PMC:4301621 / 6498-28945 JSONTXT

Annnotations TAB JSON ListView MergeView

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/4301621","sourcedb":"PMC","sourceid":"4301621","source_url":"https://www.ncbi.nlm.nih.gov/pmc/4301621","text":"Results and discussions\n\nCharacterization of the M. alpina GSMM iCY1106\nThe genome-scale metabolic model iCY1106 reconstructed in this study included 1106 genes representing 9.51% of the 11631 protein-coding genes in the genome of M. alpina ATCC 32222. The model consisted of 1854 reactions and 1732 metabolites (Table 1). Compared with the previous model constructed by Vongsangnak [21], iCY1106 is more detailed and contains a higher degree of functional information that can be used in in silico simulations. During the construction of iCY1106, some missing gene were added from the Vongsangnak’s model such as MA-090-247, which encodes an asparaginase (EC: 3.5.1.1), and catalyzes the transformation of L-asparagine into L-aspartate. When compared with the existing model, many improvements in iCY1106 are apparent. (1) A total of 139 transport proteins were annotated based on TCDB and TransportDB. (2) A total of 216 exchange reactions were added into model iCY1106. (3) Biomass composition of M. alpina was calculated based on literature mining. (4) Additional chemical reactions were also included. (5) The reversibility of some reactions was redefined according to MetaCyc and BioPath databases. For example, in the published model, the reaction catalyzed by aldehyde dehydrogenase (EC: 1.2.1.3) was reversible, whereas this reaction is irreversible according to BioPath. The flux distribution of iCY1106 was normal following this change.\nTable 1 General features of model i CY1106\nFeatures i CY1106 Existed model a\nGenome feature\n Genome size (Mb) 38.38 38.38\n Total open reading frames (ORFs) 11631 11631\nMetabolic model\n Total reactions 1854 1183\n Biochemical reactions 1391 1183\n Transport reactions 247 none\n Exchange reactions 216 none\n Metabolites 1732 1660\n ORFs associated in model 1106 1042\n ORF coverageb (%) 9.51 8.95\naThe existed model was constructed by Vongsangnak et al. [21]; bThe number of ORFs incorporated in model iCY1106 divided by the total number of ORFs.\nIn model iCY1106, four compartments (extracellular, cytoplasmic, peroxisome and mitochondrial compartments) were linked by 141 trans-plasma membrane transport reactions, 98 cytoplasmic-mitochondrial transport reactions and eight cytoplasmic-peroxisome transport reactions (Additional file 1). The extracellular compartment contained 267 reactions, including 216 exchange reactions and 51 metabolic reactions. These metabolic reactions catalyzed by extracellular enzymes may enable M. alpina to adapt to different environmental conditions. For example, M. alpina grows on different carbohydrates such as glycerol [24], sucrose [24], d-mannose [25], and raffinose [25] using various extracellular galactosidases. The peroxisome compartment contained 34 reactions mostly related to glyoxylate metabolism.\nMetabolic reactions in the model iCY1106 were divided into ten metabolic subsystems (Figure 1). The largest subsystem was lipid metabolism (fatty acid biosynthesis, fatty acid elongation, steroid, glycerolipid, glycerophospholipid and sphingolipid metabolism) that accounted for 23.53% of the total reactions. Reactions involving the synthesis of PUFAs such as DGLA, ARA, and EPA are listed in model iCY1106. For most subsystems, most categories were more abundant in iCY1106 than in the Vongsangnak’s model, with the exception of Amino Acid Metabolism and Nucleotide Metabolism (Figure 1a). Since there were six compartments (cytosol, mitochondria, peroxisome, extracellular, plasma membrane and unclassified compartment) in Vongsangnak’s model, some reactions occur in multiple compartments at the same time. When ignoring compartments, not including transport reactions and exchange reactions, these two models shared 566 reactions (Additional file 2). Vongsangnak’s model and iCY1106 included 179 and 621 unique reactions, respectively (Figure 1b). Compared with model iYL619_PCP of the oleaginous yeast Y. lipolytica [20], lipid metabolism accounted for a higher proportion (16.7%, 191 of total 1142 reactions) of reactions in iCY1106. Amino acid and carbohydrate metabolism subsystems were 13.17% and 11.66%, respectively, in iCY1106.\nFigure 1 The comparison between Vongsangnak’s model and i CY1106. a: reaction distribution in subsystems between these two models; b: shared reactions and unique reactions between models.\n\nIdentification of essential genes and reactions in iCY1106\nWhen cultured on minimal growth medium (MG) containing glucose, KH2PO4, MgSO4 and NaNO3 [23], 86 M. alpina genes (7.78% of the total) were identified as essential for growth using the FBA method (Additional file 3). In contrast, on yeast extract (YE) medium (based on MG but including all 20 regular amino acids) only 49 genes were identified as essential. On MG medium, over half of the essential genes were involved in amino acid (36.05%) and nucleotide metabolism (23.26%; Figure 2). However on YE medium, most of the essential genes were associated with nucleotide metabolism (38.78%).\nFigure 2 Essential genes identified by MG and YE medium in model i CY1106.