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The authors dedicate this paper to G. T. Babcock and M. P. Klein. J.B. and S.I. acknowledge support from the Biotechnology and Biological Research Science Council. We thank the Centre for Structural Biology and Bioinformatics Facility at Imperial College London for technical support; and C. Schulze-Briese and T. Tomizaki at PX06SA/SLS Paul Scherrer Institute Villigen Switzerland; B. Shepard at ID29; and M. Iwata of the ERATO ATP System Project for help with data collections. T.M.I. has been a Life Sciences Research Foundation Fellow of the Howard Hughes Medical Institute a European Molecular Biology Organization Long-Term Fellow and a Ruth L. Kirschstein National Research Science Award Fellow and acknowledges support from D. C. Rees during the initial stages of the study. We wish to thank P. Siegbahn M. Lundberg P. Nixon C. Dismukes and A. Telfer for comments and helpful discussions. The coordinates together with the structure factors native Mn-edge and long wavelength (2.25 Å for Ca detection) and the experimental multi-wavelength anomalous dispersion (MAD) phases have been deposited in the Protein Data Bank (entry 1S5L) and will be available upon publication.