List of publications


International (refereed) journals





91. Croce R. and van Amerongen H. (2014) "Natural strategies for photosynthetic  light harvesting" Nat. Chem. Biol. , 10, 492-501.


90. Ünlü C., Drop B., Croce R. and van Amerongen H. (2014) “State transitions in Chlamydomonas reinhardtii strongly modulate the size of Photosystem II but not of Photosystem I” Proc.Nat.Am.Sci. USA, 111(9):3460-3465.


89. Drop B., Yadav S., Boekema E.J. and Croce R. (2014) “Consequences of state transitions on the structural and functional organization of Photosystem I in the green alga Chlamydomonas reinhardtiiPlant Journal, 78, 181-191


88. Passarini F., Xu P., Caffarri S., Hille J. and Croce R. (2014) “Towards in vivo mutation analysis: Knock-out of specific chlorophylls bound to the light-harvesting complexes of Arabidopsis thaliana - The case of CP24 (Lhcb6)” BBA-Bioenergetics – 1837, 1500-1506


87. Dinc E., Ramundo S., Croce R. and Rochaix, JD (2014) "Repressible chloroplast gene expression in Chlamydomonas: A new tool for the study of the photosynthetic apparatus" BBA – bioenergetics , 1837, 1548-1552


86. Drop B., Webber-Birungi M, Yadav S., Filipowicz-Szymanska A., Fusetti F., Boekema E.J. and Croce R. (2014) “Light-harvesting complex II (LHCII) and its supramolecular organization in Chlamydomonas reinhardtiiBBA – bioenergetics,  1837 63–72





85.    Liguori N., Roy L.M., Opacic M., Durand G. and Croce R. (2013) “Regulation of light harvesting in the green alga Chlamydomonas reinhardtii: the C-terminus of LHCSR is the knob of a dimmer switch” J.Am.Chem.Soc  , 135 (49)  18339–18342


83.    Wientjes E., Drop B., Kouril R., Boekema E.J. and Croce R. (2013) “Photosystem II supercomplex organization does not change during state transitions in higher plant” J. Biol. Chem.  288, 32821-32826.


84.    Croce R. and van Amerongen H (2013) “Light-harvesting in Photosystem I”, Photosynt. Res., 116:153–166.


81.    van Amerongen H and Croce R (2013) “Light harvesting in Photosystem II” Photosynt. Res., 116:251–263.


80.    Wientjes E., van Amerongen H. and Croce R. (2013) “Quantum yield of charge separation in Photosystem II: functional effect of changes in the antenna size upon light acclimation” (2013) J. Phys. Chem B. 117, 11200−11208.


79.    Wientjes E., van Amerongen H. and Croce R. (2013) “LHCII is an antenna of both photosystems after long term acclimation” BBA- bioenergetics, 1827,  420–426


78.    Kouril R., Wientjes E., Bultema J.B, Croce R. and Boekema E.J. (2013) “High-light vs. low-light: Effect of light acclimation on photosystem II composition and organization in Arabidopsis thaliana” BBA- bioenergetics, 1827,  411–419





77.    Hogewoning SW, Wientjes E., Douwstra P., Trouwborst G., van Iperen W., Croce R.  and Harbinson J. (2012) “Photosynthetic Quantum Yield Dynamics: from Photosystems to Leaves” Plant Cell, 24: 1921–1935.


76.    Wientjes E. and Croce R. (2012) “PMS: Photosystem I electron donor or fluorescence quencher” Photosyn. Res., 111, 185-191


75.    Wientjes E., Roest G. and Croce R. (2012) “From blue to red to far-red in Lhca4: how does the protein modulate the spectral properties of the pigments?” BBA- bioenergetics 1817, 711-717





74.    Drop B., Webber-Birungi M., Fusetti F., Kouřil R., Redding KE, Boekema EJ and Croce R. (2011) “Photosystem I of Chlamydomonas reinhardtii is composed of nine Light-harvesting complexes (Lhca) located on one side of the core”. J. Biol. Chem.  286: 44878-44887.


73.    Krüger T.P.J, Wientjes E., Croce R. and van Grondelle R. (2011) “Conformational switching explains the intrinsic multifunctionality of plant light-harvesting complexesPNAS, 108, 1351613521.


