Research Progress of Citric Acid Metabolism in the Fruit

doi:10.16420/j.issn.0513-353x.2021-0709

Acta Horticulturae Sinica ›› 2022, Vol. 49 ›› Issue (12): 2579-2596.doi: 10.16420/j.issn.0513-353x.2021-0709

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Research Progress of Citric Acid Metabolism in the Fruit

ZHAO Yong, ZHU Hongju, YANG Dongdong, GONG Chengsheng, LIU Wenge()

Zhengzhou Fruit Research Institute，Chinese Academy of Agricultural Sciences，Zhengzhou 450009，China

Received:2022-04-08 Revised:2022-06-13 Online:2022-12-25 Published:2023-01-02
Contact: LIU Wenge E-mail:liuwenge@caas.cn

Abstract

Abstract:

Citric acid is one of the important organic acids in the Horticultural crop fruit. Citric acid significantly affects the taste and flavor of the fruit. This paper reviews the status and function of citric acid synthesis，transport，storage and utilization in the accumulation of fruit. This paper emphasized the influences of the proton transfer of proton pump in the tonoplast and the regulation of citric acid accumulation. In addition，it gives an overview to the status of transcription factors，environmental and climatic factors，cultivation management measures and postharvest treatments in the metabolism and accumulation of citric acid. Based on the comprehensive understanding of the metabolic mechanism of citric acid synthesis and accumulation，the accumulation of citric acid in the fruit can be finely regulated by gene editing，to obtain the fruit with more suitable sugar-to-acid ratio and better taste.

Key words: citric acid, transport protein, metabolism, fruit acidity

CLC Number:

ZHAO Yong, ZHU Hongju, YANG Dongdong, GONG Chengsheng, LIU Wenge. Research Progress of Citric Acid Metabolism in the Fruit[J]. Acta Horticulturae Sinica, 2022, 49(12): 2579-2596.

Figures/Tables 4

Fig. 1 Schematic diagram of synthesis，storage and metabolism of citric acid in pulp cells The purple font represents the enzymes in the reaction process. The red font represents the intermediate product of the TCA cycle process. The brown font represents the carrier of transmembrane transport. The large arrows of the double arrows represent the main direction of the reversible reaction，and the small arrows of the double arrows represent the secondary direction of the reversible reaction. The blue dotted arrows represent the direction of product formation after multiple reactions. The dotted lines of double arrows represent materials transported for carriage of the transmembrane transport. The pink double ellipses represent transmembrane channels or carriers. The blue circles represent the proton pump. CS：citrate synthase;ACO：aconitase;NAD-IDH：NAD+-isocitrate dehydrogenase;NADP-IDH：NADP+-isocitrate dehydrogenase;NAD-a-KGDH：NAD+-a-oxoglutarate dehydrogenase;STK：succinic thiokinase;SDH：succinic dehydrogenase;FH：fumarase;NAD-MDH：NAD+-Malate Dehydrogenase;NAD-ME：NAD+-malic enzyme;NADP-ME：NADP+-malic enzyme;PDH：pyruvate dehydrogenase;PPDK：pyruvate orthophosphate dikinase;PEPC：phosphoenolpyruvate carboxylase;PEPCK：phosphoenolpyruvate carboxykinase;ACL：ATP-citrate lyase;GAD：glutamate decarboxylase;GS：glutamine synthetase. （Shimada et al.，2006;Sweetlove et al.，2010;Etienne et al.，2013;Guo et al.，2016;Sadka et al.，2019）.

Fig. 2 Schematic diagram of the TCA cycle process and its intermediate metabolites embedded in other metabolic networks The blue font represents the enzymes in the TCA cycle reaction process. The arrows outside the TCA cycle network point to the type of the consumption or the production of intermediate metabolites in the TCA cycle. The intermediate metabolites of the TCA cycle pathway can be used as a carbon skeleton to participate in other metabolic pathways，such as the citric acid firstly via pathway 1，and finally enter into the secondary metabolite synthesis or the fatty acid extension metabolism of the mevalonate pathway. The malic acid can be generated by the glyoxylic acid cycle of the third pathway. The fumaric acid can be generated by the pathway of the amino acid synthesis of the fourth pathway and the pathway of the purine metabolism of the fifth pathway. The OAA participates in the pathway of the amino acid synthesis and decomposition of the second pathway. The 2-OG can participate in the pathway of the nitrogen assimilation of the sixth and seventh pathways，and can also participate in the synthesis and decomposition of amino acids by the pathway from the eighth to eighteenth. （Sweetlove et al.，2010）

Fig. 3 Major carboxylic acid transport proteins in the mitochondria The green triangle represents the electrochemical gradients of protons that drives the transport of the organic acids between the cytoplasm and mitochondria. The arrows represent the direction of material transport. The pink arrow represent the direction of citrate transport. The double ellipses represent transmembrane channels or carriers. PiC：phosphate carrier;PTP：pyruvate transport protein;DTP：dicarboxylate transport protein;CTP：citrate transport protein;DTC：dicarboxylate-tricarboxylate carrier;OMT：oxoglutarate-malate transporter;OAT：oxaloacetate-malate transporter. （Laloi，1999;Etienne et al.，2002，2013;Picault et al.，2002）.

Fig. 4 Citrate homeostasis in the vacuole of citrus juice sac cells The pink double ellipses represent transmembrane channels or carriers. The blue circles represent the proton pump. VHA：vacuolar H+-ATPase;VHP：Vacuolar H+-pyrophosphatase;PH：Plasma membrane H+-ATPase. （Guo et al.，2016;Shimada et al.，2006）.

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Research Progress of Citric Acid Metabolism in the Fruit

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