园艺植物质子泵及其对有机酸积累调控的研究进展

doi:10.16420/j.issn.0513-353x.2021-0936

摘要/Abstract

摘要：

质子泵是一类促使质子（H⁺）跨膜运输造成膜内外电势差而参与多种生命进程的重要酶类。质子泵促进质子跨膜运输，同时伴随着能量的释放，可以为有机酸等物质跨膜运输贮藏于液泡中提供动力。本文中重点阐述园艺植物中与酸转运相关的质子泵类型及其结构特征和调控因素，以及不同类型质子泵对有机酸积累的调控作用，以期为园艺植物质子泵及其在有机酸积累方面的调控研究提供参考，进而为园艺植物的品质调控提供理论依据。

关键词: 园艺植物, 有机酸, 质子泵, 果实, 品质调控

Abstract:

Proton pumps are types of important enzymes that promote proton（H⁺）transmembrane transport and cause potential difference between the membrane and participate in a variety of life processes. Organic acids are mainly stored in vacuoles and proton pump promotes proton transmembrane transport accompanied by energy release，which then provides power for the transmembrane transport and storage of organic acids in vacuoles. This paper focuses on the categories，structural characteristics and regulatory factors of proton pumps related to acid transport in horticultural plants，as well as the research status of different types of proton pumps in the regulation of organic acid accumulation，in order to provide reference for the research and regulation of proton pumps and organic acid accumulation in horticultural plants，and then provide a theoretical basis for regulating the quality of horticultural plants.

Key words: horticultural plant, organic acid, proton pump, fruit, regulating of quality

中图分类号:

S6
Q 945

石彩云, 刘丽, 魏志峰, 高登涛, 刘永忠. 园艺植物质子泵及其对有机酸积累调控的研究进展[J]. 园艺学报, 2022, 49(12): 2611-2621.

SHI Caiyun, LIU Li, WEI Zhifeng, GAO Dengtao, LIU Yongzhong. Research Progress of Proton Pumps and Their Regulation in Organic Acid Accumulation in Horticultural Plants[J]. Acta Horticulturae Sinica, 2022, 49(12): 2611-2621.

图/表 2

图1 质子泵在柠檬酸和苹果酸贮藏液泡中的作用示意图（改编自Guo 等，2016;Zheng 等，2021） OAA：草酰乙酸;ACL：ATP柠檬酸裂解酶;ACO：顺乌头酸酶;GAD：谷氨酸脱羧酶;GABA：γ-氨基丁酸;GS：谷氨酰胺合成酶;PEP：磷酸烯醇式丙酮酸;PEPC：磷酸烯醇式丙酮酸羧化酶;PEPCK：磷酸烯醇式丙酮酸羧激酶;Citrate2-，Citrate3-：柠檬酸根离子;Malate2-：苹果酸根离子;CsCit1：柠檬酸/H+同向转运蛋白;ATP：三磷酸腺苷;Pi：磷酸基团;PPi：焦磷酸;P：磷酸;ADP：二磷酸腺苷;H+：氢离子;NADP-ME：烟酰胺腺嘌呤二核苷酸磷酸-苹果酸酶;ALMT9：铝激活苹果酸转运蛋白;tDT：液泡膜二羧酸转运蛋白;NAD-MDH1：NAD-苹果酸脱氢酶1。

Fig. 1 Role of proton pumps in citric acid and malic acid storage in vacuole（Adapted from Guo et al.，2016;Zheng et al.，2021） OAA：Oxaloacetate;ACL：ATP citrate lyase;ACO：Aconitase;GAD：Glutamate decarboxylase;GABA：γ-aminobutyric acid;GS：Glutamine synthetase;PEP：Phosphoenolpyruvate;PEPC：Phosphoenolpyruvate carboxylase;PEPCK：Phosphoenolpyruvate carboxykinase;Citrate2-，Citrate3-：citrate ion;Malate2-：Malate ion;CsCit1：Citrate/H+ symporter;ATP：Adenosine triphosphate;Pi：Phosphate group;PPi：Pyrophosphate;P：Phosphoric acid;ADP：Adenosine diphosphate;H+：Hydrogen ion;NADP-ME：Nicotinamide adenine dinucleotide phosphate-malic enzyme;ALMT9：AL-activated malate transporter 9;tDT：Tonoplast dicarboxylate transporter;NAD-MDH1：Nicotinamide adenine dinucleotide-malate dehydrogenase 1.

表1 园艺植物中已报道活性或基因数量的质子泵

Table 1 Types and gene number of proton pumps reported in horticultural plants

园艺作物 Horticultural crop	基因数或活性 Gene number or enzyme activity			参考文献Reference
园艺作物 Horticultural crop	PHA	VHA	VHP	参考文献Reference
柑橘Citrus	8	20	3	Shi et al.，2015，2018;Hussain et al.，2020
苹果Apple	15	48	1	Yao et al.，2011;Dong et al.，2013;Hu et al.，2015;Ma et al.，2020;Zhou et al.，2020
梨Pear	11 ~ 16	43	活性Activity	Suzuki et al.，1999;Amemiya，2005;Zhang et al.，2020;Zhou et al.，2020
桃Peach	9	23	2	Etienne et al.，2002a;Zhang et al.，2020;Zhou et al.，2020
李Plum	0	25	0	Zhou et al.，2020
杏Apricot	9	0	0	Zhang et al.，2020
葡萄Grape	活性Activity	活性Activity	2	Terrier et al.，2001;Venter et al.，2006;Liu et al.，2008
枇杷Loquat	0	2	1	Yang et al.，2011，2021;李大培等，2019
核桃Walnut	0	1	0	Xu et al.，2017
草莓Strawberry	7	20	0	Zhang et al.，2020;Zhou et al.，2020
番茄Tomato	7/8	23	3	Ewing & Bennett，1994;Coker et al.，2003;Mohammed et al.，2012;
黄瓜Cucumber	10	2	1	Kaba?a et al.，2014;Wdowikowska & Klobus，2016
南瓜Pumpkin	0	0	1	Maruyama et al.，1998
番木瓜Papaya	活性Activity	0	0	Azevedo et al.，2008
矮牵牛Petunia	2	0	0	Veiweij et al.，2008;Faraco et al.，2014
白花马蔺White flower iris	0	1	0	Wang et al.，2016
冰叶日中花Ice plant	0	1	0	Tsiantis et al.，1996

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