72.    Wientjes E., van Stokkum IHM, van Amerongen H. and Croce R. (2011) “The role of the individual Lhca’s in Photosystem I excitation energy trapping” Biophys. J.101, 745-754.


71.    Marin A., Passarini F., van Stokkum IHM, van Grondelle R. and Croce R. (2011). “Minor complexes at work: light-harvesting by Carotenoids in the Photosystem II antenna complexes CP24 and CP26” Biophys. J. 100, 2829–2838


70.    Croce R. and van Amerongen H. (2011). “Light-harvesting and structural organization of Photosystem II: from individual complexes to thylakoid membrane” . J. Photochem.Photobio. B  Biology  104, 142–153


69.    Caffarri S., Broess K, Croce R. and van Amerongen H. (2011) “Excitation energy transfer and trapping in higher plant Photosystem II complexes with different antenna sizes”. Biophys. J. 100, 2094–2103.


79.    Wientjes E., van Stokkum IHM, van Amerongen H. and Croce R. (2011) ”Excitation energy transfer dynamics of higher plant Photosystem I light harvesting complexes”. Biophys. J., 100, 1372–1380.


67.    Wientjes E. and Croce R. (2011) “The light-harvesting complexes of higher plant Photosystem I: Lhca1/4 and Lhca2/3 form two red-emitting heterodimers” Biochemical Journal 433, 477-485.





66.    Marin A., Passarini F., Croce R. and van Grondelle R. (2010)” The energy transfer pathways in the CP24 and CP26 antenna complexes of higher plant Photosystem II. A comparative study." Biophys. J. ,99, 4056-4065. 


65.    Passarini F., Wientjes E., van Amerongen H. and Croce R. (2010) “Photosystem I light-harvesting complex Lhca4 adopts multiple conformations: red forms and excited-state quenching are mutually exclusive”. Biochim.Biophys.Acta - Bioenergetics. 1797(4):501-508.


64.    Müller M.G., Lambrev P., Reus M., Wientjes E., Croce R., and Holzwarth A.R. (2010) “Singlet energy dissipation in Photosystem II light-harvesting complex does not involve energy transfer to carotenoids”. ChemPhysChem 11, 1289 – 1296.


63.    Van Oort B., Alberts M., de Bianchi S., Dall’Osto L., Bassi R., Trinkunas G., Croce R. and van Amerongen H (2010) “Effect of antenna-depletion in Photosystem II on excitation energy transfer in Arabidopsis thaliana”. Biophys. J.  98(5), 922-31.


62.    Mozzo M., Mantelli M., Passarini F., Caffarri S., Croce R. and Bassi R. (2010) “Functional analysis of phoyosystem I light-harvesting complexes (Lhca) gene products of Chlamydomonas reinhardtii”. ”. Biochim.Biophys.Acta - Bioenergetics 1797, 212–221.





61.    Passarini F., Wientjes E., Hienerwadel R. and Croce R (2009) “Molecular Basis of Light Harvesting and Photoprotection in CP24.  UNIQUE FEATURES OF THE MOST RECENT ANTENNA COMPLEX” J. Biol. Chem, 43, 29536–29546.


60.    Caffarri S., Kouril R., Kereiche S., Boekema EJ and Croce R. (2009) “Functional architecture of photosystem II supercomplexesEMBO Journal. 28, 3052-3063


59.    A.A. Kavalenka, R.B. Spruijt, C.J.A.M. Wolfs, J. Štrancar, R. Croce, M.A. Hemminga, H. van Amerongen (2009) “Site-Directed Spin Labeling Study of CP29” Biophys. J. 96(9) 3620 – 3628


58.    Ballottari M., Mozzo M., Croce R., Morosinotto T. and Bassi R. (2009) “Pigment binding sites occupancy and functional architecture of the Photosystem II antenna complex Lhcb5” J. Biol. Chem. 284, 8103–8113


57.    Wientjes E., Oostergetel G.T., Jansson S., Boekema E.J. and Croce R. (2009) “The role of Lhca complexes in the supramolecular organization of Photosystem I” J. Biol. Chem. 284  7803–7810.


