Kiefer Patrick, Heinzle Elmar, Zelder Oskar, Wittmann Christoph
Biochemical Engineering, Saarland University, Saarbrucken, Germany.
Appl Environ Microbiol. 2004 Jan;70(1):229-39. doi: 10.1128/AEM.70.1.229-239.2004.
A comprehensive approach to (13)C tracer studies, labeling measurements by gas chromatography-mass spectrometry, metabolite balancing, and isotopomer modeling, was applied for comparative metabolic network analysis of lysine-producing Corynebacterium glutamicum on glucose or fructose. Significantly reduced yields of lysine and biomass and enhanced formation of dihydroxyacetone, glycerol, and lactate in comparison to those for glucose resulted on fructose. Metabolic flux analysis revealed drastic differences in intracellular flux depending on the carbon source applied. On fructose, flux through the pentose phosphate pathway (PPP) was only 14.4% of the total substrate uptake flux and therefore markedly decreased compared to that for glucose (62.0%). This result is due mainly to (i) the predominance of phosphoenolpyruvate-dependent phosphotransferase systems for fructose uptake (PTS(Fructose)) (92.3%), resulting in a major entry of fructose via fructose 1,6-bisphosphate, and (ii) the inactivity of fructose 1,6-bisphosphatase (0.0%). The uptake of fructose during flux via PTS(Mannose) was only 7.7%. In glucose-grown cells, the flux through pyruvate dehydrogenase (70.9%) was much less than that in fructose-grown cells (95.2%). Accordingly, flux through the tricarboxylic acid cycle was decreased on glucose. Normalized to that for glucose uptake, the supply of NADPH during flux was only 112.4% on fructose compared to 176.9% on glucose, which might explain the substantially lower lysine yield of C. glutamicum on fructose. Balancing NADPH levels even revealed an apparent deficiency of NADPH on fructose, which is probably overcome by in vivo activity of malic enzyme. Based on these results, potential targets could be identified for optimization of lysine production by C. glutamicum on fructose, involving (i) modification of flux through the two PTS for fructose uptake, (ii) amplification of fructose 1,6-bisphosphatase to increase flux through the PPP, and (iii) knockout of a not-yet-annotated gene encoding dihydroxyacetone phosphatase or kinase activity to suppress overflow metabolism. Statistical evaluation revealed high precision of the estimates of flux, so the observed differences for metabolic flux are clearly substrate specific.
采用一种综合方法,即(13)C示踪研究、气相色谱 - 质谱法进行标记测量、代谢物平衡以及同位素异构体建模,对在葡萄糖或果糖上生产赖氨酸的谷氨酸棒杆菌进行比较代谢网络分析。与葡萄糖相比,果糖上赖氨酸和生物量的产量显著降低,二羟基丙酮、甘油和乳酸的生成增加。代谢通量分析表明,根据所应用的碳源不同,细胞内通量存在显著差异。在果糖上,通过戊糖磷酸途径(PPP)的通量仅占总底物摄取通量的14.4%,因此与葡萄糖相比(62.0%)明显降低。这一结果主要归因于:(i)果糖摄取依赖磷酸烯醇丙酮酸的磷酸转移酶系统(PTS(Fructose))占主导(92.3%),导致果糖主要通过1,6 - 二磷酸果糖进入细胞;(ii)果糖1,6 - 双磷酸酶无活性(0.0%)。通过PTS(甘露糖)途径摄取果糖的通量仅为7.7%。在以葡萄糖生长的细胞中,通过丙酮酸脱氢酶的通量(70.9%)远低于以果糖生长的细胞(95.2%)。因此,在葡萄糖上三羧酸循环的通量降低。相对于葡萄糖摄取量进行归一化后,果糖通量期间NADPH的供应量仅为112.4%,而葡萄糖上为176.9%,这可能解释了谷氨酸棒杆菌在果糖上赖氨酸产量显著较低的原因。平衡NADPH水平甚至揭示了果糖上NADPH明显不足,这可能通过苹果酸酶的体内活性得以克服。基于这些结果,可以确定潜在靶点以优化谷氨酸棒杆菌在果糖上的赖氨酸生产,包括:(i)改变通过两种果糖摄取PTS的通量;(ii)扩增果糖1,6 - 双磷酸酶以增加通过PPP的通量;(iii)敲除一个尚未注释的编码二羟基丙酮磷酸酶或激酶活性的基因以抑制溢流代谢。统计评估显示通量估计具有高精度,因此观察到的代谢通量差异明显是底物特异性的。