsova N, Allahverdiyeva Y, Battchikova N, Nixon PJ, Aro EM. 2008. Expression of inducible inorganic carbon acquisition complexes is under the control of the FtsH protease in Synechocystis sp. PCC 6803 In: Allen JF., Gantt E., Goldbeck JH. and Osmond B. (eds.) Photosynthesis. Energy from the sun. 14th International Gongress of Photosynthesis. Springer, Heidelberg, ISBN: 978-1-4020-6707-5, pp. 829-833


Allahverdiyeva Y, Mamedov F, Suorsa M, Styring S, Vass I, Aro EM. 2007.  Insights into the function of PsbR protein in Arabidopsis thaliana. Biochim Biophys Acta – Bioenerg. 1767: 677-685
Battchikova N, Aro EM. 2007.  Cyanobacterial NDH-1 complexes: multiplicity in function and subunit composition. Physiol Plant. 131: 22-32
Bishop CL, Ulas S, Baena-Gonzalez E, Aro EM, Purton S, Nugent JHA, Mäenpää P. 2007. The PsbZ subunit of Photosystem II in Synechocystis sp. PCC 6803 modulates electron flow through the photosynthetic electron transfer chain. Photos Res. 93: 139-147
Cardona T, Battchikova N, Agervald Å, Zhang P, Nagel E, Styring S, Aro EM, Lindblad P, Magnusson A. 2007. Isolation and characterization of thylakoid membranes from the filamentous cyanobacterium Nostoc punctiforme. Physiol Plant. 131: 622-34
Hoffman A, Milde S, Desel C, Hümpel A, Kaiser H, Hammes E, Piippo M, Soitamo A, Aro EM, Gerendas J, Sattelmacher B, Hansen UP. 2007. N form-dependent growth retardation of Arabidopsis thaliana seedlings as revealed from physiological and microrray studies. J Plant Nutr Soil Sci. 170:1-11
Kanervo E, Suorsa M, Aro EM. 2007. Assembly of protein complexes in plastids. In: Bock R (ed.). Topics in Current Genetics: Cell and Molecular Biology of Plastids. SpringerLink, Vol. 19, pp. 283-313, ISBN: 978-3-540-75375-9
Lintala M, Allahverdiyeva Y, Kidron H, Piippo M, Battchikova N, Suorsa M, Rintamäki E, Salminen TA, Aro EM, Mulo P. 2007. Structural and functional characterization of ferredoxin-NADP+-oxidoreductase using knock-out mutants of Arabidopsis. Plant J. 49: 1041-52
Sirpiö S, Allahverdiyeva Y, Suorsa M, Paakkarinen V, Vainonen J, Battchikova N, Aro EM. 2007. TLP18.3, a novel thylakoid lumen protein regulating Photosystem II repair cycle. Biochem J. 406: 415-425
Suorsa M, Aro EM. 2007.  Expression, assembly and auxiliary functions of photosystem II oxygen-evolving proteins in higher plants. Photos Res 93: 89-100
Zhang P, Sicora CI, Vorontsova N, Allahverdiyeva Y, Battchikova N, Nixon PJ, Aro EM. 2007. FtsH protease is required for induction of inorganic carbon acquisition complexes in Synechocystis sp. PCC 6803. Mol Microbiol. 65: 728-740