56.    van Oort B., Murali S., Wientjes E., Koehorst R., Spruijt R., van Hoek A., Croce R. and van Amerongen H. (2009) ” Ultrafast resonance energy transfer from a site-specifically attached fluorescent chromophore reveals the folding of the N-terminal domain of CP29”. Chem. Phys. 357, 113–119


55.    Romero E., Mozzo M., van Stokkum I.H.M., Dekker J.P., van Grondelle  and R. Croce R. (2009) ” The Origin of the Low-Energy Form of Photosystem I Light-Harvesting Complex Lhca4: Mixing of the Lowest Exciton with a Charge-Transfer State”. Biophys. J. 96(5), L35 - L37





54.    van Oort B., Amunts A., Borst J.W., van Hoek A., Nelson N., van Amerongen H. and Croce R. (2008) “Picosecond fluorescence of intact and dissolved PSI‑LHCI crystals”. Biophys. J. 95, 5851–5861.


53.    Miloslavina Y., Wehner A., Wientjes E., Reus M., Lambrev P., Garab G., Croce R., and Holzwarth A.R. (2008) “Far-red fluorescence: A direct spectroscopic marker for  LHC II oligomers forming a charge transfer state in non-photochemical quenching in vivo” . FEBS Lett. 582, 3625–3631


52.    Mozzo M., Passarini F., Bassi R., van Amerongen H. and Croce R. (2008) “Photoprotection in higher plants: The putative quenching site is conserved in all outer light-harvesting complexes of Photosystem II ”.  Biochim.Biophys.Acta - Bioenergetics 1777(10): 1263-1267.


51.    Broess K., Trinkunas G., van Hoek A, Croce R., and van Amerongen H. (2008) “Excitation energy transfer within photosystem II is much faster than in isolated light-harvesting complexes”. ”. Biochim.Biophys.Acta - Bioenergetics 1777(5):404-409.


50.    Mozzo M., Dall’Osto L., Hienerwadel R., Bassi R. and Croce R. (2008) “Photoprotection in the antenna complexes of Photosystem II: role of individual xanthophylls in Chlorophyll triplet quenching”. J. Biol. Chem. 283, 6184-6192.





49.    Caffarri S., Passarini F., Bassi R. and Croce R. (2007) “A specific binding site for neoxanthin in the monomeric antenna proteins CP26 and CP29 of Photosystem II”. FEBS Lett. 581, 4704-4710.


48.    Croce R., Chojnicka A., Morosinotto T., Ihalainen J.A., van Mourik F., Dekker J.P., Bassi R. and van Grondelle R. (2007) “The low-energy forms of Photosystem I light-harvesting complexes:  spectroscopic properties and pigment-pigment interaction characteristics” Biophys. J. 93,  2418-2428.


47.    S. Georgakopoulou, G. van der Zwan, R. Bassi, R. van Grondelle,  H. van Amerongen and R Croce (2007) “Understanding the circular dichroism of Light Harvesting Complex II upon Varying its Pigment Composition and organisation”. Biochemistry 46, 4745-4754.


46.    Croce R., Mozzo M., Morosinotto T., Romeo A., Hienerwadel R. and Bassi R. (2007) “Singlet and triplet state transitions of carotenoids in the antenna complexes of higher plants Photosystem I”. Biochemistry 46, 3846-3855.





45.    Mozzo M., Morosinotto T., Bassi R. and Croce R. (2006) “Probing the structure of Lhca3 by mutation analysis” ”. Biochim.Biophys.Acta - Bioenergetics, 1757, 1607-1613.





44.    Ihalainen J.A., Croce R., Morosinotto T., van Stokkum I.H.M., Bassi R., Dekker J.P. and van Grondelle R. (2005) “Excitation decay pathways of Lhca proteins – A time resolved fluorescence study”. J. Phys. Chem. B 109, 21150-21158.


43.    Sener M.K., Jolley C., Ben-Shem A., Fromme P., Nelson N., Croce R. and Schulten K. (2005) “Comparison of the light harvesting networks of plants and cyanobacterial Photosystem I” Biophys. J. 89,1630-1642.


42.    Consoli E., Croce R., Dunlap D.D. and  Finzi L. (2005), “Diffusion of Light Harvesting Complex II in the thylakoid membranes”. EMBO Report 6, 782-786.


