果实抗坏血酸生物合成、代谢循环及其调控的研究进展

doi:10.16420/j.issn.0513-353x.2022-0490

园艺学报 ›› 2023, Vol. 50 ›› Issue (9): 1899-1915.doi: 10.16420/j.issn.0513-353x.2022-0490

果实抗坏血酸生物合成、代谢循环及其调控的研究进展

何义仲¹^,^*, 庞尧²^,^*, 孙浩谦², 李欣宇², 王振豪², 钱卫², 张印², 何发³, 尹杭¹, 赖恒鑫¹, 淳长品¹, 付行政¹, 彭良志¹^,^**()

¹ 西南大学/中国农业科学院柑桔研究所，国家柑桔工程技术研究中心，重庆 400712
² 西南大学园艺园林学院，重庆 400715
³ 南充市农业科学院，四川南充 637000

收稿日期:2023-04-02 修回日期:2023-08-13 出版日期:2023-09-25 发布日期:2023-09-26
通讯作者:
^**（E-mail：pengliangzhi@cric.cn）
作者简介:
^* 共同第一作者
基金资助:
国家自然科学基金项目(31902084); 重庆市自然科学基金项目(CSTB2022NSCQ-MSX1499); 现代农业产业技术体系建设专项; 国家重点研发计划项目(2020YFD1000102); 重庆市级大学生创新创业训练计划项目(S202110635085); 重庆市级大学生创新创业训练计划项目(S202210635141)

Recent Advances in the Biosynthesis，Recycling and Regulation of Ascorbic Acid in Fruit

HE Yizhong¹, PANG Yao², SUN Haoqian², LI Xinyu², WANG Zhenhao², QIAN Wei², ZHANG Yin², HE Fa³, YIN Hang¹, LAI Hengxin¹, CHUN Changpin¹, FU Xingzheng¹, PENG Liangzhi¹^,^**()

¹ National Citrus Engineering Research Center，Citrus Research Institute，Southwest University/Chinese Academy of Agricultural Sciences，Chongqing 400712，China
² College of Horticulture and Forestry，Southwest University，Chongqing 400715，China
³ Nanchong Academy of Agricultural Sciences，Nanchong，Sichuan 637000，China

Received:2023-04-02 Revised:2023-08-13 Published:2023-09-25 Online:2023-09-26
Contact: ^**（E-mail：pengliangzhi@cric.cn）

摘要/Abstract

摘要：

对不同果实抗坏血酸（AsA，维生素C）积累特点、生物学功能和合成途径等进行了综述，分析了AsA受遗传与外在因素如光照、糖和激素等影响的共性问题，阐述转录因子、microRNA、DNA甲基化和翻译后修饰方式调控果实AsA合成与循环途径相关基因的模式，以期为后续果实AsA形成机制研究提供参考。

关键词: 果实, 生物学功能, 生物合成与再生, 调控机制

Abstract:

The biological function，biosynthesis and accumulation characteristics of ascorbic acid（AsA or vitamin C）in fruit germplasm were summarized. It addresses such questions as the both genetic and environmental factors that affect its accumulation，such as sugars，hormones，light，and temperature. This study elaborates the regulatory mechanism of AsA biosynthesis and recycling by the transcription factors，microRNAs，DNA methylation and post-translational patterns. The goal is to provide a reference for the formation of fruit AsA in the future.

Key words: fruit, biological function, biosynthesis and regeneration, regulation mechanism

何义仲, 庞尧, 孙浩谦, 李欣宇, 王振豪, 钱卫, 张印, 何发, 尹杭, 赖恒鑫, 淳长品, 付行政, 彭良志. 果实抗坏血酸生物合成、代谢循环及其调控的研究进展[J]. 园艺学报, 2023, 50(9): 1899-1915.

HE Yizhong, PANG Yao, SUN Haoqian, LI Xinyu, WANG Zhenhao, QIAN Wei, ZHANG Yin, HE Fa, YIN Hang, LAI Hengxin, CHUN Changpin, FU Xingzheng, PENG Liangzhi. Recent Advances in the Biosynthesis，Recycling and Regulation of Ascorbic Acid in Fruit[J]. Acta Horticulturae Sinica, 2023, 50(9): 1899-1915.