Andaluz S, Lopez-Millan AF, De Las Rivas J, Aro EM, Abadia J, Abadia A. 2006. Proteomic profiles of thylakoid membranes and changes in response to iron deficiency. - Photos Res 89: 141-155
Arteni AA, Zhang P, Battchikova N, Ogawa T, Aro EM, Boekema,E. 2006. Structural characterisation of NDH-1 complexes of Thermosynechococcus elongatus by single particle electron microscopy. Biochim Biophys Acta – Bioenerg. 1757: 1469-1475
Danielsson R, Suorsa M, Paakkarinen V, Albertsson PÅ, Styring S, Aro EM, Mamedov F. 2006. Dimeric and monomeric organization of photosystem II: Distribution of five distinct complexes in the different domains of the thylakoid membrane. J Biol Chem 281: 14241-14249
Piippo M, Allahverdiyeva Y, Paakkarinen V, Suoranta UM, Battchikova N, Aro EM. 2006. Chloroplast-mediated regulation of nuclear genes in Arabidopsis thaliana in the absence of light stress. Physiol Genomics. 25: 142-152
Srivastava R, Battchikova N, Norling B, Aro EM. 2006. Plasma membrane of Synechocystis PCC 6803: A heterogeneous distribution of membrane proteins. Arch Microbiol. 185: 238-243
Suorsa M, Sirpiö S, Allahverdiyeva Y, Paakkarinen V, Mamedov F, Styring S, Aro EM. 2006. PsbR – a missing link in the assembly of the oxygen evolving complex of plant photosystem II. J Biol Chem. 281: 145-150
Tikkanen M, Piippo M, Suorsa M, Sirpiö S, Mulo P, Vainonen J, Vener AV, Allahverdiyeva Y, Aro EM. 2006. State transitions revisited - a buffering system for dynamic low light acclimation of Arabidopsis. Plant Mol Biol. 62: 779-793


Allahverdiyeva Y, Mamedov F, Mäenpää P, Vass I, Aro EM. 2005. Modulation of photosynthetic electron transport in the absence of terminal electron acceptors. Characterisation of the rbcL deletion mutant of tobacco. Biochim Biophys Acta – Bioenerg. 1709: 69-83
Aro EM, Suorsa M, Rokka A, Allahverdiyeva Y, Paakkarinen V, Saleem A, Battchikova N, Rintamäki E. 2005. Dynamics of Photosystem II – a proteomic approach to thylakoid protein complexes. J Exp Bot. 56: 347-356
Battchikova N, Zhang PP, Rudd S, Ogawa T, Aro EM. 2005. Identification of NdhL and Ssl1690 (NdhO) in NDH-1L and NDH-1M complexes of Synechocystis sp. PCC 6803. J Biol Chem. 280: 2587-2595
Chow WS, Aro EM. 2005. The photoinactivation of photosystem II and mechanisms of recovery. In: Wydrzynski T. and Satoh K. (eds.). Photosystem II – The light-driven water plastoquinone oxidoreductase. Advances in photosynthesis and respiration, chapter 68, pp. 628-643, ISBN: 1-4020-4249-3
Herranen M, Tyystjärvi T, Aro EM. 2005. Regulation of Photosystem I Reaction Center Genes in Synechocystis sp. Strain PCC6803 during Light Acclimation. Plant Cell Physiol. 46: 1484-1493
Kanervo E, Suorsa M, Aro EM. 2005. Functional flexibility and acclimation of the thylakoid membrane.  Photochem Photobiol Sci. 4: 1072-1080
Khrouchtchova A, Hansson M, Paakkarinen V, Vainonen J, Janssen PE, Scheller HV, Aro EM, Haldrup A. 2005. A previously found thylakoid membrane protein of 14 kDa (TMP14) is a novel subunit of plant photosystem I and is designed as PSI-P. FEBS Lett. 579: 4808-4812
Rokka A, Suorsa M, Saleem A, Battchikova N, Aro EM. 2005. Synthesis and assembly of thylakoid protein complexes. Multiple assembly intermediates of photosystem II. Biochem J. 388: 159-168
Singh M, Yamamoto Y, Satoh K, Aro EM, Kanervo E. 2005. Post-illumination-related loss of photochemical efficiency of photosystem II and degradation of the D1 protein are temperature-dependent. J Plant Physiol. 162: 1246-1253
Zhang PP, Battchikova N, Paakkarinen V, Katoh H, Iwai M, Ikeuchi M, Pakrasi HB, Ogawa T, Aro EM. 2005. Isolation, subunit composition and interaction of the NDH-1 complexes from Thermosynechococcus elongatus BP-1. Biochem J. 390: 513-520


