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. 2020 Nov 18;9(1):1–13. doi: 10.1093/gastro/goaa066

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© The Author(s) 2020. Published by Oxford University Press and Sixth Affiliated Hospital of Sun Yat-sen University

This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

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Figure 2. — Schematic of glutamine synthesis, nucleotide biosynthesis, and the pentose-phosphate pathway. Nucleotide metabolism and its supporting processes such as the pentose-phosphate pathways and glutamine synthesis are significantly altered in HCC. Nucleotides are important precursors for DNA and RNA, as well as the energy currency ATP, which are needed for cell growth and proliferation. The exact mechanisms by which these pathways promote HCC tumorigenesis remain unclear however. Metabolite nomenclature: 6PG, 6-phosphogluconate; 6PGL, 6-phosphogluconolactone; ADP, adenosine diphosphate; AICAR, 5-aminoimidazole-4-carboxamide ribonucleotide; dADP, deoxyadenosine diphosphate; dGDP, deoxyguanosine diphosphate; dTMP, deoxythymidine monophosphate; dUMP, deoxyuridine monophosphate; E4P, erythrose-4-phosphate; F6P, fructose-6-phosphate; FAICAR, 5-formamidoimidazole-4-carboxamide ribonucleotide; G3P, glyceraldehyde-3-phosphate; G6P, glucose-6-phosphate; GDP, guanosine diphosphate; NADPH, reduced nicotinamide adenine dinucleotide phosphate; R5P, ribose-5-phosphate; Ru5P, ribulose-5-phospahte; S7P, sedoheptulose-7-phosphate; SAICAR, succinyl 5-aminoimidazole-4-carboxamide ribonucleotide; Xu5P, xylulose-5-phospahte. Enzyme nomenclature: 5NT, 5’-nucleotidase; 6PGD, 6-phosphogluconate dehydrogenase; 6PGL, 6-phosphogluconolactonase; ADSL, adenylosuccinate lyase; ADSS, adenylosuccinate synthetase; AK2, adenylate kinase 2; ATIC, 5-aminoimidazole-4-carboxamiade ribonucleotide formyltransferase/IMP cyclohydrolase; CAD, carbamoyl phosphate synthetase 2 aspartate transcarbamylase, and dihydroorotase; CMPK1, cytidine/uridine monophosphate kinase 1; CTPS1/2, cytidine triphosphate synthase 1/2; DHODH, dihydroorotate dehydrogenase; DTYML, deoxythymidylate kinase; G6PD, glucose-6-phosphate dehydrogenase; GART, glycinamide ribonucleotide synthetase, aminoimidazole ribonucleotide synthetase, and glycinamide ribonucleotide transformylase; GLS1, glutaminase 1; GMPS, guanine monophosphate synthase; GS, glutamine synthetase; GUK1, guanylate kinase 1; IMPDH, inosine monophosphate dehydrogenase; NDPK, nucleoside diphosphate kinase; PAICS, phosphoribosylaminoimidazole carboxylase and phosphoribosylaminoimidazole succinocarboxamide synthase; PFAS, phosphoribosylformylglycinamide synthase; PNP, purine nucleoside phosphorylase; PPAT, phosphoribosyl pyrophosphate amidotransferase; PRPS1, phosphoribosyl pyrophosphate synthetase 1; RPE, ribulose-5-phosphate-3-epimerase; RPI, ribose-5-phosphate isomerase; RRM1/2, ribonucleoside diphosphate reductase 1/2; TALDO1, transaldolase 1; TK1, thymidine kinase 1; TKT, transketolase; TYMP, thymidine phosphorylase; TYMS, thymidylate synthase; UCK1/2, uridine-cytidine kinase 1/2; UMPS, uridine monophosphate synthetase; UPP1, uridine phosphorylase 1.