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Acknowledgements JRS was supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (grant XDB13030000), a ‘1000 talents’ professorship from the Ministry of Science and Technology (MOST) of the Chinese government, and a Wolfson award from the Royal Society. SEM was supported by the US National Institute of Health grant R01AG043972 and MM was supported by a TWAS studentship of the Chinese Academy of Sciences, during the preparation of this manuscript. We are grateful to three anonymous referees for their constructive and helpful comments.