\nThe identified genes were compared with the database of essential genes (DEG), and 40 genes were found to share high homology with genes identified by DEG (identity ≥ 40%, E-value ≤ 1.0E-30; Table 2). One gene, UTP-glucose-1-phosphate uridylyltransferase (MA-090-485), is involved in the synthesis of the cell wall component 1,3-beta-D-glucan (Table 2). MA-090-485 is highly homologous with the Saccharomyces cerevisiae enzyme (sequence identity = 62.58%, e-value = 0) identified in the DEG search. Similarly, nucleoside diphosphate kinase (MA-120-261), which is involved in the synthesis of CTP and GTP, also shared high homology with one of the essential genes identified by DEG (identity = 66.89%, e-value = 2.0E-68). Besides, in MG, there were 46 genes that did not match with DEG (Additional file 3). Of these, when nitrate was used as nitrogen source, nitrate reductase (MA-291-2) and nitrite reductase (MA-291-3), which together convert nitrate into NH3 for cell growth, were essential under this condition. Another two essential genes, delta 5 desaturase (MA-326-160) and delta 12 desaturase (MA-334-414) are vital for the synthesis of ARA in M. alpina. In addition, most of these essential genes were distributed in the Amino Acid Metabolism category (28.3%). This was understandable since essential gene identified by DEG were in a rich medium in which genes involving amino acid synthesis may not be essential.\nTable 2 Essential genes identified on different mediums which are high homology with DEG\nGene Enzyme Essentiality Subsystem\nMG YE\nMA-133-4 Ribose-5-phosphate isomerase E E Carbohydrate metabolism\nMA-090-485 UTP-glucose-1-phosphate uridylyltransferase E E\nMA-101-200 Mannose-6-phosphate isomerase E E\nMA-120-141 Phosphoacetylglucosamine mutase E E\nMA-184-380 Acetyl-CoA carboxylase E E\nMA-072-196 Argininosuccinate lyase E NE Amino acid metabolism\nMA-213-50 Adenylosuccinate synthase E E\nMA-213-539 Adenylosuccinate lyase E E\nMA-326-6 Aspartate carbamoyltransferase E E\nMA-334-434 Phosphoribosylanthranilate isomerase E NE\nMA-120-148 2-Acetolactate methylmutase E NE\nMA-090-434 Dihydroxy-acid dehydratase E NE\nMA-139-331 Homoaconitate hydratase E NE\nMA-323-77 Saccharopine dehydrogenase E NE\nMA-326-106 Ornithine carbamoyltransferase E NE\nMA-101-393 Imidazole-4-carboxamide isomerase E NE\nMA-184-558 Imidazoleglycerol-phosphate dehydratase E NE\nMA-090-452 Glutamine amidotransferase:cyclase E NE\nMA-320-159 3-dehydroquinate synthase E NE\nMA-139-157 Chorismate synthase E NE\nMA-073-62 3-deoxy-7-phosphoheptulonate synthase E NE\nMA-213-547 Tryptophan synthase E NE\nMA-297-40 Chorismate mutase E NE\nMA-120-157 Thioredoxin reductase E E\nMA-173-30 Purine-nucleoside phosphorylase E E Nucleotide metabolism\nMA-213-65 IMP dehydrogenase E E\nMA-120-261 Nucleoside diphosphate kinase E E\nMA-139-347 Guanylate kinase E E\nMA-120-138 Ribonucleoside-diphosphate reductase E E\nMA-055-211 Ribonucleoside-diphosphate reductase E E\nMA-323-58 Ribonucleoside-diphosphate reductase E E\nMA-334-356 Thymidylate synthase E E\nMA-153-455 3(2),5-bisphosphate nucleotidase E NE\nMA-111-23 phosphoadenylyl-sulfate reductase E NE Energy metabolism\nMA-182-360 6,7-dimethyl-8-ribityllumazine synthase E E Cofactors and vitamins\nMA-184-368 Riboflavin synthase E E\nMA-210-311 Aspartate dehydrogenase E E\nMA-055-340 Fatty-acyl-CoA synthase E E Lipid metabolism\nMA-162-131 Palmitoyl-protein thioesterase E E\nMA-334-239 Very-long-chain enoyl-CoA reductase E E\nE: essential gene; NE: non-essential gene. MG: minimal medium; YE: yeast extract medium.\n\nSimulation and verification of model iCY1106\nFBA with constraints was used to investigate the metabolic properties of model iCY1106. YE medium was used to simulate the growth rate and ARA production, the maximum uptake rate of all 20 amino acids were set at 0.01 mmol gDW−1 h-1 [26], and the glucose consumption rate was set at 0.8 mmol gDW−1 h-1 [8], and 2.0 mmol gDW−1 h-1 [1] respectively, and the in silico growth rate were closed to experimental values (differences were only 0.8% and 9.48%; Table 3). To investigate ARA production in silico, the growth rate was fixed at 0.03 h-1 [8], and the ARA exchange reaction was set as the objective function. Under these conditions, the ARA rate was 0.128 mmol gDW−1 h−1 by FBA. This in silico value was lower than the experimentally determined value of 0.149 mmol gDW−1 h−1 (14.09%) [8], and this may due to the use of two different M. alpina strains (ATCC 32222 and ME-1) for modeling and experimental components, with ME-1 being the higher ARA producing strain [27]. Furthermore, the difficulties associated with accurately modeling the yeast extract may have resulted in a nutritional deficiency in the model.\nTable 3 Comparison of in silico and in vivo growth rates of M . alpina\nMedia condition (mmol gDW −1  h −1 ) Growth rate (h −1 ) Ref.\nin vivo in silico\nGlc (v = 0.8) 0.0696 0.0690 [8]\nGlc (v = 2.0) 0.1708 0.1546 [1]\nin vivo: experimental results; in silico: simulation results.