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. 2018 Jan 11;8:518. doi: 10.1038/s41598-017-19074-3

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© The Author(s) 2018

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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The higher-order Te2 cortex provides essential information to LA and BA during auditory memory retrieval. (a) Lesions were centered in the secondary auditory cortex Te2 (n = 9). Scale bar, 500 μm. (b) A mixed-design ANOVA indicated a significant main effect of the zone (F_(1,34) =5.36, P = 0.027), significant main effects of the group in zone 1 (F_(2,34) = 6.48, P = 0.004) and zone 6 (F_(2,34) =6.27, P = 0.005). Pairwise comparisons showed significant differences between Te2-lesioned and conditioned animals in zones 1 (P = 0.015) and 6 (P = 0.044). Indeed, the distribution of the time spent in zones 1 and 6 was similar in Te2-lesioned and naive animals (P > 0.05). (c) Avoidance behavior was weaker (F_(2,34) =6.68, P = 0.004) in Te2-lesioned animals relative to conditioned rats (P = 0.020). (d) The freezing responses of Te2-lesioned rats were similar to naive animals (F_(2,34) = 25.6, P < 0.001; naive versus Te2-lesioned animals, P > 0.05; Te2-lesioned versus conditioned rats, P < 0.001). (e) The Te2 region of the rats was injected with rAAV5/CamKIIa-eNpHR3.0-mCherry-WPRE. (f) Bilateral placement of optic fiber tips within the LA and BA. Orange dots represent eNpHR-mCherry-injected animals, while yellow dots represent mCherry control animals. (g) Representative micrographs showing the expression of eNpHR3.0-mCherry (in red) in Te2 and its terminals in either LA and BA. Nuclei were counterstained with DAPI. Scale bars, 300, 200, 20, 20 μm respectively. (h) eNpHR-mCherry-injected animals (n = 8) spent significantly less time in zone 1 and significantly more time in zone 6 than mCherry controls (n = 8). (i, j) Animals injected with eNpHR-mCherry exhibited significantly lower (i) avoidance and (j) freezing than mCherry controls. (k) There were no differences in the freezing response to the presentation of an innate threat stimulus. *P < 0.05; **P < 0.01; ***P < 0.005. All values are reported as the mean ± SEM. See also Supplementary Fig. S4.