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ACKNOWLEDGMENTS
We thank F. Hara, K. Umeno, N. Hamaoka, Y. Misumi, J. Y. Kim,
C. König, M. Völkel, and R. Oworah-Nkruma for excellent technical
assistance and Y. Miyanoiri, E. Yamashita, and A. Nakagawa for support
during data collection by x-ray and NMR analysis. J.M.S. is grateful
to E. Conti for scientific independence and great mentorship and to
J. M. Plitzko and W. Baumeister for access to the cryo-EM
infrastructure and early career support. P.S. thanks A. Boussac and
B. Lagoutte for purification of PSI and Fd. M.M.N. is grateful to his
mentor M. Rögner for generous support. Synchrotron radiation
experiments were performed at BL44XU of SPring-8, Japan (proposal
no. 2017B6500).Funding:Financial support was provided by the
Max Planck Society, the Cluster of Excellence RESOLV (EXC 1069 to
W.L., N.C., and M.M.N.) funded by the German Research Council
(DFG), the DFG research unit FOR2092 (EN 1194/1-1 to B.D.E.
and NO 836/3-2 to M.M.N.), the DFG priority program 2002
(NO 836/4-1 to M.M.N.), DFG grant NO 836/1-1 (to M.M.N.), the
Australian Research Council (FT140100834 to N.C.), JST-CREST
(JPMJCR13M4 to G.K.), MEXT-KAKENHI (16H06560 to G.K.), the
French Infrastructure for Integrated Structural Biology/FRISBI
ANR-10-INSB-05 (P.S.), and the International Joint Research
Promotion Program, Osaka University (G.K. and M.M.N.).Author
contributions:J.M.S., J.A.B., N.C., W.L., P.S., B.D.E., G.K., and

M.M.N. conceived the research, prepared the figures, and
wrote the manuscript with the contributions of all other
authors.
M.M.N. coordinated the activities. Preparation of complex I,
biochemical analysis, and mass spectrometry were done
by H.W., J.T., and M.M.N. J.M.S. and B.D.E. performed the
cryo-EM analysis and built the structural model with the
help of S.K.S. and G.K. Flash-absorption spectroscopy was
carried out by P.S. EPR experiments were conducted by J.A.B.
and N.C. and NMR and x-ray analysis by H.T., T.K., T.I., and G.K.
All authors approved the final version of the manuscript.
Competing interests: The authors declare no competing
interests.Data and materials availability:The cryo-EM
density map is deposited in the Electron Microscopy
Data Bank under accession number EMD-0281. The atomic
models of the cryo-EM and x-ray structures are deposited
in the worldwide Protein Data Bank under accession
numbers 6HUM and 6A7K, respectively. All other data are
available in the manuscript or the supplementary materials.
SUPPLEMENTARY MATERIALS
http://www.sciencemag.org/content/363/6424/257/suppl/DC1
Materials and Methods
Supplementary Text
Figs. S1 to S19
Tables S1 to S6
References ( 33 – 88 )

20 September 2018; accepted 6 December 2018
Published online 20 December 2018
10.1126/science.aau3613

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