\nThe capability of utilizing 14 different carbon sources for cell growth was qualitatively predicted by the model iCY1106 using FBA (Additional file 4). Each substance was the sole carbon source and the rate of uptake was 1.0 mmol gDW−1 h−1. Initially, there were four types of carbon source (ethanol, xylose, maltose, and rhamnose), and the model could not achieve growth with any individual carbon source. For ethanol, the reaction catalyzed by alcohol dehydrogenase (EC: 1.1.1.1) was irreversible, but should be reversible according to the MetaCyc database. For rhamnose, the lack of L-rhamnulose-1-phosphate lactaldehyde-lyase (EC: 4.1.2.19) meant that rhamnose could not generate glycerone phosphate, which could otherwise be used by the glycolysis pathway. After elimination of initial discrepancies using the continuous gap filling process, a 100% match was acquired. This indicated that model iCY1106 could predict catabolic pathways of various carbon sources including common sugars and alcohols. A similar FBA simulation of nitrogen sources (ammonium, nitrate, urea, and amino acids) also generated results that were consistent with experimental fermentation data. Both glutamate and glycine supported M. alpina growth, confirming that the model performed well for predicting the utilization of different carbon and nitrogen sources.\n\nPUFA biosynthetic pathways based on GSMM\nTo further investigate the pathways associated with PUFA biosynthesis and metabolism, genome annotation, literature mining and KEGG database analysis were performed. ARA is a ω-6 PUFA synthesized by M. alpina (Figure 3). ARA (c204(6)) is synthesized from acetyl-CoA in 38 enzymatic steps, including the de novo synthesis of palmitic acid (c160) and the synthesis of very long chain fatty acids. During PUFA biosynthesis, all fatty acids should be in the form of acyl-CoA [2]. Desaturase and elongase were necessary for the conversion of octadecanoyl-CoA (c180coa) to ARA. Δ9 desaturase [28] (MA-055-51, MA-101-533, MA-184-235), Δ12 desaturase [29] (MA-334-414) and Δ6 desaturase [30,31] (MA-268-54, MA-101-36) catalyzed the conversion of c180coa into γ-linolenoyl-CoA (c183(6)coa) which was subsequently lengthened to 8,11,14-eicosatrienoyl-CoA (c203(6)coa) by an elongase such as ELO/GLELO [30] (MA-189-257), MAELO [32] (MA-184-206) or MALCE1 [33] (MA-073-327, MA-320-221). Δ5 desaturase [10] (MA-326-160) catalyzed c203(6)coa to arachidonyl-CoA (c204(6)coa), which was finally hydrolyzed to c204(6) by choloyl-CoA hydrolase (MA-049-4, MA-139-47).\nFigure 3 Synthesis pathway of PUFAs in M . alpina . (accoa: acetyl-CoA, acp: acyl-carrier protein, acacp: Acetyl-[acyl-carrier protein], malcoa: malonyl-CoA, malacp: malonyl-[acp], aacacp: acetoacetyl [acp], c4hacp: (3R)-3-Hydroxybutanoyl-[acp], c4dacp: but-2-enoyl-[acp], c40acp: butanoyl-[acp], c6oacp: 3-oxohexanoyl-[acp], c6hacp: 3-hydroxyhexanoyl-[acp], c6dacp: hex-2-enoyl-[acp], c60acp: hexanoyl-[acp], c8oacp: 3-oxooctanoyl-[acp], c8hacp: 3-hydroxyoctanoyl-[acp], c8dacp: oct-2-enoyl-[acp], c80acp: octanoyl-[acp], c10oacp: 3-oxodecanoyl-[acp], c10hacp: 3-hydroxydecanoyl-[acp], c10dacp: dec-2-enoyl-[acp], c100acp: decanoyl-[acp], c12oacp: 3-oxododecanoyl-[acp], c12hacp: 3-hydroxy-dodecanoyl-[acp], c12dacp: dodec-2-enoyl-[acp], c120acp: dodecanoyl-[acp], c14oacp: 3-oxomyristoyl-[acp], c14dacp: tetradec-2-enoyl-[acp], c14hacp: 3-hydroxymyristoyl-[acp], c140acp: myristoyl-[acp], c16oacp: 3-oxohexadecanoyl-[acp], c16hacp: (3R)-3-Hydroxypalmitoyl-[acp], c16dacp: hexadec-2-enoyl-[acp], c160acp: hexadecanoyl-[acp], c160: palmitic acid, c160coa: hexadecanoyl-CoA, c180coa: octadecanoyl-CoA, c181coa: octadecenoyl-CoA, c182coa: linoleoyl-CoA, c183(6)coa: gamma-Linolenoyl-CoA, c203(6)coa: 8,11,14-Eicosatrienoyl-CoA, c204(6)coa: Arachidonyl-CoA, c204(6): ARA, c183(3)coa: alpha-Linolenoyl-CoA, c184(3)coa: Stearidonoyl-CoA, c204(3)coa: (11Z,14Z,17Z)-Icosatrienoyl-CoA, c205(3)coa: eicosapentaenoyl-CoA, c225(3)coa: (7Z,10Z,13Z,16Z,19Z)-Docosapentaenoyl-CoA, c245(3)coa: (9Z,12Z,15Z,18Z,21Z)-Tetracosaheptaenoyl-CoA, c246(3)coa: (6Z,9Z,12Z,15Z,18Z,21Z)-Tetracosahexaenoyl-CoA, c226(3)coa: (4Z,7Z,10Z,13Z,16Z,19Z)-Docosahexaenoyl-CoA).\nM. alpina is capable of producing ARA and EPA, and EPA can be produced via the ω-3 or the ω-6 PUFA biosynthetic pathways through ARA desaturation [34]. During EPA biosynthesis, Δ15 (ω3) desaturase (MA-072-10) is the key enzyme and was the first fungal desaturase known that uses both 18-carbon and 20-carbon ω-6 PUFAs as substrates [35]. In the ω-3 pathway, (11Z,14Z,17Z)-icosatrienoyl-CoA (c204(3)coa) was used as substrate, while the ω-6 pathway utilized c204(6)coa.\nBoth the maximum and minimum flux of oxygen absorption of DGLA, ARA and EPA production in silico were found to be 1.552, 1.674 and 1.891 mmol gDW−1 h−1 respectively, using FVA. This indicated that the degree of PUFA unsaturation was dependent on oxygen content. Robustness analysis was used to elucidate the effects of oxygen uptake rate on PUFA biosynthesis. The oxygen uptake rate was perturbed by constraints-based flux analysis between 0 and 20 mmol gDW−1 h−1. The minimum growth rate was set at 0.03 h−1, with ARA exchange reaction as the objective function. When oxygen absorption was lower than 2.0 mmol gDW−1 h−1, ARA production increased with oxygen absorption rate (Additional file 5a). However, beyond this range, ARA production gradually declined with increasing oxygen uptake rate. As reported by Higashiyama et al., when the DO concentration range was 10-15 ppm, the ARA yield was enhanced 1.6-fold over a DO of 7 ppm, and a DO between 20 and 50 ppm drastically decreased ARA production [36]. This indicated that excessive oxygen exposure during fermentation could impair ARA overproduction. During ARA biosynthesis, high oxygen concentrations may affect metabolism and cell growth in the filamentous mycelia, and beta-oxidation of fatty acids may be needed to obtain the extra energy required for adaptation to the high oxygen conditions.