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. 2017 Jun 6;7:2885. doi: 10.1038/s41598-017-03119-8

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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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Summary of present study. Mechanism of muscle thermogenesis starts with shivering as first defense against exposure to cold. Thereafter, Sln ^{22, 50} and Ucp1 ^{51, 52} are respectively driven to elicit non-shivering thermogenesis in skeletal muscle and BAT. Exposure to cold also induces emergence of brown adipocytes in WAT by increasing expression of Ucp1, Cidea, and Pgc1a genes^{52, 53}. Meanwhile, the present study found induced circadian Slc25a25 gene expression in skeletal muscle, but not in BAT and WAT of mice fed with ketogenic diet (KD), which induces torpor. Messenger RNA expression of thermogenesis-related genes such as Ucp1 was not affected by KD in BAT and WAT. Sciatic denervation abolished Slc25a25 expression and augmented KD-induced hypothermia. These results suggest that skeletal muscle is involved in thermogenesis during KD feeding, and that Slc25a25 and Ucp3 are candidates for muscle thermogenesis. Less Slc25a25 was expressed in skeletal muscle of aged, compared with adult mice, whereas Ucp1 expression was not altered by aging at thermoneutrality⁵⁴. SLC25A25 might help to maintain Tb in aged mammals.