园艺作物抗坏血酸生物合成中光的调控作用研究综述

doi:10.16420/j.issn.0513-353x.2022-0486

园艺学报 ›› 2022, Vol. 49 ›› Issue (11): 2502-2518.doi: 10.16420/j.issn.0513-353x.2022-0486

园艺作物抗坏血酸生物合成中光的调控作用研究综述

彭银霞¹, 张颖¹, 朱康友¹, 孙鑫³, 张克敏⁴, 孙周平¹, 齐明芳¹, 李天来¹^,², 王峰¹^,²^,^*()

¹沈阳农业大学园艺学院，沈阳 110866
²设施园艺教育部重点实验室，北方园艺设施设计与应用技术国家地方联合工程研究中心（辽宁），沈阳 110866
³沈阳农业大学土地与环境学院，沈阳 110866
⁴河北省辛集市植保植检站，河北辛集 052360

收稿日期:2022-06-09 修回日期:2020-09-29 出版日期:2022-11-25 发布日期:2022-11-25
通讯作者: 王峰 E-mail:fengwang@syau.edu.cn
基金资助:
国家重点研发计划项目(2019YFD1000300);国家自然科学基金项目(32122081);国家自然科学基金项目(32272698);辽宁省自然科学基金优秀青年基金项目;国家重点研发计划项目(2018YFD1000800);沈阳市中青年科技创新人才支持计划项目(RC200449);国家现代农业产业技术体系建设专项资金项目(CARS-23)

Light Regulation of Ascorbic Acid Biosynthesis in Horticultural Crops

PENG Yinxia¹, ZHANG Ying¹, ZHU Kangyou¹, SUN Xin³, ZHANG Kemin⁴, SUN Zhouping¹, QI Mingfang¹, LI Tianlai¹^,², WANG Feng¹^,²^,^*()

¹College of Horticulture，Shenyang Agricultural University，Shenyang 110866，China
²Key Laboratory of Protected Horticulture，Ministry of Education，Local Joint Engineering Research Center of Northern Horticultural Facilities Design & Application Technology（Liaoning），Shenyang 110866，China
³College of Land and Environment，Shenyang Agricultural University，Shenyang 110866，China
⁴Plant Protection and Inspection Station，Xinji，Hebei 052360，China

Received:2022-06-09 Revised:2020-09-29 Online:2022-11-25 Published:2022-11-25
Contact: WANG Feng E-mail:fengwang@syau.edu.cn

摘要/Abstract

摘要：

总结了光对园艺作物抗坏血酸生物合成与代谢的调控作用，解析了光合作用和呼吸作用间的相互作用对抗坏血酸合成和代谢的影响，重点阐明了光调节关键因子对园艺作物抗坏血酸生物合成的调控机制和信号网络，以期为解析光调控植物抗坏血酸生物合成和代谢的分子机制及调控网络奠定理论基础，为利用基因工程和LED补光技术增加园艺作物抗坏血酸含量和培育功能性园艺产品提供理论和实践指导。

关键词: 光, 抗坏血酸, 园艺作物, 光合作用, 呼吸作用

Abstract:

This review summarized the regulation of light on ascorbic acid biosynthesis and metabolism in horticultural plants，analyzed the interaction between photosynthesis and respiration on ascorbic acid biosynthesis and metabolism，and focused on elucidating the regulation mechanism and signaling network of key light-regulated factors on ascorbic acid biosynthesis in horticultural plants. It is expected to lay a theoretical foundation for analyzing the molecular mechanism and regulatory network of light regulation of plant ascorbic acid biosynthesis and metabolism，and provide theoretical and practical guidance for increasing the content of ascorbic acid inhorticultural crops，and cultivating functional horticultural products by using genetic engineering and LED light supplement technology.

Key words: light, ascorbate acid, horticultural crops, photosynthesis, respiration

中图分类号:

Q945.1

彭银霞, 张颖, 朱康友, 孙鑫, 张克敏, 孙周平, 齐明芳, 李天来, 王峰. 园艺作物抗坏血酸生物合成中光的调控作用研究综述[J]. 园艺学报, 2022, 49(11): 2502-2518.

PENG Yinxia, ZHANG Ying, ZHU Kangyou, SUN Xin, ZHANG Kemin, SUN Zhouping, QI Mingfang, LI Tianlai, WANG Feng. Light Regulation of Ascorbic Acid Biosynthesis in Horticultural Crops[J]. Acta Horticulturae Sinica, 2022, 49(11): 2502-2518.