\n\nRegulation of PUFA biosynthesis and metabolism based on GSMM\nAcetyl-CoA is the essential two-carbon donor molecule for fatty acid synthesis. In model iCY1106, there were ten pathways that could produce acetyl-CoA, including amino acid degradation (from aspartate, isoleucine, leucine, and lysine), fatty acid beta-oxidation, and other acetyl-CoA generation processes (phosphoenolpyruvate [PEP], malate, L-lactate, acetate, and citrate) (Figure 4). However, only the pathway from PEP to pyruvate could generate sufficient flux for synthesis of PUFAs on the fermentation medium. During the growth stage, the flux of acetyl-CoA generated by pyruvate was 1.22 mmol gDW−1 h−1, while this value was 1.56 mmol gDW−1 h−1 during the product synthesis stage. This corresponds to a 27.87% increase, which would ensure acetyl-CoA was available for ARA production due to the enhanced activity of the pyruvate dehydrogenase complex (EC: 1.2.4.1, 2.3.1.12, 1.8.1.4) [37]. Although aspartate, isoleucine, leucine, and lysine could all be used to generate acetyl-CoA for ARA production. Asparate has four carbon, while isoleucine, leucine, and lysine all contains six carbon. When the maximum uptake rate of asparate and the other amino acids were set at 0.15 mmol gDW−1 h−1 and 0.1 mmol gDW−1 h−1, respectively. FBA showed only that leucine and lysine could increase ARA production by 14.06% and 13.28%.\nFigure 4 The sources of acetyl-CoA in M . alpina . (cit: citrate, ac: acetate, llac: l-lactate, mal: malate, pep: phosphoenolpyruvate, pyr: pyruvate, accoa: acrtyl-CoA, asp: l-aspartate, bala: beta-alanine, oppa: 3-oxopropanoate, ile: l-isoleucine, omval: (s)-3-nethyl-2-oxopentanoate, mbcoa: 2-methylbutanoyl-CoA, mcrocoa: 2-methylcrotonoyl-CoA, metbycoa: (2S,3S)-3-Hydroxy-2-methylbutanoyl-CoA, mcecoa: 2-Methylacetoacetyl-CoA, lys: l-lysine, sacp: l-saccharopine, amasa: l-2-aminoadipate 6-semialdehyde, ama: l-2-aminoadipate, aka: 2-oxoadipate, gltcoa: glutaryl-CoA, c4dcoa: crotonoyl-coa, c4hcoa: (S)-3-Hydroxybutanoyl-CoA, aaccoa: acetoacetyl-CoA, leu: l-leucine, 4mop: 4-methyl-2-oxopentanoate, ivacoa: 3-isovaleryl-coa, mcrcoa: 3-nethylcrotonyl-CoA, mgcoa: trans-3-methylglutaconyl-CoA, hmgcoa: (s)-3-hydroxy-3-methylglutaryl-coa).\nIn model iCY1106, acetyl-CoA can be consumed by amino acid synthesis, amino sugar metabolism and fatty acid synthesis as well as ARA production (Additional file 5b). During the synthesis of six amino acids (threonine, methionine, lysine, cysteine, ornithine, and leucine), acetyl-CoA was required. FBA results showed that during the growth stage, 96.5% of the acetyl-CoA flux was used to synthesize fatty acids and 2.58% was used for amino acid synthesis. In contrast, during the product synthesis stage, the flux of acetyl-CoA used for fatty acid synthesis accounted for 99.52%, which indicates that amino acid synthesis was inhibited during ARA production. Limiting the nitrogen source can be an effective strategy to control amino acids biosynthesis [38], in which the lipid yield may be increased despite decreases in mycelia concentration.\nDuring the growth stages, the flux of acetyl-CoA used to synthesize malonyl-CoA, catalyzed by acetyl-CoA carboxylase (ACC, EC 6.4.1.2), was 0.79 mmol gDW−1 h−1. In contrast during the ARA production stage, the corresponding flux was 1.43 mmol gDW−1 h−1 (81.0%). ACC catalyzes the first committal step of the fatty acid biosynthetic pathway, and should be overexpressed to maximize ARA production [39]. Glutamate has been shown to activate ACC, and adding glutamate to the culture medium led to an increase in total lipid (21.81%) and ARA yield (66.07%) [40]. As the M. alpina growth rate in batch culture ranged from 0.06 h−1 to 0.19 h-1 [1,7,8,41,42], the glutamate uptake rates used for the in silico analysis should be set between 0 and 1.8 mmol gDW−1 h−1. When the glutamate uptake rate was increased, ARA production increased from 0.128 mmol gDW−1 h−1 to 0.355 mmol gDW−1 h−1 (Additional file 5c).\nDuring PUFA biosynthesis, NADPH is a necessary cofactor for de novo fatty acid synthesis involving desaturation and elongation. In model iCY1106, there were 172 reactions involving NADPH (Additional file 6). With ARA production fixed at 0.059 mmol gDW−1 h−1 (μ = 0.06 h−1) and 0.128 mmol gDW−1 h−1 (μ = 0.03 h−1), the flux of NADPH reactions were investigated. In total, 53 NADPH reactions exhibited flux changes of \u003e 10−6 mmol gDW−1 h−1 (Figure 5).\nFigure 5 Changes of reactions involving NADPH at different ARA levels. (Different colors represent different changes on NADPH flux. Flux obviously changed reactions are listed. R2: D-glucose 6-phosphate + NADP ⬄ 6-phospho-D-glucono-1,5-lactone + NADPH + H, R3: 6-phospho-D-gluconate + NADP - \u003e D-ribulose 5-phosphate + CO2 + NADPH + H, R4: (S)-malate + NADP - \u003e pyruvate + CO2 + NADPH, R5: L-glutamate 5-semialdehyde + NAD P+ H2O - \u003e L-glutamate + NADPH + H, R6: 1-pyrroline-5-carboxylate + NADPH + H ⬄ L-proline + NADP, R16: Nitrite + 3 NADPH + 5 H - \u003e Ammonia + 3 NADP + 2 H2O, R51: N-acetyl-L-glutamyl 5-phosphate + NADPH + H ⬄ N-acetyl-L-glutamate 5-semialdehyde + NADP + Phosphate, R53: (S)-3-hydroxy-3-methylglutaryl-CoA + 2 NADPH + 2 H ⬄ (R)-mevalonate + CoA + 2 NADP).