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. 2020 May 5;11:2217. doi: 10.1038/s41467-020-15840-6

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© The Author(s) 2020

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 1 — a Representative example of a CA1 pyramidal cell recorded juxtacellularly from a head-fixed running mouse. See average action potential waveform and autocorrelation at right, and morphological identification as deep cell at bottom. Scale bar 50 µm. b Theta phase-locked firing histogram of the cell shown in a. Raw local field potential (LFP) traces and juxtacellular signals are shown below: SO, stratum oriens; SP, stratum pyramidale; SR, stratum radiatum; SLM, stratum lacunosum moleculare. c Theta phase firing histogram from single cells recorded in awake head-fixed mice, ranked according to their preferred phase (n = 12 cells). Individual (gray) and mean (black) theta filtered LFP signals at top are z-scored (Z). The population histogram at bottom represents the distribution of mean preferred phases from individual cells. d Same for single pyramidal cells recorded juxtacellularly in freely moving rats (n = 28 cells). e Individual and mean ± SD values of vector length per cells reported before in head-fixed and freely moving conditions. No differences between groups. f Awake/REM sleep (left) and RUN/no-RUN (running vs. other motor behavior) dependency of theta phase preference of single pyramidal cells. See legend in (g). g Multivariate analysis of morphologically identified cells demonstrate effects per location (Harrison–Kanji test for deep-superficial location: χ²(2) = 10.7, p = 0.0136) and preparation (χ²(2) = 8.5, p = 0.0046); no interaction (χ²(1) = 2.1, p = 0.148). Phase preferred data from each morphologically identified cell is depicted separately for deep and superficial cells in both preparations. h Optogenetic tagging of single units using micro-LED optoelectrodes in two transgenic lines allowed identifying superficial (Calb1-cre + AAV5-DIO-ChR2-YFP) and deep CA1 pyramidal cells (Thy1-ChR2). Raster at right shows response of one representative unit to 79 trials of nanowatt blue light stimulation in a Thy1-ChR2 mouse. Bottom, ChR2 signal (white) and Calbindin immunostaining (magenta) are shown in false colors (one confocal plane from each line). Scale bar 25 µm. i Phase-locked firing histograms from Calbindin+ superficial (n = 7 units from 3 mice) and Thy1+ deep opto-tagged units (n = 14 units from 2 mice; statistically different Watson–Williams test, F(1,20) = 17.7, p = 0.00048. Note that bimodal distribution of preferred theta phases can be explained by different cell-type contribution.