Article
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Acknowledgements We thank M. Mukundan, B. Wong and L. Bao for technical support;
R. Beltramo for help with extracellular recordings; J. Isaacson, G. Keller, R. Nicoll and
M. Heindorf for comments on the manuscript; the members of the Scanziani laboratory for
discussions of this project as well as for comments on the manuscript; and M. Rio for software
support. This project was supported by the National Institutes of Health grant U19NS107613,
the Howard Hughes Medical Institute and the Swiss National Science Foundation grants
P300PA_177882 and P2EZP3_162284 to A.J.K. and P300PA_177898 to M.M.R. Confocal images
were acquired at the Nikon Imaging Center at the University of California San Francisco.
Author contributions A.J.K. and M.S. designed the study. A.J.K. and M.M.R. conducted all
experiments and analysis. M.S., A.J.K. and M.M.R. wrote the manuscript.
Competing interests The authors declare no competing interests.
Additional information
Supplementary information is available for this paper at https://doi.org/10.1038/s41586-020-
2319-4.
Correspondence and requests for materials should be addressed to A.J.K. or M.S.
Peer review information Nature thanks Richard Born, Mark Hübener and the other,
anonymous, reviewer(s) for their contribution to the peer review of this work.
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