\nFurther research showed that only five reactions (R2, R3, R4, R5, and R6) involving accumulation of NADPH were significantly altered. When ARA production was increased, three reactions (R4, R5, and R6) underwent flux increases, which corresponded to enhanced enzyme activities due to up-regulation of the genes encoding the enzymes. Of these, malic enzyme (ME, EC 1.1.1.40), which catalyzes the conversion of malate to pyruvate (R4), is considered a key enzyme in lipogenesis in M. alpina. ME expression and enzyme activity are enhanced during ARA production [43], and the minimization of metabolic adjustment (MOMA) method was used to investigate the role of this enzyme. With the biomass and ARA flux maximized by ME deletion using MOMA, 410 (22.11%) reactions showed flux changes (Additional file 6) compared with the wild type model. The ARA exchange reaction flux was lowered from 0.128 mmol gDW−1 h−1 to 0.079 mmol gDW−1 h−1, a decrease of 38.28%. Additionally, all reactions involving NADPH consumption reactions other than R13, R16, R51, and R53 were associated with lipid metabolism. These results indicated that the increase in ARA production was directly correlated with the NADPH consumption rate.","divisions":[{"label":"title","span":{"begin":0,"end":23}},{"label":"sec","span":{"begin":25,"end":4323}},{"label":"title","span":{"begin":25,"end":71}},{"label":"p","span":{"begin":72,"end":1992}},{"label":"table-wrap","span":{"begin":1448,"end":1992}},{"label":"label","span":{"begin":1448,"end":1455}},{"label":"caption","span":{"begin":1456,"end":1491}},{"label":"p","span":{"begin":1456,"end":1491}},{"label":"table","span":{"begin":1492,"end":1841}},{"label":"tr","span":{"begin":1492,"end":1527}},{"label":"th","span":{"begin":1492,"end":1501}},{"label":"th","span":{"begin":1501,"end":1510}},{"label":"th","span":{"begin":1511,"end":1527}},{"label":"tr","span":{"begin":1528,"end":1542}},{"label":"td","span":{"begin":1528,"end":1542}},{"label":"tr","span":{"begin":1543,"end":1572}},{"label":"td","span":{"begin":1543,"end":1560}},{"label":"td","span":{"begin":1561,"end":1566}},{"label":"td","span":{"begin":1567,"end":1572}},{"label":"tr","span":{"begin":1573,"end":1618}},{"label":"td","span":{"begin":1573,"end":1606}},{"label":"td","span":{"begin":1607,"end":1612}},{"label":"td","span":{"begin":1613,"end":1618}},{"label":"tr","span":{"begin":1619,"end":1634}},{"label":"td","span":{"begin":1619,"end":1634}},{"label":"tr","span":{"begin":1635,"end":1661}},{"label":"td","span":{"begin":1635,"end":1651}},{"label":"td","span":{"begin":1652,"end":1656}},{"label":"td","span":{"begin":1657,"end":1661}},{"label":"tr","span":{"begin":1662,"end":1694}},{"label":"td","span":{"begin":1662,"end":1684}},{"label":"td","span":{"begin":1685,"end":1689}},{"label":"td","span":{"begin":1690,"end":1694}},{"label":"tr","span":{"begin":1695,"end":1724}},{"label":"td","span":{"begin":1695,"end":1715}},{"label":"td","span":{"begin":1716,"end":1719}},{"label":"td","span":{"begin":1720,"end":1724}},{"label":"tr","span":{"begin":1725,"end":1753}},{"label":"td","span":{"begin":1725,"end":1744}},{"label":"td","span":{"begin":1745,"end":1748}},{"label":"td","span":{"begin":1749,"end":1753}},{"label":"tr","span":{"begin":1754,"end":1776}},{"label":"td","span":{"begin":1754,"end":1766}},{"label":"td","span":{"begin":1767,"end":1771}},{"label":"td","span":{"begin":1772,"end":1776}},{"label":"tr","span":{"begin":1777,"end":1812}},{"label":"td","span":{"begin":1777,"end":1802}},{"label":"td","span":{"begin":1803,"end":1807}},{"label":"td","span":{"begin":1808,"end":1812}},{"label":"tr","span":{"begin":1813,"end":1841}},{"label":"td","span":{"begin":1813,"end":1831}},{"label":"td","span":{"begin":1832,"end":1836}},{"label":"td","span":{"begin":1837,"end":1841}},{"label":"table-wrap-foot","span":{"begin":1842,"end":1992}},{"label":"p","span":{"begin":1842,"end":1992}},{"label":"p","span":{"begin":1992,"end":2793}},{"label":"p","span":{"begin":2794,"end":4323}},{"label":"figure","span":{"begin":4135,"end":4323}},{"label":"label","span":{"begin":4135,"end":4143}},{"label":"caption","span":{"begin":4144,"end":4323}},{"label":"p","span":{"begin":4144,"end":4323}},{"label":"sec","span":{"begin":4325,"end":8586}},{"label":"title","span":{"begin":4325,"end":4383}},{"label":"p","span":{"begin":4384,"end":5049}},{"label":"figure","span":{"begin":4974,"end":5049}},{"label":"label","span":{"begin":4974,"end":4982}},{"label":"caption","span":{"begin":4983,"end":5049}},{"label":"p","span":{"begin":4983,"end":5049}},{"label":"p","span":{"begin":5050,"end":8587}},{"label":"table-wrap","span":{"begin":6472,"end":8587}},{"label":"label","span":{"begin":6472,"end":6479}},{"label":"caption","span":{"begin":6480,"end":6561}},{"label":"p","span":{"begin":6480,"end":6561}},{"label":"table","span":{"begin":6562,"end":8497}},{"label":"tr","span":{"begin":6562,"end":6599}},{"label":"th","span":{"begin":6562,"end":6567}},{"label":"th","span":{"begin":6567,"end":6574}},{"label":"th","span":{"begin":6575,"end":6588}},{"label":"th","span":{"begin":6589,"end":6599}},{"label":"tr","span":{"begin":6600,"end":6606}},{"label":"th","span":{"begin":6600,"end":6603}},{"label":"th","span":{"begin":6603,"end":6606}},{"label":"tr","span":{"begin":6607,"end":6672}},{"label":"td","span":{"begin":6607,"end":6615}},{"label":"td","span":{"begin":6616,"end":6644}},{"label":"td","span":{"begin":6645,"end":6646}},{"label":"td","span":{"begin":6647,"end":6648}},{"label":"td","span":{"begin":6649,"end":6672}},{"label":"tr","span":{"begin":6673,"end":