图/表 3

表1 不同植物材料的果实或其他食用部位的AsA含量

Table 1 The content of AsA in the fruit or other edible tissues of different types of plant materials

科 Family	材料 Material	每100 g或100 mL含量/mg Content per 100 g or 100 mL	参考文献 Reference
禾本科Gramineae	水稻Rice（Foliar）	ND ~ 4.65	Ji et al.,2013
	小麦Wheat	0.54 ~ 0.38	Nicolò et al.,2006
	玉米Maize	10.11 ~ 22.17	杨若明和李玉田,2001
豆科Legume	大豆Soybean	3.52 ~ 7.47	Kumar et al.,2010
茄科Solanaceae	马铃薯Potato	7.54 ~ 28.58	Jiménez et al.,2009
蔷薇科Rosaceae	刺梨Chestnut rose	1 290.00 ~ 2 000.00	Huang et al.,2014
	玫瑰果Rose hip	1 273.64 ~ 2 264.00	韩云和管正学,1991
	金樱子Cherokee rose	700.00 ~ 900.00	佘祥威等,1988
	木瓜Papaya	31.50 ~ 62.50	Farina et al.,2020
	草莓Strawberry	28.80 ~ 88.70	Mezzetti et al.,2016；Gundogdu et al.,2021
	苹果Apple	1.05 ~ 27.85	Mellidou et al.,2012；Fang et al.,2017
	樱桃Cherry	8.40 ~ 17.60	Gündogdu & Bilge,2012；Suekawa et al.,2019
	枇杷Loquat	6.50 ~ 14.80	Cai et al.,2011
	桃Peach	1.00 ~ 14.00	Gil et al.,2002；Cantín et al.,2009
	梨Pear	1.32 ~ 3.59	Curi et al.,2021
	李Plum	3.00 ~ 10.00	Gil et al.,2002
金虎尾科Malpighiaceae	针叶樱桃Acerola	800.00 ~ 4 000.00	Nagy & Shaw,1980
猕猴桃科Actinidiaceae	猕猴桃Kiwifruit	29.00 ~ 2 140.00	Huang et al.,2004；Ichiro et al.,2004；Lin et al.,2022
胡桃科Juglandaceae	核桃Walnut	410.00 ~ 1 800.00	Pyke et al.,1942
胡颓子科Elaeagnaceae	沙棘Sea-buckthorn	10.00 ~ 1 320.00	吕荣森,1990；Kallio et al.,2002
鼠李科Rhamnaceae	枣Chinese jujube	166.47 ~ 808.83	刘孟军和汪民,2009；Zhang et al.,2016
芸香科Rutaceae	柑橘Citrus	15.90 ~ 62.00	Claudie et al.,2005；He et al.,2019
桃金娘科Myrtaceae	番石榴 Guava	136.50 ~ 220.40	Adrees et al.,2010；Singh et al.,2016
柿科Ebenaceae	柿Persimmon	11.66 ~ 12.32	Cardoso et al.,2011
漆树科Anacardiaceae	杧果Mango	17.01 ~ 50.71	Sellamuthu et al.,2013；Vithana et al.,2019
无患子科Sapindaceae Juss	龙眼Longan	44.65 ~ 79.23	Wall,2006b
无患子科Sapindaceae Juss	荔枝Litchi	21.00 ~ 36.00	Wall,2006b
葡萄科Vitaceae	葡萄Grape	12.33 ~ 30.80	Derradji et al.,2014；Alrashdi et al.,2017
酢浆草科Oxalidaceae	杨桃Carambola	17.50 ~ 20.50	Luximon et al.,2003
凤梨Bromeliaceae	菠萝Pineapple	5.08 ~ 33.57	Lu et al.,2014
芭蕉科Musaceae	香蕉Banana	2.10 ~ 18.70	Wall,2006a
番木瓜科 Caricaceae	番木瓜Papaya	36.30 ~ 67.80	Wall,2006a
茄科Solanaceae	番茄Tomato	1.07 ~ 38.80	Bhandari et al.,2016；Alegre et al.,2020

表1 不同植物材料的果实或其他食用部位的AsA含量

Table 1 The content of AsA in the fruit or other edible tissues of different types of plant materials

