The binding of photosystem I (PS I) from Thermosynechococcus elongatus to the native cytochrome (cyt) c6 and cyt c from horse heart (cyt cHH) was analyzed by oxygen consumption measurements, isothermal titration calorimetry (ITC), and rigid body docking combined with electrostatic computations of binding energies. Although PS I has a higher affinity for cyt cHH than for cyt c6, the influence of ionic strength and pH on binding is different in the two cases. ITC and theoretical computations revealed the existence of unspecific binding sites for cyt cHH besides one specific binding site close to P700 Binding to PS I was found to be the same for reduced and oxidized cyt cHH Based on this information, suitable conditions for cocrystallization of cyt cHH with PS I were found, resulting in crystals with a PS I:cyt cHH ratio of 1:1. A crystal structure at 3.4-Å resolution was obtained, but cyt cHH cannot be identified in the electron density map because of unspecific binding sites and/or high flexibility at the specific binding site. Modeling the binding of cyt c6 to PS I revealed a specific binding site where the distance and orientation of cyt c6 relative to P700 are comparable with cyt c2 from purple bacteria relative to P870 This work provides new insights into the binding modes of different cytochromes to PS I, thus facilitating steps toward solving the PS I-cyt c costructure and a more detailed understanding of natural electron transport processes.

Biosystems Technology Institute for Applied Life Sciences University of Applied Sciences Wildau Hochschulring 1 15745 Wildau Germany

Department of Chemical Physics and Optics Charles University Ke Karlovu 3 CZ 121 16 Praha 2 Czech Republic

Department of Theoretical Biophysics Institute for Theoretical Physics Johannes Kepler University Linz Altenberger Strasse 69 4040 Linz Austria and

From the Biophysics of Photosynthesis Institute for Biology Humboldt Universität zu Berlin Philippstrasse 13 10115 Berlin Germany

Lawrence Berkeley National Laboratory Berkeley California 94720

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PDB
1l9b, 1jb0, 1hrc, 1c6s