Aro EM, Rokka A, Vener AV. 2004. Determination of phosphoproteins in higher plant thylakoids. In: Carpentier R. (ed.) Photosynthesis Research Protocols. Methods Mol Biol. 274: 271-285. Humana Press, NY, ISBN: 1-58829-232-0, 1-59259-799-8
Herranen M, Battchikova N, Zhang P, Graf A, Sirpiö S, Paakkarinen V, Aro EM. 2004. Towards functional proteomics of membrane protein complexes in Synechocystis sp. PCC 6803. Plant Physiol. 134: 470-481
Kanervo E, Aro EM. 2004. Photosystem II assembly and turnover of the D1 protein. In: Lennarz W. and Lane MD. (eds.) Encyclopedia of Biological Chemistry, Vol 3, Elsevier, Oxford, ISBN: 0-12-443710-9, pp. 363-366
Suorsa M, Regel RE, Paakkarinen V, Battchikova N, Herrmann RG, Aro EM. 2004. Protein assembly of photosystem II and accumulation of subcomplexes in the absence of low molecular weight subunits PsbL and PsbJ. Eur J Biochem. 271: 96-107
Tyystjärvi T, Sirpiö S, Aro EM. 2004. Post-translational regulation of the psbA gene family in the cyanobacterium Synechocystis sp. PCC 7942. FEBS Lett 576: 211-215
Zhang PP, Battchikova N, Jansen T, Appel J, Ogawa T, Aro EM. 2004. Expression and functional roles of the two distinct NDH-1 complexes and the carbon acquisition complex NdhD3/NdhF3/CupA/Sll1735 in Synechocystis sp. PCC 6803. Plant Cell 16: 3326-3340


Aro EM, Ohad I. 2003. Redox regulation of thylakoid protein phosphorylation. Antioxid Redox Signal. 5: 55-67
Baena-Gonzalez E, Allahverdhieva Y, Svab Z, Maliga P, Josse EM, Kuntz M, Mäenpää P, Aro EM. 2003. Deletion of the tobacco plastid psbA gene triggers an up-regulation of the thylakoid-associated NAD(P)H complex and the plastid terminal oxidase (PTOX). Plant J. 35: 704-716
Hiriart JB, Aro EM, Lehto K. 2003. Dynamics of the VIGS-mediated chimeric silencing of the Nicotiana benthamiana ChlH gene and of the tobacco mosaic virus. Mol Plant Microbe Int. 16: 99-106
Hou XC, Rintamäki E, Aro EM. 2003. Ascorbate-mediated LHCII protein phosphorylation - LHCII kinase regulation in light and in darkness. Biochemistry 42: 5828-5836
Kanervo E, Spetea C, Nishiyama Y, Murata N, Andersson B, Aro EM. 2003. Dissecting a cyanobacterial proteolytic system: Efficiency in inducing degradation of the D1 protein of photosystem II in cyanobacteria and plants. Biochim Biophys Acta – Bioenerg. 1607: 131-140
Keränen M, Aro EM, Tyystjärvi E. 2003. Automatic plant identification with chlorophyll fluorescence fingerprinting. Precis Agric. 4: 53-67
Lehto K, Tikkanen M, Hiriart JB, Paakkarinen V, Aro EM. 2003. Depletion of the photosystem II complex in mature tobacco leaves infected by the Flavum strain of Tobacco mosaic virus. Mol Plant Microbe Int. 16: 1135-1144
Martinsuo P, Pursiheimo S, Aro EM, Rintamäki E. 2003. Dithiol oxidant and disulfide reductant dynamically regulate the phosphorylation of light-harvesting complex II proteins in thylakoid membranes. Plant Physiol. 133: 37-46
Mulo P, Pursiheimo S, Hou CX, Tyystjärvi T, Aro EM. 2003. Multiple effects of antibiotics on chloroplast and nuclear gene expression. Funct Plant Biol. 30: 1097-1103
Pursiheimo S, Martinsuo P, Rintamäki E, Aro EM. 2003. Photosystem II protein phosphorylation follows four different regulatory patterns induced by environmental cues. Plant Cell Environ. 26: 1995-2003
Sakurai I, Hagio M, Gombos Z, Tyystjärvi T, Paakkarinen V, Aro EM, Wada H. 2003. Requirement of phosphatidylglycerol for maintenance of photosynthetic machinery. Plant Physiol. 133: 1376-1384


