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. 2020 Jul 7;10:11130. doi: 10.1038/s41598-020-67974-8

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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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Metabolic pathway for the synthesis of gamma-hydroxybutyric acid (GHB) and glutamate/glutamine shuttle in neurons. Succinic semi-aldehyde (SSA) is formed in the mitochondria by GABA-transaminase (GABA-T). SSA can either be oxidized to succinate by succinic semialdehyde dehydrogenase (SSADH) in the mitochondria or transported to the cytosol where it is reduced to gamma-aminobutyric acid (GABA) by succinic semialdehyde reductase (SSR). Glutamate can form GABA by glutamate decarboxylase (GAD). Glutamine is broken down to glutamate under the action of glutaminase. Glutamate is converted to glutamine by glutamine synthetase (GS). In the low dose tramadol group, the content of SSA was down regulated and the content of GHB was up regulated. In the high dose tramadol group, the content of glutamine was down regulated and the content of GHB was up regulated.