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. 2017 Aug 15;7:8282. doi: 10.1038/s41598-017-08331-0

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© The Author(s) 2017

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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Layer 2b is the main source of feedback from the APC to the OB. (A) Schematic representation of injection of the tracer into the OB, and imaging from sagittal sections of the APC. (B) Injections were targeted to the granule cell layer of the OB. *, injection site. Red: retrograde tracer; blue: DAPI. (C) Representative sagittal section used for APC imaging. Left, DAPI labeling shows the main anatomical landmarks: dense cell layer 2 of the APC; rf: rhinal fissure; OT: olfactory tubercle; Hc: hippocampus; Ctx: neocortex and Str: striatum. Right, Retrogradely-labeled cells were found mainly in the APC in those sections. Some labeling was also observed in the MCPO: magnocellular preoptic nucleus and nLOT: nucleus of the lateral olfactory tract. (D) Higher magnification image showing retrogradely labeled cells across APC layers. Superficial limit of the layer 2 (border between layers 1 and 2a) is defined with a depth of 0 while a depth of 1 is the deep end of that layer (limit between layers 2b and 3). Red: retrograde tracer. Blue: DAPI. (E) Bar graph showing the relative fractions of retrogradely labeled neurons, normalized by the number of DAPI cells in each layer. OB-projecting neurons were heterogeneously distributed across APC layers (p < 0.0001, total cell count: 546 Retrobeads + , 4474 DAPI + cells, n = 13 sections, 11 mice, Friedman test). Within layer 2, cells were more densely found in layer 2b than in layer 2a (p = 0.005, n = 84 2a cells vs. 205 2b cells, Dunn’s multiple comparisons post-hoc test). **P < 0.01. (F) Cumulative distribution of the OB-projecting cells within layer 2. On the x-axis, 0 indicates the border between layers 1 and 2a, while 1 is the limit between layers 2b and 3 (see panel D). The light green and red curves show the results from the counting obtained in a single optical section with green and red Retrobeads, respectively. Distributions obtained from green and red Retrobeads were not significantly different (p = 0.47, Kolmogorov-Smirnov test, n = 856 cells, 14 sections, 6 mice for the green Retrobeads, n = 598 cells, 13 sections, 6 mice for the red Retrobeads). The distribution of all the bead-labeled cells (thick red trace) was shifted to deeper part of the layer 2 compared to the distribution of the DAPI + cells (thick blue trace; p < 0.0001, Kolmogorov-Smirnov test, n = 658 DAPI + cells).