图/表 3

图1 植物抗坏血酸生物合成与代谢途径 HXK：己糖激酶;PGI：磷酸葡萄糖异构酶;PMI：磷酸甘露糖异构化;PMM：磷酸甘露糖突变酶;GMP：GDP-甘露糖焦磷酸化酶;GME：GDP-甘露糖-3,5-差向异构酶;GGP：GDP-L-半乳糖磷酸化酶;GPP：L-半乳糖-1-磷酸磷酸酶;GalDH：L-半乳糖脱氢酶;GLDH：L-半乳糖-1,4-内酯脱氢酶;GalUR：半乳糖醛酸还原酶;MIOX：肌醇加氧酶;APX：抗坏血酸过氧化物酶;MDAR：单脱氢抗坏血酸还原酶;AO：抗坏血酸氧化酶;GR：谷胱甘肽还原酶;DHAR：脱氢抗坏血酸还原酶;GSH：谷胱甘肽;GSSG：氧化型谷胱甘肽。数字代表AsA合成及代谢步骤。

Fig. 1 Biosynthetic and metabolic pathway of ascorbic acid in plants HXK：hexokinase;PGI：phosphoglucose isomerase;PMI：phosphomannose isomerase;PMM：phosphomannomutase;GMP：GDP-D-mannose pyrophosphorylase;GME：GDP-D-mannose 3’,5’-epimerase;GGP：GDP-L-galactose phosphorylase;GPP：L-galactose-1-P phosphatase;GalDH：L-galactose dehydrogenase;GLDH：L-galactono-1,4-lactone dehydrogenase;GalUR：D-galacturonate reductase;MIOX：myo-inositol oxygenase;APX：ascorbate peroxidase;MDAR：monodehydroascorbate reductase;AO：ascorbate oxidase;GR：glutathione reductase;DHAR：dehydroascorbate reductase;GSH：glutathione;GSSG：oxidized glutathione. Numbers represent AsA synthetic and metabolic steps.

表1 植物抗环血酸生物合成与代谢途径相关基因

Table 1 Ascorbic acid biosynthesis and metabolic pathway related genes in plants

编号 Code	酶 Enzyme	缩写 Abbreviation	基因编号 ccession number	参考文献 Reference
1	己糖激酶Hexokinase	HXK	At1g50460，At3g20040，At4g29130，At2g19860，At1g47840	2009
2	磷酸葡萄糖异构酶Phosphoglucoisomerase	PGI	At5g42740，At4g24620	2009
3	磷酸甘露糖异构化Phosphomannose isomerise	PMI	At1g67070，At3g02570	2008
4	磷酸甘露糖突变酶Phosphomannomutase	PMM	At2g45790	2008
5	GDP-甘露糖焦磷酸化酶 GDP-mannose pyrophosphorylase	GMP/VTC1	At2g39770，At3g55590，At4g30570，At1g74910	2000
6	GDP-甘露糖-3,5-差向异构酶 GME GDP-mannose-3,5-epimerase	GME	At5g28840	2007
7	GDP-L-半乳糖磷酸化酶 GDP-L-galactose phosphorylase	GGP/VTC2/ VTC5	At4g26850，At5g55120	Wolucka & van 2003
8	L-半乳糖-1-磷酸磷酸酶 L-Galactose-1-phosphate phosphatase	GPP/VTC4	At3g02870	2009
9	L-半乳糖脱氢酶L-Galactose dehydrogenase	GalDH	At4g33670	1998
10	L-半乳糖-1,4-内酯脱氢酶 L-Galactono-1,4-lactone dehydrogenase	GLDH	At3g47930	2005
11	半乳糖醛酸还原酶L-galactose dehydrogenase	GaIUR	At4g33670	2012
12	肌醇加氧酶Inositol oxygenase 4	MIOX4	At4g26260	2012
13	单脱氢抗坏血酸还原酶 Monodehydro ascorbate reductase	MDAR	At1g63940，At3g52880，At3g27820，At5g03630，At3g09940	2003
14	抗坏血酸过氧化物酶Ascorbate peroxidase	APX	At1g07890，At1g77490，At3g09640，At4g32320，At4g35970，At4g09010 At4g35000，At4g08390	2011
15	抗坏血酸氧化酶Ascorbate Oxidase	AO	At4g39830，At5g21105，At5g21100，At5g48450	1984
16	脱氢抗坏血酸还原酶 Dehydro ascorbate reductase	DHAR	At1g19570，At1g19550，At1g75270，At5g16710，At5g36270	2006
17	谷胱甘肽还原酶Glutathione reductase	GR	At3g24170，At3g54660	1998