41.    Morosinotto T., Mozzo M., Bassi R. and Croce R. (2005) “Pigment-Pigment interactions in Lhca4 Antenna complex of Higher Plants”. J. Biol. Chem. 280: 20612 – 20619.


40.    Gibasiewicz K., Croce R.,.Morosinotto T, Ihalainen J., van Stokkum I., Dekker J.P, Bassi R. and van Grondelle R. (2005) “Excitation energy transfer pathways in Lhca4”. Biophys. J. 88, 1959-1969.





39.    Croce R. , Morosinotto T., Ihalainen J.A., Chojnicka A., Breton J., Dekker J. P, van Grondelle R. and Bassi R..(2004) “Origin of the 701 nm fluorescence emission of the Lhca2 subunit of higher plant    Photosystem I ” J. Biol. Chem. 279: 48543 – 48549.


38.    Caffarri S., Croce R., Cattivelli L. and Bassi R. (2004) “A look within LHCII: differential analysis of the Lhcb 1-3 gene products composing the major antenna complex of higher plant photosynthesis”. Biochemistry 43(29)  9467 – 9476.





37.    Morosinotto T., Breton J., Bassi R. and Croce R. (2003) “The nature of a chlorophyll ligand in Lhca proteins determines the far red fluorescence emission typical of Photosystem I”. J. Biol. Chem. 278(49):49223-9.


36.    Ihalainen JA, Ratsep M., Jensen PE, Scheller HV, Croce R, Bassi R, Korppi-Tommola JEI and Freiberg A (2003) “Red spectral forms of Chlorophylls in green plant PSI – A site selective and High pressure Spectroscopy Study”. J. Phys. Chem. B 107, 9086-9093.


35.    Gastaldelli M, Canino G., Croce R. and Bassi R. (2003) “ Xanthophyll binding sites of the CP29 (Lhca4) subunit of higher plants photosystem II investigated by domine swapping and mutation analysis” J. Biol.Chem 278, 19190-19198.


34.    Castelletti S, Morosinotto T, Robert B, Caffarri S., Bassi R. and Croce R. (2003)” The recombinant Lhca2 and Lhca3 subunits of the photosystem I antenna system” Biochemistry 42, 4226-4234.


33.    Croce R., Müller M.G., Caffarri S., Bassi R. and Holzwarth A.R. (2003) “Selective study of Carotenoid to Chlorophyll Energy transfer pathways in the minor antenna of Photosystem II CP29. A femtosecond study on mutant and WT complexes”. Biophys. J. 84, 2517-2532.


32.    Croce R., Müller M.G., Bassi R. and Holzwarth A.R. (2003) “Energy transfer processes between Chls b and Chls a in the minor antenna of Photosystem II CP29. A comparative femtosecond study between native and  reconstituted complex”. Biophys. J. 84, 2508-2516.


31.    Jennings RC, Zucchelli G., Croce R. and Garlaschi FM. (2003) “Thermally activated energy transfer from the red absorbing chlorophyll states to the bulk antenna in PSI-LHCI”. ”. Biochim.Biophys.Acta - Bioenergetics 1557, 91-98.





30.    Croce R., Morosinotto T., Castelletti S., Breton J. and Bassi R. (2002) “The Lhca antenna complexes of higher plants Photosystem I” ”. Biochim.Biophys.Acta - Bioenergetics 1556, 29-40.


29.    Morosinotto T., Castelletti S., Breton J., Bassi R. and Croce R. (2002) “Mutation analysis of Lhca1 complex: the low energy absorption forms originate from pigment-pigment interaction.” J. Biol.Chem. 277, 36253-36261.


28.    Croce R., Canino G., Ros F and Bassi R. (2002) “Chromophores organization of the higher plants photosynthetic antenna complex CP26”. Biochemistry 41, 7334-7343.


27.    Pascal A., Caffarri, S., Croce R., Sandonà D., Bassi R. and Robert B. (2002) “A Structural investigation of the central Chlorophyll a-binding sites in the Minor Photosystem II Antenna protein, Lhcb4. Biochemistry 41, 2305-2310.