科 Family	材料 Material	每100 g或100 mL含量/mg Content per 100 g or 100 mL	参考文献 Reference
禾本科Gramineae	水稻Rice（Foliar）	ND ~ 4.65	Ji et al.,2013
	小麦Wheat	0.54 ~ 0.38	Nicolò et al.,2006
	玉米Maize	10.11 ~ 22.17	杨若明和李玉田,2001
豆科Legume	大豆Soybean	3.52 ~ 7.47	Kumar et al.,2010
茄科Solanaceae	马铃薯Potato	7.54 ~ 28.58	Jiménez et al.,2009
蔷薇科Rosaceae	刺梨Chestnut rose	1 290.00 ~ 2 000.00	Huang et al.,2014
	玫瑰果Rose hip	1 273.64 ~ 2 264.00	韩云和管正学,1991
	金樱子Cherokee rose	700.00 ~ 900.00	佘祥威等,1988
	木瓜Papaya	31.50 ~ 62.50	Farina et al.,2020
	草莓Strawberry	28.80 ~ 88.70	Mezzetti et al.,2016；Gundogdu et al.,2021
	苹果Apple	1.05 ~ 27.85	Mellidou et al.,2012；Fang et al.,2017
	樱桃Cherry	8.40 ~ 17.60	Gündogdu & Bilge,2012；Suekawa et al.,2019
	枇杷Loquat	6.50 ~ 14.80	Cai et al.,2011
	桃Peach	1.00 ~ 14.00	Gil et al.,2002；Cantín et al.,2009
	梨Pear	1.32 ~ 3.59	Curi et al.,2021
	李Plum	3.00 ~ 10.00	Gil et al.,2002
金虎尾科Malpighiaceae	针叶樱桃Acerola	800.00 ~ 4 000.00	Nagy & Shaw,1980
猕猴桃科Actinidiaceae	猕猴桃Kiwifruit	29.00 ~ 2 140.00	Huang et al.,2004；Ichiro et al.,2004；Lin et al.,2022
胡桃科Juglandaceae	核桃Walnut	410.00 ~ 1 800.00	Pyke et al.,1942
胡颓子科Elaeagnaceae	沙棘Sea-buckthorn	10.00 ~ 1 320.00	吕荣森,1990；Kallio et al.,2002
鼠李科Rhamnaceae	枣Chinese jujube	166.47 ~ 808.83	刘孟军和汪民,2009；Zhang et al.,2016
芸香科Rutaceae	柑橘Citrus	15.90 ~ 62.00	Claudie et al.,2005；He et al.,2019
桃金娘科Myrtaceae	番石榴 Guava	136.50 ~ 220.40	Adrees et al.,2010；Singh et al.,2016
柿科Ebenaceae	柿Persimmon	11.66 ~ 12.32	Cardoso et al.,2011
漆树科Anacardiaceae	杧果Mango	17.01 ~ 50.71	Sellamuthu et al.,2013；Vithana et al.,2019
无患子科Sapindaceae Juss	龙眼Longan	44.65 ~ 79.23	Wall,2006b
无患子科Sapindaceae Juss	荔枝Litchi	21.00 ~ 36.00	Wall,2006b
葡萄科Vitaceae	葡萄Grape	12.33 ~ 30.80	Derradji et al.,2014；Alrashdi et al.,2017
酢浆草科Oxalidaceae	杨桃Carambola	17.50 ~ 20.50	Luximon et al.,2003
凤梨Bromeliaceae	菠萝Pineapple	5.08 ~ 33.57	Lu et al.,2014
芭蕉科Musaceae	香蕉Banana	2.10 ~ 18.70	Wall,2006a
番木瓜科 Caricaceae	番木瓜Papaya	36.30 ~ 67.80	Wall,2006a
茄科Solanaceae	番茄Tomato	1.07 ~ 38.80	Bhandari et al.,2016；Alegre et al.,2020