图2 光信号调控植物抗坏血酸合成的信号网络 ERF98：乙烯响应因子98;CSN5B：COP9 signalosome subunit 5B;CSN8：COP9 signaling complex subunit 8;ZF3：Cys2/His2型锌蛋白;KJC1/2：核苷糖焦磷酸蛋白;VTC1：GDP-D-甘露糖焦磷酸化酶;VTC2：GDP-L-半乳糖磷酸化酶;HZ24：HD-ZIP I;ABI4：脱落酸不敏感4;HY5：光信号转录因子;CML10：钙调蛋白;AMR1：抗坏血酸甘露糖途径调节剂1;AMR1L1：AMR1类似蛋白;GBF3：GBF3转录因子;MYBS1：MYBS转录因子;bHLH59：碱性螺旋—环—螺旋蛋白59;NFYA10：血红素激活蛋白;PMM：磷酸甘露糖突变酶;VTC4：L-半乳糖-1-磷酸磷酸酶;GLDH：L-半乳糖-1,4-内酯脱氢酶;GME：GDP-甘露糖-3’,5’差向异构酶;GGP：GDP-L-半乳糖磷酸化酶;D-mal/L-gal pathway：D-甘露糖和L-半乳糖途径;

Fig. 2 Signal networks of light signaling regulation of ascorbic acid biosynthesis in plants ERF98：Ethylene response factor 98;CSN5B：COP9 signalosome subunit 5B;CSN8：COP9 signaling complex subunit 8;ZF3：Cys2/His2-type zinc-finger 3;KJC1/2：KONJAC1/2;VTC：L-galactose-1-phosphate phosphatase，GPP/VTC;HZ24：HD-ZIP I;ABI4：Abscisic acid insensitive 4;HY5：Long hypocotyl 5;CML10：Calmodulin-like 10;AMR1：Ascorbate mannose pathway modulator 1;AMR1L1：Ascorbate mannose pathway modulator 1 like 1;GBF3：G-box binding factor 3;MYBS1：myeloblastosis（MYB）S1;bHLH59：Basic helix-loop-helix protein 59;NFYA10：Nuclear factor Y-box 10;PMM：Phosphomannomutase;VTC4：L-galactose-1-phosphate phosphatase;GLDH：L-galactose-1,4-lactone dehydrogenase;GME：GDP-D-mannose 3’,5’-epimerase;GGP，GDP-L-galactose phosphorylase.