Baena-Gonzalez E, Aro EM. 2002. Biogenesis, assembly and turnover of photosystem II units. Philos Trans Roy Soc B. 357: 1439-1440
Bergo E, Pursiheimo S, Paakkarinen V, Giagometti GM, Donella-Deana A, Andreucci F, Barbato R, Aro EM. 2002. Rapid and highly specific monitoring of reversible thylakoid protein phosphorylation by polyclonal antibody to phosphothreonine-containing proteins. J Plant Physiol. 159: 371-377
Hiriart JB, Lehto K, Tyystjärvi E, Junttila T, Aro EM. 2002. Suppression of a key enzyme involved in chlorophyll biosynthesis using virus-inducing gene silencing. Plant Mol Biol. 50: 213-224
Hou CX, Rintamäki E, Aro EM. 2002. Environmental and metabolic control of PHCII protein phosphorylation: revealing the mechanisms for dual regulation of the LHCII kinase. Plant Cell Environ. 25: 1515-1525
Jansen T, Kanervo E, Aro EM, Mäenpää P. 2002. Localisation and processing of the precursor form of photosystem II protein D1 in Synechocystis 6803. J Plant Physiol. 159: 1205-1211
Mamedov F, Rintamäki E, Aro EM, Andersson B, Styring S. 2002. Influence of protein phosphorylation on the electron-transfer properties of Photosystem II. Photos Res. 74: 61-72
Pätsikkä E, Kairavuo M, Sersen F, Aro EM, Tyystjärvi E. 2002. Excess copper predisposes photosystem II to photoinhibition in vivo by outcompeting iron and causing decrease in leaf chlorophyll. Plant Physiol. 129: 1359-1367
Tyystjärvi T, Tuominen I, Herranen M, Aro EM, Tyystjärvi E. 2002. Action spectrum of psbA gene transcription is similar to that of photoinhibition in Synechocystis sp. PCC 6803. FEBS Lett. 516: 167-171
Zhang L, Aro EM. 2002. Synthesis, membrane insertion and assembly of the chloroplast-encoded D1 protein into Photosystem II. FEBS Lett. 512: 13-18


Aro EM, Andersson B. 2001. Photodamage and D1 protein turnover in photosystem II. In: Aro EM. and Andresson B (eds.) Regulation on Photosynthesis. Kluwer Academic Publishers, Dortrecht, Netherlands. pp. 377-393
Aro EM, Zhang L, Suorsa M, Paakkarinen V, Battchikova N. 2001. Maintenance of functional Photosystem II by D1 protein turnover. In PS2001 Proceedings: 12th International Congress on Photosynthesis. CSIRO Publishing, Melbourne, Australia. CD-rom.
Baena-Gonzalez E, Baginsky S, Mulo P, Summer H, Aro EM, Link G. 2001. Chloroplast transcription at different light intensities: Glutathione-mediated phosphorylation of the major RNA polymerase involved in redox-regulated organellar gene expression. Plant Physiol. 127: 1044-1052​1104/​pp.​010168
Baena-Gonzalez E, Gray JC, Tyystjärvi E, Aro EM, Mäenpää P. 2001. Abnormal regulation of photosynthetic electron transport in a chloroplast ycf9 inactivation mutant. J Biol Chem. 276: 20795-20802
Baena-Gonzalez E, Gray JC, Tyystjärvi E, Battchikova N, Mäenpää P, Aro EM. 2001. The ORF62 protein of the thylakoid membrane is involved in regulation of photosynthetic electron transport. PS2001 Proceedings: 12th International Congress of Photosynthesis. CSIRO Publishing, Australia. CD-rom
Herranen M, Aro EM, Tyystjärvi T. 2001. Two distinct mechanisms regulate the transcription of photosystem II genes in Synechocystis sp. PCC 6803. Physiol Plant. 112: 531-539
Herranen M, Tyystjärvi T, Aro EM. 2001. Light-regulation of photosystem I reaction centre genes in the cyanobacterium Synechocystis sp. PCC 6803. PS2001 Proceedings: 12th International Congress of Photosynthesis. CSIRO Publishing, Australia. CD-rom
Pursiheimo S, Mulo P, Rintamäki E, Aro EM. 2001. Coregulation of light-harvesting complex II phosphorylation and lhcb mRNA accumulation in winter rye. Plant J. 26: 317-327
Pätsikkä E, Aro EM, Tyystjärvi E. 2001. Mechanism of copper-induced photoinhibition in thylakoid membranes. Physiol Plant. 113: 142-150