图1 AsA的生物合成、代谢循环（A）和细胞内转运（B）途径（刘永立等，2006；Mellidou & Kanellis，2017；Fenech et al.，2019；Foyer et al.，2020；Chaturvedi et al.，2022）。 PGM：磷酸葡萄糖变位酶；UDPGP：UDP-葡萄糖焦磷酸化酶；UDPGDH：UDP-葡萄糖6-脱氢酶；GAUT：半乳糖醛酸转移酶；PME：果胶酯酶；PG：聚半乳糖醛酸酶；GalUR：半乳糖醛酸还原酶；GNL：葡萄糖酸内酯酶；GLDH：L-半乳糖-1,4-内酯脱氢酶；GPI：6-磷酸葡萄糖异构酶；PMI：6-磷酸甘露糖异构酶；PMM：磷酸甘露糖变位酶；VCT1/GMP：GDP-D-甘露糖焦磷酸酶；GME：GDP-D-甘露糖3,5-差向异构酶；VCT2/VCT5/GGP：GDP-L-半乳糖磷酸化酶；VCT4/GPP：L-半乳糖-1-磷酸磷酸酶；GalDH：L-半乳糖脱氢酶；PDE：磷酸二酯酶；SUPH：糖磷酸酶；ALDH：L-古洛糖酸-1,4-内酯脱氢酶；GulLO：L-古洛糖酸内酯氧化酶；PsUSP：UDP-糖焦磷酸化酶；UGT：UDP-葡萄糖醛酸转移酶；GLUK：葡萄糖醛酸激酶；GLUR：葡萄糖醛酸还原酶；MIPS：肌醇3-磷酸合酶；IMP：肌醇单磷酸酶；MIOX：肌醇加氧酶；APX：L-抗坏血酸过氧化物酶；AAO：L-抗坏血酸氧化酶；MDHAR：单脱氢抗坏血酸还原酶；DHAR：脱氢抗坏血酸还原酶；GSHR：谷胱甘肽还原酶。

Fig. 1 AsA biosynthesis，regeneration（A）and intracellular transport（B） PGM：Phosphoglucomutase；UDPGP：UDP-glucose pyrophosphorylase；UDPGDH：UDP-glucose 6-dehydrogenase；GAUT：Galacturonosyltransferase；PME：Pectinesterase；PG：Polygalacturonase；GalUR：Galacturonic acid reductase；GNL：Gluconolactonase；GLDH：L-galactono-1,4-lactone dehydrogenase；GPI：Glucose-6-phosphate isomerase；PMI：Mannose-6-phosphate isomerase；PMM：Phosphomannomutase；VCT1/GMP：GDP-D-mannose pyrophorylase；GME：GDP-D-mannose 3,5-epimerase；VCT2/VCT5/GGP：GDP-L-galactose phosphorylase；VCT4/GPP：L-galactose-1-phosphate phosphatase；GalDH：L-galactose dehydrogenase；PDE：Phosphodiesterase；SUPH：Sugar-phosphatase；ALDH：L-gulono-1,4-lactone dehydrogenase；GulLO：L-gulonolactone oxidase；PsUSP：UDP-sugar pyrophosphorylase；UGT：UDP-glucuronosyltransferase；GLUK：Glucuronokinase；GLUR：Glucuronate reductase；MIPS：myo-Inositol-3-phosphate synthase；IMP：myo-Inositol monophosphatase；MIOX：myo-Inositol oxygenase；APX：L-ascorbate peroxidase；AAO：L-ascorbate oxidase；MDHAR：Monodehydroascorbate reductase；DHAR：Dehydroascorbate reductase；GSHR：Glutathione reductase.

图2 AsA的生物合成与代谢循环途径的调控网络蓝白色椭圆：转录因子；红白色椭圆：翻译后调控因子；棕白色椭圆：小RNA。

Fig. 2 The regulatory network of AsA biosynthesis and regeneration White-blue ellipse：transcription factor；White-red ellipse：post-translational regulation factor；White-blue ellipse：microRNAs. （Zhang et al.，2012；Mellidou & Kanellis，2017；Yu et al.，2021；Liu et al.，2022）.

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果实抗坏血酸生物合成、代谢循环及其调控的研究进展

Recent Advances in the Biosynthesis，Recycling and Regulation of Ascorbic Acid in Fruit

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果实抗坏血酸生物合成、代谢循环及其调控的研究进展

Recent Advances in the Biosynthesis，Recycling and Regulation of Ascorbic Acid in Fruit

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