参考文献 105

[1]	Agius F, Amaya I, Botella M A, Valpuesta V. 2005. Functional analysis of homologous and heterologous promoters in strawberry fruits using transient expression. Journal of Experimental Botany, 56 (409):37-46. doi: 10.1093/jxb/eri004 pmid: 15533885
[2]	Agius F, González-Lamothe R, Caballero J L, Muñoz-Blanco J, Botella M A, Valpuesta V. 2003. Engineering increased vitamin C levels in plants by overexpression of a D-galacturonic acid reductase. Nature Biotechnology, 21 (2):177-181. doi: 10.1038/nbt777 pmid: 12524550
[3]	Badejo A A, Wada K, Gao Y, Maruta T, Sawa Y, Shigeoka S, Ishikawa T. 2012. Translocation and the alternative D-galacturonate pathway contribute to increasing the Ascorbate level in ripening tomato fruits together with the D-mannose/L-galactose pathway. Journal of Experimental Botany, 63 (1):229-239. doi: 10.1093/jxb/err275 pmid: 21984649
[4]	Bartoli C G, Pastori G M, Foyer C H. 2000. Ascorbate biosynthesis in mitochondria is linked to the electron transport chain between complexes III and IV. Plant Physiology, 123 (1):335-344. pmid: 10806250
[5]	Bartoli C G, Tambussi E A, Diego F, Foyer C H. 2009. Control of ascorbic acid synthesis and accumulation and glutathione by the incident light red/far red ratio in Phaseolus vulgaris leaves. FEBS Letters, 583 (1):118-122. doi: 10.1016/j.febslet.2008.11.034 pmid: 19059408
[6]	Bartoli C G, Yu J, Gomez F, Fernández L, McIntosh L, Foyer C H. 2006. Interrelationships between light and respiration in the control of ascorbic acid synthesis and accumulation in Arabidopsis thaliana leaves. Journal of Experimental Botany, 57 (8):1621-1631. pmid: 16714304
[7]	Bienger I, Schopfer P. 1970. Photomodulation by phytochrome of the rate of accumulation of ascorbic acid in mustard seedlings(Sinapis alba L.). Planta, 93 (2):152-159. doi: 10.1007/BF00387122 pmid: 24496710
[8]	Bulley M, Rassam M, Hoser D, Otto W, Schünemann N, Wright M, MacRae E, Gleave A, Laing W. 2009. Gene expression studies in kiwifruit and gene over-expression in Arabidopsis indicates that GDP-L-galactose guanyl transferase is a major control point of vitamin C biosynthesis. Journal of Experimental Botany, 60 (3):765-778. doi: 10.1093/jxb/ern327 pmid: 19129165
[9]	Bulley S, Wright M, Rommens C, Yan H, Rassam M, Wang K L, Andre C, Brewster D, Karunairetnam S, Allan A C, Laing W A. 2012. Enhancing ascorbate in fruits and tubers through over-expression of the L-galactose pathway gene GDP-L-galactose phosphorylase. Plant Biotechnology Journal, 10 (4):390-397. doi: 10.1111/j.1467-7652.2011.00668.x pmid: 22129455
[10]	Chen W, Hu T, Ye J, Wang B, Liu G Z, Wang Y, Yuan L, Li J M, Li F M, Ye Z B, Zhang Y Y. 2020. A CCAAT-binding factor,SlNFYA10,negatively regulates ascorbate accumulation by modulating the D-mannose/L-galactose pathway in tomato. Horticulture Research, 17 (1):200.
[11]	Chen Z, Young T E, Ling J, Chang S, Gallie D R. 2003. Increasing vitamin C content of plants through enhanced ascorbate recycling. Proceedings of the National Academy of Sciences of the United States of America, 100 (6):3525-3530.
[12]	Chew O, Whelan J, Millar A H. 2003. Molecular definition of the Ascorbate-glutathione cycle in Arabidopsis mitochondria reveals dual targeting of antioxidant defenses in plants. Journal of Biological Chemistry, 278 (47):46869-46877. doi: 10.1074/jbc.M307525200 URL
[13]	Cho K M, Nguyen H T K, Kim S Y, Shin J S, Cho D H, Hong S B, Shin J S, Ok S H. 2015. CML10,a variant of calmodulin,modulates ascorbic acid synthesis. New Phytologist, 209 (2):664-678. doi: 10.1111/nph.13612 URL
[14]	Conklin P L, DePaolo D, Wintle B, Schatz C, Buckenmeyer G. 2013. Identification of Arabidopsis VTC 3 as a putative and unique dual function protein kinase:protein phosphatase involved in the regulation of the ascorbic acid pool in plants. Journal of Experimental Botany, 64 (10):2793-2804. doi: 10.1093/jxb/ert140 URL
[15]	Conklin P L, Pallanca J E, Last R L, Smirnoff N. 1997. L-ascorbic acid metabolism in the ascorbate-deficient Arabidopsis mutant vtc7. Plant Physiology, 115:1277-1285. pmid: 9390448
[16]	Dowdle J, Ishikawa T, Gatzek S, Rolinski S, Smirnoff N. 2007. Two genes in Arabidopsis thaliana encoding GDP-L-galactose phosphorylase are required for ascorbate biosynthesis and seedling viability. The Plant Journal, 52 (4):673-689. doi: 10.1111/j.1365-313X.2007.03266.x URL