Rintamäki E, Aro EM. 2001. Phosphorylation of photosystem II proteins. In: Aro EM. and Andersson B. (eds.). Regulation of Photosynthesis - Advances in Photosynthesis Vol. 11. Kluwer Academic Publishers, Dordrecht. pp. 395-418
Rokka A, Zhang L, Aro EM. 2001. Rubisco activase, an enzyme with a temperature-dependent dual function. Plant J. 25: 463-471
Suorsa M, Zhang L, Battchikova N, Paakkarinen V, Regel E, Herrmann R, Aro EM. 2001. Assembly of photosystem II in chloroplast psbEFLJ operon mutants. PS2001 Proceedings: 12th International Congress of Photosynthesis. CSIRO Publishing, Australia. CD-rom
Tyystjärvi T, Herranen M, Aro EM. 2001. Regulation of translation elongation in cyanobacteria: membrane targeting of the ribosome nascent-chain complexes controls the synthesis of D1 protein. Mol Microbiol. 40: 467-484
Yang DH, Andersson B, Aro EM, Ohad I. 2001. The redox state of the plastoquinone pool controls the level of the light-harvesting chlorophyll a/b binding protein complex II (LHCII) during photoacclimation. Photos Res. 68: 163-174
Zhang L, Paakkarinen V, Suorsa M, Aro EM. 2001. A SecY homologue is involved in chloroplast-encoded D1 protein biogenesis. J Biol Chem. 276: 37809-37814


Mäenpää P, Baena-Gonzalez E, Chen L, Khan MS, Gray JC, Aro EM. 2000. The ycf 9 [ofr 62] gene in the plant chloroplast genome encodes a hydrophobic protein of stromal thylakoid membranes. J Exp Bot. 51: 375-382
Rintamäki E, Martinsuo P, Pursiheimo S, Aro EM. 2000. Cooperative regulation of light-harvesting complex II phosphorylation via the plastoquinol and ferredoxin-thioredoxin system in chloroplasts. Proc Natl Acad Sci USA. 10: 11644-11649
Rokka A, Aro EM, Herrmann RG, Andersson B, Vener A. 2000. Dephosphorylation of photosystem II reaction center proteins in plant photosynthetic membranes as an immediate response to abrupt elevation of temperature. Plant Physiol. 123: 1525-1535
Sippola K, Aro EM. 2000. Expression of psbA genes is regulated at multiple levels in the cyanobacterium Synechococcus sp. PCC 7942. Photochem Photobiol. 71: 706-714
Zhang L, Paakkarinen V, van Wijk KJ, Aro EM. 2000. Biogenesis of the chloroplast-encoded D1 protein: Regulation of translation elongation, insertion and assembly into photosystem II. Plant Cell 12: 1769-1782

1999 and earlier publications

Around 100, please contact Prof. Aro for complete list