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. 2022 Mar 24;13:1601. doi: 10.1038/s41467-022-29208-5

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© The Author(s) 2022

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. 3 — a–e Artificial iSOS in neocortex during training can rescue fear memory deficits in HPC-lesioned mice. a Scheme of the co-activation LEC fiber during encoding. The iSOS were induced by LED on cortical surface to activate oCHiEF-expressing widespread axons from LEC L5. Activation of cortical fibers but not LEC cell bodies is to avoid the unspecific activation of circuits from LEC to other brain regions. b Flow of the behavioral experiment. All four groups of mice received contextual fear conditioning (CFC) training. Two groups of HPC-lesioned mice were given iSOS during training trials. Long-term memory was tested 24 h later without iSOS. HPC intact group means no HPC lesion, no iSOS applied and no virus infected. c Travel distance of open field test upon given iSOS or not. (HPC-lesioned mice, N = 13, two-sided paired t-test, t(12) = 1.6, P = 0.1449). d Immediate freezing after foot shock (N_{HPC intact} = 15, N_{HPC lesion} = 19, N_iSOS-10Hz = 8; N_iSOS-30Hz = 8; ANOVA, F(3, 46) = 9.1, P < 0.0001; Bonferroni post-hoc test, P_{Control vs. HPC-lesion} = 0.0005, P_{iSOS-10Hz vs. HPC-lesion} = 0.0035, P_{iSOS-30Hz vs. HPC-lesion} = 0.0005). e Memory test in the conditioned context (same mice as d, ANOVA, F (3, 46) = 13.5, P < 0.0001; Bonferroni post-hoc test, P_{Control vs. HPC-lesion} < 0.0001, P_{iSOS-10Hz vs. HPC-lesion} < 0.0001, P_{iSOS-30Hz vs. HPC-lesion} < 0.0001). f–j Artificial iSOS rescued spatial memory deficit in HPC-lesioned mice. f Flow of the behavioral experiment. g Learning curves for the spatial memory task (mice number: N_HPC-intact = 24, N_HPC-lesion = 15, N_{iSOS (30Hz,EYFP)} = 6, N_{iSOS (10Hz,oCHiEF)} = 11; N_{iSOS (30Hz,oCHiEF)} = 22; ANOVA, Time factor: F(3, 219) = 52.5, P < 0.0001; Group factor: F(4, 73) = 10.6, P < 0.0001; Interaction, F(12, 219) = 1.4, P = 0.1453; Bonferroni post-hoc test, P_{HPC-intact vs. HPC-lesion} < 0.0001, P _{iSOS (30Hz,EYFP) vs. HPC-lesion} > 0.9999, P _{iSOS (10Hz,oCHiEF) vs. HPC-lesion} < 0.0001, P _{iSOS (30Hz,oCHiEF) vs. HPC-lesion} < 0.0001). h Averaged occupancy maps for memory recall in day5. i Quantification of occupancy in food zone (same mice as g. ANOVA, F(4, 73) = 5.0, P = 0.0012; Bonferroni post-hoc test, P_{HPC-intact vs. HPC-lesion} = 0.0048, P _{iSOS (30Hz,EYFP) vs. HPC-lesion} > 0.9999, P _{iSOS (10Hz,oCHiEF) vs. HPC-lesion} = 0.0243, P _{iSOS (30Hz,oCHiEF) vs. HPC-lesion} = 0.0098). j Quantification of occupancy in four quadrants (same mice as g ANOVA for each quadrant, Target quadrant: F(4, 73) = 5.0, P = 0.0013; Bonferroni post-hoc test, P_{HPC-intact vs. HPC-lesion} = 0.0159, P _{iSOS (30Hz,EYFP) vs. HPC-lesion} = 0.9803, P _{iSOS (10Hz,oCHiEF) vs. HPC-lesion} = 0.0102, P _{iSOS (30Hz,oCHiEF) vs. HPC-lesion} = 0.0070, Opposite quadrant: F(4, 73) = 4.3, P = 0.0038, Bonferroni post-hoc test, P _{iSOS (30Hz,oCHiEF) vs. HPC-lesion} = 0.0190; Left quadrant: F(4, 73) = 1.4, P = 0.2529; Right quadrant: F(4, 73) = 1.0, P = 0.4043). *P < 0.05, **P < 0.01, ***P < 0.001. Each dot represents one mouse. Error bar shows lines shows S.E.M. For all box plot, whiskers show min and max, box shows 25th, median and 75th percentile.