6731}},{"label":"td","span":{"begin":6673,"end":6683}},{"label":"td","span":{"begin":6684,"end":6727}},{"label":"td","span":{"begin":6728,"end":6729}},{"label":"td","span":{"begin":6730,"end":6731}},{"label":"tr","span":{"begin":6732,"end":6776}},{"label":"td","span":{"begin":6732,"end":6742}},{"label":"td","span":{"begin":6743,"end":6772}},{"label":"td","span":{"begin":6773,"end":6774}},{"label":"td","span":{"begin":6775,"end":6776}},{"label":"tr","span":{"begin":6777,"end":6823}},{"label":"td","span":{"begin":6777,"end":6787}},{"label":"td","span":{"begin":6788,"end":6819}},{"label":"td","span":{"begin":6820,"end":6821}},{"label":"td","span":{"begin":6822,"end":6823}},{"label":"tr","span":{"begin":6824,"end":6861}},{"label":"td","span":{"begin":6824,"end":6834}},{"label":"td","span":{"begin":6835,"end":6857}},{"label":"td","span":{"begin":6858,"end":6859}},{"label":"td","span":{"begin":6860,"end":6861}},{"label":"tr","span":{"begin":6862,"end":6923}},{"label":"td","span":{"begin":6862,"end":6872}},{"label":"td","span":{"begin":6873,"end":6896}},{"label":"td","span":{"begin":6897,"end":6898}},{"label":"td","span":{"begin":6899,"end":6901}},{"label":"td","span":{"begin":6902,"end":6923}},{"label":"tr","span":{"begin":6924,"end":6963}},{"label":"td","span":{"begin":6924,"end":6933}},{"label":"td","span":{"begin":6934,"end":6959}},{"label":"td","span":{"begin":6960,"end":6961}},{"label":"td","span":{"begin":6962,"end":6963}},{"label":"tr","span":{"begin":6964,"end":7001}},{"label":"td","span":{"begin":6964,"end":6974}},{"label":"td","span":{"begin":6975,"end":6997}},{"label":"td","span":{"begin":6998,"end":6999}},{"label":"td","span":{"begin":7000,"end":7001}},{"label":"tr","span":{"begin":7002,"end":7045}},{"label":"td","span":{"begin":7002,"end":7010}},{"label":"td","span":{"begin":7011,"end":7041}},{"label":"td","span":{"begin":7042,"end":7043}},{"label":"td","span":{"begin":7044,"end":7045}},{"label":"tr","span":{"begin":7046,"end":7098}},{"label":"td","span":{"begin":7046,"end":7056}},{"label":"td","span":{"begin":7057,"end":7093}},{"label":"td","span":{"begin":7094,"end":7095}},{"label":"td","span":{"begin":7096,"end":7098}},{"label":"tr","span":{"begin":7099,"end":7142}},{"label":"td","span":{"begin":7099,"end":7109}},{"label":"td","span":{"begin":7110,"end":7137}},{"label":"td","span":{"begin":7138,"end":7139}},{"label":"td","span":{"begin":7140,"end":7142}},{"label":"tr","span":{"begin":7143,"end":7185}},{"label":"td","span":{"begin":7143,"end":7153}},{"label":"td","span":{"begin":7154,"end":7180}},{"label":"td","span":{"begin":7181,"end":7182}},{"label":"td","span":{"begin":7183,"end":7185}},{"label":"tr","span":{"begin":7186,"end":7225}},{"label":"td","span":{"begin":7186,"end":7196}},{"label":"td","span":{"begin":7197,"end":7220}},{"label":"td","span":{"begin":7221,"end":7222}},{"label":"td","span":{"begin":7223,"end":7225}},{"label":"tr","span":{"begin":7226,"end":7267}},{"label":"td","span":{"begin":7226,"end":7235}},{"label":"td","span":{"begin":7236,"end":7262}},{"label":"td","span":{"begin":7263,"end":7264}},{"label":"td","span":{"begin":7265,"end":7267}},{"label":"tr","span":{"begin":7268,"end":7314}},{"label":"td","span":{"begin":7268,"end":7278}},{"label":"td","span":{"begin":7279,"end":7309}},{"label":"td","span":{"begin":7310,"end":7311}},{"label":"td","span":{"begin":7312,"end":7314}},{"label":"tr","span":{"begin":7315,"end":7364}},{"label":"td","span":{"begin":7315,"end":7325}},{"label":"td","span":{"begin":7326,"end":7359}},{"label":"td","span":{"begin":7360,"end":7361}},{"label":"td","span":{"begin":7362,"end":7364}},{"label":"tr","span":{"begin":7365,"end":7420}},{"label":"td","span":{"begin":7365,"end":7375}},{"label":"td","span":{"begin":7376,"end":7415}},{"label":"td","span":{"begin":7416,"end":7417}},{"label":"td","span":{"begin":7418,"end":7420}},{"label":"tr","span":{"begin":7421,"end":7471}},{"label":"td","span":{"begin":7421,"end":7431}},{"label":"td","span":{"begin":7432,"end":7466}},{"label":"td","span":{"begin":7467,"end":7468}},{"label":"td","span":{"begin":7469,"end":7471}},{"label":"tr","span":{"begin":7472,"end":7513}},{"label":"td","span":{"begin":7472,"end":7482}},{"label":"td","span":{"begin":7483,"end":7508}},{"label":"td","span":{"begin":7509,"end":7510}},{"label":"td","span":{"begin":7511,"end":7513}},{"label":"tr","span":{"begin":7514,"end":7549}},{"label":"td","span":{"begin":7514,"end":7524}},{"label":"td","span":{"begin":7525,"end":7544}},{"label":"td","span":{"begin":7545,"end":7546}},{"label":"td","span":{"begin":7547,"end":7549}},{"label":"tr","span":{"begin":7550,"end":7602}},{"label":"td","span":{"begin":7550,"end":7559}},{"label":"td","span":{"begin":7560,"end":7597}},{"label":"td","span":{"begin":7598,"end":7599}},{"label":"td","span":{"begin":7600,"end":7602}},{"label":"tr","span":{"begin":7603,"end":7638}},{"label":"td","span":{"begin":7603,"end":7613}},{"label":"td","span":{"begin":7614,"end":7633}},{"label":"td","span":{"begin":7634,"end":7635}},{"label":"td","span":{"begin":7636,"end":7638}},{"label":"tr","span":{"begin":7639,"end":7671}},{"label":"td","span":{"begin":7639,"end":7648}},{"label":"td","span":{"begin":7649,"end":7666}},{"label":"td","span":{"begin":7667,"end":7668}},{"label":"td","span":{"begin":7669,"end":7671}},{"label":"tr","span":{"begin":7672,"end":7708}},{"label":"td","span":{"begin":7672,"end":7682}},{"label":"td","span":{"begin":7683,"end":7