26.    Gobets B., Kennis J.T.M., Ihalainen J.A., Brazzoli M., Croce R., van Stokkum I.H.M., Bassi R., Dekker J.P., van Amerongen H., Fleming G.R. and van Grondelle R. (2001) “Excitation energy transfer in dimeric light-harvesting complex I: a combined streak-camera/fluorescence up-conversion study” J. Phys. Chem. B 105, 10132-10139.


25.    Caffarri S., Croce R., Breton J. and Bassi R. (2001) “The major antenna complex of Photosystem II (LHCII) has a Xanthophyll Binding Site not Involved in Ligth-Harvesting”. J. Biol. Chem. 276, 35924-35933.


24.    Frank H.A., Das S. K., Bautista J.A., Bruce D., Vasil´ev S., Crimi M., Croce R. and Bassi R. (2001) “The Photochemical Behavior of Xanthopylls in the Recombinant Photosystem II Antenna Complex CP26”. Biochemistry, 40, 1020-1025.


23.    Croce R., Müller M.G., Bassi R. and Holzwarth A.R. (2001) “Carotenoid to Chlorophyll Energy transfer in recombinant Major Light-Harvesting Complex (LHCII) of Higher Plants. 1. Femtosecond Transient Absorption Measurements” Biophys. J. 80, 901-915.




22.    Cinque G., Croce R. and Bassi R. (2000) “ Absorption spectra of chlorophyll a and b in Lhcb protein environment” Photosynth. Res. 64, 233-242.


21.    Croce R., Cinque G., Holzwarth A.R. and Bassi R. (2000) “The Soret absorption properties of carotenoids and chlorophylls in antenna complexes of higher plants” Photosynth. Res. 64, 221-231.


20.    Cinque G., Croce R., Holzwarth, A.R. and Bassi R. (2000) “Förster energy transfer in the chlorophyll a/b complex CP29” Biophys.J. 79, 1706-1717.


19.    Ihalainen J.A., Gobets B., Sznee K., Brazzoli, M., Croce R., Bassi R., van Grondelle R., Korppi-Tommola J.E.I. and Dekker J.P. (2000) “Evidence for two spectroscopically different dimers of light-harvesting complex I from green plants” Biochemistry 39, 8625-8631.


18.    Croce R., Dorra D., Holzwarth A.R. and Jennings R. (2000) “Fluorescence decay and spectral evolution in intact Photosystem I of higher plants” Biochemistry 39, 6341-6348.




17.    Remelli R., Varotto C., Sandona´ D., Croce R. and Bassi R. (1999) “Chlorophyll binding sites of monomeric light harvesting complex (LHCII) reconstituted in vitro: a mutation analysis of chromophore binding residues”. J. Biol. Chem. 274, 33510-33521.


16.    Simonetto R., Crimi M., Sandona´ D., Croce R., Cinque G.F., Breton J. and Bassi R. (1999) “Orientation of chlorophyll transition moments in the higher plants light harvesting complex CP29”. Biochemistry 38 (40), 12974-12983.


15.    Croce R., Weiss S. and Bassi R. (1999) “Carotenoid-binding sites in the major light-harvesting complex (LHCII) of higher plants” J. Biol. Chem 274(42), 29613-29623.


14.    Bassi R., Croce R., Cugini D. and Sandona´ D. (1999) “Structure of the higher plant Light-Harvesting protein CP29 probed by mutation analysis”. Proc.Nat.Am.Sci. USA, 96: 10056-10061.


13.    Croce R., Remelli R., Varotto C., Breton J. and Bassi R. (1999) “The neoxanthin binding site of the major light harvesting complex (LHCII) form higher plants”. FEBS Lett. 456 (1), 1-6.


12.    Verhoeven A. S., Adams III W. W., Demmig-Adams B., Croce R. and Bassi R. (1999) “Xanthophyll cycle pigment localization and dynamics during exposure to low temperatures and light stress in low and high light-acclimated Vinca major”. Plant Physiol. 120 (3), 727-737.




11.    Croce R., Zucchelli G., Garlaschi F.M. and Jennings R.C. (1998) “A thermal broadening study of the antenna chlorophylls in PSI-200, LHCI and PSI-core”. Biochemistry 37 (50), 17355-17360.


