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Obligate heterofermentative lactic acid bacteria (LAB) are well-known for their beneficial health effects in humans. To delineate the incompletely characterized metabolism that currently limits their exploitation, at systems-level, we developed a genome-scale metabolic model of the representative obligate heterofermenting LAB , Leuconostoc mesenteroides  ( i LME620). Constraint-based flux analysis was then used to simulate several qualitative and quantitative phenotypes of  L. mesenteroides , thereby evaluating the model validity. With established predictive capabilities, we subsequently employed  i LME620 to elucidate unique metabolic characteristics of  L. mesenteroides , such as the limited ability to utilize amino acids as energy source, and to substantiate the role of malolactic fermentation (MLF) in the reduction of pH-homeostatic burden on F 0 F 1 -ATPase. We also reported new hypothesis on the MLF mechanism that could be explained via a substrate channelling-like phenomenon mainly influenced by intracellular redox state rather than the intermediary reactions. Model simulations further revealed possible proton-symporter dependent activity of the energy efficient glucose-phosphotransferase system in obligate heterofermentative LAB. Moreover, integrated transcriptomic analysis allowed us to hypothesize transcriptional regulatory bias affecting the intracellular redox state. The insights gained here about the low ATP-yielding metabolism of  L. mesenteroides , dominantly controlled by the cellular redox state, could potentially aid strain design for probiotic and cell factory applications.

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Genome-scale modeling and transcriptome analysis of Leuconostoc mesenteroides unravel the redox governed metabolic states in obligate heterofermentative lactic acid bacteria