704}},{"label":"td","span":{"begin":7705,"end":7706}},{"label":"td","span":{"begin":7707,"end":7708}},{"label":"tr","span":{"begin":7709,"end":7776}},{"label":"td","span":{"begin":7709,"end":7718}},{"label":"td","span":{"begin":7719,"end":7750}},{"label":"td","span":{"begin":7751,"end":7752}},{"label":"td","span":{"begin":7753,"end":7754}},{"label":"td","span":{"begin":7755,"end":7776}},{"label":"tr","span":{"begin":7777,"end":7808}},{"label":"td","span":{"begin":7777,"end":7786}},{"label":"td","span":{"begin":7787,"end":7804}},{"label":"td","span":{"begin":7805,"end":7806}},{"label":"td","span":{"begin":7807,"end":7808}},{"label":"tr","span":{"begin":7809,"end":7853}},{"label":"td","span":{"begin":7809,"end":7819}},{"label":"td","span":{"begin":7820,"end":7849}},{"label":"td","span":{"begin":7850,"end":7851}},{"label":"td","span":{"begin":7852,"end":7853}},{"label":"tr","span":{"begin":7854,"end":7885}},{"label":"td","span":{"begin":7854,"end":7864}},{"label":"td","span":{"begin":7865,"end":7881}},{"label":"td","span":{"begin":7882,"end":7883}},{"label":"td","span":{"begin":7884,"end":7885}},{"label":"tr","span":{"begin":7886,"end":7937}},{"label":"td","span":{"begin":7886,"end":7896}},{"label":"td","span":{"begin":7897,"end":7933}},{"label":"td","span":{"begin":7934,"end":7935}},{"label":"td","span":{"begin":7936,"end":7937}},{"label":"tr","span":{"begin":7938,"end":7989}},{"label":"td","span":{"begin":7938,"end":7948}},{"label":"td","span":{"begin":7949,"end":7985}},{"label":"td","span":{"begin":7986,"end":7987}},{"label":"td","span":{"begin":7988,"end":7989}},{"label":"tr","span":{"begin":7990,"end":8040}},{"label":"td","span":{"begin":7990,"end":7999}},{"label":"td","span":{"begin":8000,"end":8036}},{"label":"td","span":{"begin":8037,"end":8038}},{"label":"td","span":{"begin":8039,"end":8040}},{"label":"tr","span":{"begin":8041,"end":8076}},{"label":"td","span":{"begin":8041,"end":8051}},{"label":"td","span":{"begin":8052,"end":8072}},{"label":"td","span":{"begin":8073,"end":8074}},{"label":"td","span":{"begin":8075,"end":8076}},{"label":"tr","span":{"begin":8077,"end":8125}},{"label":"td","span":{"begin":8077,"end":8087}},{"label":"td","span":{"begin":8088,"end":8120}},{"label":"td","span":{"begin":8121,"end":8122}},{"label":"td","span":{"begin":8123,"end":8125}},{"label":"tr","span":{"begin":8126,"end":8192}},{"label":"td","span":{"begin":8126,"end":8135}},{"label":"td","span":{"begin":8136,"end":8169}},{"label":"td","span":{"begin":8170,"end":8171}},{"label":"td","span":{"begin":8172,"end":8174}},{"label":"td","span":{"begin":8175,"end":8192}},{"label":"tr","span":{"begin":8193,"end":8270}},{"label":"td","span":{"begin":8193,"end":8203}},{"label":"td","span":{"begin":8204,"end":8243}},{"label":"td","span":{"begin":8244,"end":8245}},{"label":"td","span":{"begin":8246,"end":8247}},{"label":"td","span":{"begin":8248,"end":8270}},{"label":"tr","span":{"begin":8271,"end":8305}},{"label":"td","span":{"begin":8271,"end":8281}},{"label":"td","span":{"begin":8282,"end":8301}},{"label":"td","span":{"begin":8302,"end":8303}},{"label":"td","span":{"begin":8304,"end":8305}},{"label":"tr","span":{"begin":8306,"end":8344}},{"label":"td","span":{"begin":8306,"end":8316}},{"label":"td","span":{"begin":8317,"end":8340}},{"label":"td","span":{"begin":8341,"end":8342}},{"label":"td","span":{"begin":8343,"end":8344}},{"label":"tr","span":{"begin":8345,"end":8400}},{"label":"td","span":{"begin":8345,"end":8355}},{"label":"td","span":{"begin":8356,"end":8379}},{"label":"td","span":{"begin":8380,"end":8381}},{"label":"td","span":{"begin":8382,"end":8383}},{"label":"td","span":{"begin":8384,"end":8400}},{"label":"tr","span":{"begin":8401,"end":8446}},{"label":"td","span":{"begin":8401,"end":8411}},{"label":"td","span":{"begin":8412,"end":8442}},{"label":"td","span":{"begin":8443,"end":8444}},{"label":"td","span":{"begin":8445,"end":8446}},{"label":"tr","span":{"begin":8447,"end":8497}},{"label":"td","span":{"begin":8447,"end":8457}},{"label":"td","span":{"begin":8458,"end":8493}},{"label":"td","span":{"begin":8494,"end":8495}},{"label":"td","span":{"begin":8496,"end":8497}},{"label":"table-wrap-foot","span":{"begin":8498,"end":8587}},{"label":"p","span":{"begin":8498,"end":8587}},{"label":"sec","span":{"begin":8588,"end":11373}},{"label":"title","span":{"begin":8588,"end":8632}},{"label":"p","span":{"begin":8633,"end":10031}},{"label":"table-wrap","span":{"begin":9749,"end":10031}},{"label":"label","span":{"begin":9749,"end":9756}},{"label":"caption","span":{"begin":9757,"end":9821}},{"label":"p","span":{"begin":9757,"end":9821}},{"label":"table","span":{"begin":9822,"end":9968}},{"label":"tr","span":{"begin":9822,"end":9885}},{"label":"th","span":{"begin":9822,"end":9859}},{"label":"th","span":{"begin":9859,"end":9879}},{"label":"th","span":{"begin":9880,"end":9885}},{"label":"tr","span":{"begin":9886,"end":9904}},{"label":"th","span":{"begin":9886,"end":9894}},{"label":"th","span":{"begin":9894,"end":9904}},{"label":"tr","span":{"begin":9905,"end":9936}},{"label":"td","span":{"begin":9905,"end":9918}},{"label":"td","span":{"begin":9919,"end":9925}},{"label":"td","span":{"begin":9926,"end":9932}},{"label":"td","span":{"begin":9933,"end":9936}},{"label":"tr","span":{"begin":9937,"end":9968}},{"label":"td","span":{"begin":9937,"end":9950}},{"label":"td","span":{"begin":9951,"end":9957}},{"label":"td","span":{"begin":9958,"end":9964}},{"label":