10.    Sandona´ D. Croce R., Pagano A., Crimi M. and Bassi R. (1998) “Higher Plants Light Harvesting Proteins. Structure and Function as revealed by mutation analysis of either protein or chromophore moieties”. ”. Biochim.Biophys.Acta - Bioenergetics 1365, 207-214.




9.       Giuffra, E., Zucchelli, G., Sandonà, D., Croce, R., Cugini, D., Garlaschi, F.M., Bassi, R. and Jennings, R.C. (1997).“Analysis of some optical properties of a native and reconstituted photosystem II antenna complex, CP29: pigment binding sites can be occupied by chlorophyll a or chlorophyll b and determine spectral forms”. Biochemistry 36, 12984-12993.


8.       Jennings, R.C., Zucchelli, G, Croce, R., Valkunas, L., Finzi, L. and Garlaschi, F.M. (1997) “Model Studies on the excited state equilibrium perturbation due to reaction center trapping in photosystem I” Photosynth. Res. 52, 245-253.


7.       Bassi, R., Sandonà, D. and Croce, R. (1997) “Novel aspect of chlorophyll a/b proteins” Physiol. Plantarum. 100 (4), 769-779.


6.       Connelly, J.P., Muller, M.G., Bassi, R., Croce, R. and Holzwarth, A. (1997) “Femtosecond transient absorption study of carotenoid to chlorophyll energy transfer in the light harvesting complex II of photosystem II”. Biochemistry 36 (2), 281-287.




5.       Testi, M.G., Croce, R., Polverino, P. and Bassi, R. (1996) " A CK2 site is reversibly phosphorylated in the photosystem II subunit CP29." FEBS Lett. 399 (3), 245-250.


4.       Croce, R., Breton, J. and Bassi, R. (1996) " Conformational changes induced by phosphorylation in the photosystem II subunity CP29" Biochemistry 35, 11142-11148.


3.       Croce, R., Zucchelli, G, Garlaschi, F.M., Bassi, R. and Jennings, R.C. (1996) "Excited state equilibration in the Photosystem I-Light-Harvesting I complex: P700 is almost isoenergetic with its antenna”. Biochemistry 35, 8572-8579.


2.       Giuffra, E., Cugini, D., Croce, R. and Bassi, R.(1996) "Reconstitution and Pigment Binding Protein of Recombinant CP29".Eur. J. Biochem. 238, 112-120.





1.       Zucchelli, G., Garlaschi, F.M., Croce, R., Bassi, R. and Jennings, R.C.(1995) “A Stepanov relation analysis of steady state absorption and fluorescence spectra in the isolated D1/D2/cytochrome b-559 complex”. Biochim.Biophys.Acta - Bioenergetics 1229, 59-63.









Contributions to books


Croce R. “Chlorophylls and light-harvesting complexes” (2013) In: Encyclopedia of Biophysics. Springer. Editor: Gordon C. Roberts


Croce R. “Chlorophyll binding proteins of plants and cyanobacteria” (2011) In: Advances in Photosynthesis and Respiration. Photosynthesis: Plastid Biology, Energy Conversion and Carbon Assimilation. Edited by Julian Eaton-Rye and Baishnab C. Tripathy.


Van Amerongen H and Croce R. “Structure and function of Photosystem II light-harvesting proteins (Lhcb) of Higher Plants”.(2008) In Comprehensive Series in Photochemical and Photobiological Sciences – Primary Processes of Photosynthesis: Principles and Apparatus (ed. G. Renger). Publisher, RCS. Vol I, pp. 329-368.

Croce R., Morosinotto T. and Bassi R. “LHCI: the antenna complex of Photosystem I in plants and green algae”. (2006) In “Photosystem I: The Plastocyanin/Ferredoxin Oxidorecutase” in Photosynthesis-Edited by John Golbeck.  series: Advances in Photosynthesis and Respiration Kluwer Academic Publishers pp.119-137.


Bassi, R., Giuffra, E., Croce, R., Dainese, P. and Bergantino, E. (1996) "Biochemistry and molecular biology of pigment binding proteins". NATO Advanced Scientific Courses: "Light as energy source and information carrier in plant photophysiology", Volterra, 26.09-0610.94- ed. Jennings et al., 41-65.