"td","span":{"begin":9965,"end":9968}},{"label":"table-wrap-foot","span":{"begin":9969,"end":10031}},{"label":"p","span":{"begin":9969,"end":10031}},{"label":"p","span":{"begin":10031,"end":11373}},{"label":"sec","span":{"begin":11375,"end":16031}},{"label":"title","span":{"begin":11375,"end":11415}},{"label":"p","span":{"begin":11416,"end":14208}},{"label":"figure","span":{"begin":12565,"end":14208}},{"label":"label","span":{"begin":12565,"end":12573}},{"label":"caption","span":{"begin":12574,"end":14208}},{"label":"p","span":{"begin":12574,"end":14208}},{"label":"p","span":{"begin":14209,"end":14677}},{"label":"p","span":{"begin":14678,"end":16031}},{"label":"title","span":{"begin":16033,"end":16093}},{"label":"p","span":{"begin":16094,"end":18261}},{"label":"figure","span":{"begin":17421,"end":18261}},{"label":"label","span":{"begin":17421,"end":17429}},{"label":"caption","span":{"begin":17430,"end":18261}},{"label":"p","span":{"begin":17430,"end":18261}},{"label":"p","span":{"begin":18262,"end":19106}},{"label":"p","span":{"begin":19107,"end":20009}},{"label":"p","span":{"begin":20010,"end":21237}},{"label":"figure","span":{"begin":20460,"end":21237}},{"label":"label","span":{"begin":20460,"end":20468}},{"label":"caption","span":{"begin":20469,"end":21237}},{"label":"p","span":{"begin":20469,"end":21237}}],"tracks":[{"project":"2_test","denotations":[{"id":"25582171-23541380-14905478","span":{"begin":384,"end":386},"obj":"23541380"},{"id":"25582171-23541380-14905479","span":{"begin":1900,"end":1902},"obj":"23541380"},{"id":"25582171-21365447-14905480","span":{"begin":2605,"end":2607},"obj":"21365447"},{"id":"25582171-21365447-14905481","span":{"begin":2619,"end":2621},"obj":"21365447"},{"id":"25582171-23236514-14905482","span":{"begin":3917,"end":3919},"obj":"23236514"},{"id":"25582171-21336613-14905483","span":{"begin":4473,"end":4475},"obj":"21336613"},{"id":"25582171-22124721-14905484","span":{"begin":8867,"end":8869},"obj":"22124721"},{"id":"25582171-17439233-14905485","span":{"begin":8964,"end":8965},"obj":"17439233"},{"id":"25582171-17439233-14905486","span":{"begin":9966,"end":9967},"obj":"17439233"},{"id":"25582171-22174787-14905487","span":{"begin":11888,"end":11889},"obj":"22174787"},{"id":"25582171-16133334-14905488","span":{"begin":12003,"end":12005},"obj":"16133334"},{"id":"25582171-19217543-14905489","span":{"begin":12060,"end":12062},"obj":"19217543"},{"id":"25582171-12784608-14905490","span":{"begin":12099,"end":12101},"obj":"12784608"},{"id":"25582171-18795285-14905491","span":{"begin":12333,"end":12335},"obj":"18795285"},{"id":"25582171-19415268-14905492","span":{"begin":12361,"end":12363},"obj":"19415268"},{"id":"25582171-16233793-14905493","span":{"begin":12406,"end":12408},"obj":"16233793"},{"id":"25582171-1818539-14905494","span":{"begin":14356,"end":14358},"obj":"1818539"},{"id":"25582171-15538555-14905495","span":{"begin":14542,"end":14544},"obj":"15538555"},{"id":"25582171-10099624-14905496","span":{"begin":15680,"end":15682},"obj":"10099624"},{"id":"25582171-167775-14905497","span":{"begin":16985,"end":16987},"obj":"167775"},{"id":"25582171-16233009-14905498","span":{"begin":19016,"end":19018},"obj":"16233009"},{"id":"25582171-10893421-14905499","span":{"begin":19504,"end":19506},"obj":"10893421"},{"id":"25582171-12618051-14905500","span":{"begin":19661,"end":19663},"obj":"12618051"},{"id":"25582171-17439233-14905501","span":{"begin":19746,"end":19747},"obj":"17439233"},{"id":"25582171-10463157-14905502","span":{"begin":21798,"end":21800},"obj":"10463157"}],"attributes":[{"subj":"25582171-23541380-14905478","pred":"source","obj":"2_test"},{"subj":"25582171-23541380-14905479","pred":"source","obj":"2_test"},{"subj":"25582171-21365447-14905480","pred":"source","obj":"2_test"},{"subj":"25582171-21365447-14905481","pred":"source","obj":"2_test"},{"subj":"25582171-23236514-14905482","pred":"source","obj":"2_test"},{"subj":"25582171-21336613-14905483","pred":"source","obj":"2_test"},{"subj":"25582171-22124721-14905484","pred":"source","obj":"2_test"},{"subj":"25582171-17439233-14905485","pred":"source","obj":"2_test"},{"subj":"25582171-17439233-14905486","pred":"source","obj":"2_test"},{"subj":"25582171-22174787-14905487","pred":"source","obj":"2_test"},{"subj":"25582171-16133334-14905488","pred":"source","obj":"2_test"},{"subj":"25582171-19217543-14905489","pred":"source","obj":"2_test"},{"subj":"25582171-12784608-14905490","pred":"source","obj":"2_test"},{"subj":"25582171-18795285-14905491","pred":"source","obj":"2_test"},{"subj":"25582171-19415268-14905492","pred":"source","obj":"2_test"},{"subj":"25582171-16233793-14905493","pred":"source","obj":"2_test"},{"subj":"25582171-1818539-14905494","pred":"source","obj":"2_test"},{"subj":"25582171-15538555-14905495","pred":"source","obj":"2_test"},{"subj":"25582171-10099624-14905496","pred":"source","obj":"2_test"},{"subj":"25582171-167775-14905497","pred":"source","obj":"2_test"},{"subj":"25582171-16233009-14905498","pred":"source","obj":"2_test"},{"subj":"25582171-10893421-14905499","pred":"source","obj":"2_test"},{"subj":"25582171-12618051-14905500","pred":"source","obj":"2_test"},{"subj":"25582171-17439233-14905501","pred":"source","obj":"2_test"},{"subj":"25582171-10463157-14905502","pred":"source","obj":"2_test"}]}],"config":{"attribute types":[{"pred":"source","value type":"selection","values":[{"id":"2_test","color":"#e793ec","default":true}]}]}}