[17]	Enfissi E M, Barneche F, Ahmed I, Lichtlé C, Gerrish C, McQuinn R P, Giovannoni J J, Lopez-Juez E, Bowler C, Bramley P M, Fraser P D. 2010. Integrative transcript and metabolite analysis of nutritionally enhanced DE-ETIOLATED 1 downregulated tomato fruit. The Plant Cell, 22 (4):1190-1215. doi: 10.1105/tpc.110.073866 URL
[18]	Gilbert L, Alhagdow M, Nunes-Nesi A, Quemener B, Guillon F, Bouchet B, Faurobert M, Gouble B, Page D, Garcia V, Petit J, Stevens R, Causse M, Fernie A R, Lahaye M, Rothan C, Baldet P. 2009. GDP-d-mannose 3,5-epimerase(GME)plays a key role at the intersection of ascorbate and non-cellulosic cell-wall biosynthesis in tomato. The Plant Journal, 60 (3):499-508. doi: 10.1111/j.1365-313X.2009.03972.x pmid: 19619161
[19]	Green M A, Fry S C. 2005. Vitamin C degradation in plant cells via enzymatic hydrolysis of 4-0-oxalyl-L threonate. Nature, 433:83-87. doi: 10.1038/nature03172 URL
[20]	He Jie, Gu Xiu-rong, Wei Chun-hua, Yang Xiao-zhen, Li Hao, Ma Jian-xiang,Zhang Yong,Yang Jian-qiang,Zhang Xian. 2016. Identification and expression analysis under abiotic stresses of the bHLH transcription factor gene family in watermelon. Acta Horticulturae Sinica, 43 (2):281-294. (in Chinese)
	何洁, 顾秀容, 魏春华, 杨小振, 李好, 马建祥, 张勇, 杨建强, 张显. 2016. 西瓜bHLH转录因子家族基因的鉴定及其在非生物胁迫下的表达分析. 园艺学报, 43 (2):281-294.
[21]	Hemavathi, Upadhyaya C P,Young K E,Akula N,Kim H,Heung J J,Oh O M,Aswath C R,Chun S C,Kim D H,Park S W. 2009. Over-expression of strawberry D-galacturonic acid reductase in potato leads to accumulation of vitamin C with enhanced abiotic stress tolerance. Plant Science, 177 (6):659-667. doi: 10.1016/j.plantsci.2009.08.004 URL
[22]	Hossain M A, Nakano Y, Asada K. 1984. Monodehydroascorbate reductase in spinach chloroplasts and its participation in regeneration of ascorbate for scavenging hydrogen peroxide. Plant and Cell Physiology, 25 (3):385-395.
[23]	Hu T X, Ye J, Tao P W, Li H X, Zhang J H, Zhang Y Y, Ye Z B. 2016. Tomato HD-ZIP I transcription factor,SlHZ24,modulates ascorbate accumulation through positively regulating the D-mannose/L-galactose pathway. The Plant Journal, 85 (1):16-29. doi: 10.1111/tpj.13085 URL
[24]	Hu Tixu. 2015. Cloning and functional identification of genes regulating tomato ascorbic acid synthesis [Ph. D. Dessertation]. Wuhan:Huazhong Agriculture University. (in Chinese)
	胡体旭. 2015. 调控番茄抗坏血酸合成的基因克隆与功能鉴定[博士论文]. 武汉: 华中农业大学.
[25]	Imai T, Ban Y, Terakami S, Yamamoto T, Moriguchi T. 2009. L-Ascorbate biosynthesis in peach:cloning of six L-galactose pathway related genes and their expression during peach fruit development. Physiologia Plantarum, 136 (2):139-149. doi: 10.1111/j.1399-3054.2009.01213.x URL
[26]	Ivanov B N. 2014. Role of ascorbic acid in photosynthesis. Biochemistry, 79 (3):282-289.
[27]	Ivanov B N, Sacksteder C A, Kramer D M, Edwards G E. 2001. Light-induced ascorbate-dependent electron transport and membrane energization in chloroplasts of bundle sheath cells of the C 4 plant maize. Archives of Biochemistry and Biophysics, 385 (1):145-153. pmid: 11361011
[28]	Jain A K, Nessler C L. 2000. Metabolic engineering of an alternative pathway for ascorbic acid biosynthesis in plants. Molecular Breeding, 6 (1):73-78. doi: 10.1023/A:1009680818138 URL
[29]	Jimenez A, Hernandez J A, Río L A D, Sevilla F. 1997. Evidence for the presence of the ascorbate-glutathione cycle in mitochondria and peroxisomes of pea leaves. Plant Physiology, 114 (1):275-284. pmid: 12223704
[30]	Karpinski S, Escobar C, Karpinska B, Creissen G, Mullineaux P M. 1997. Photosynthetic electron transport regulates the expression of cytosolic ascorbate peroxidase genes in Arabidopsis during excess light stress. The Plant Cell, 9 (4):627-640.
[31]	Kerchev P I, Pellny T K, Vivancos P D, Kiddle G, Hedden P, Driscoll S, Vanacker H, Verrier P, Hancock R D, Foyer C H. 2011. The transcription factor ABI 4 Is required for the ascorbic acid-dependent regulation of growth and regulation of jasmonate-dependent defense signaling pathways in Arabidopsis. The Plant Cell, 23 (9):3319-3334. doi: 10.1105/tpc.111.090100 pmid: 21926335
[32]	Kotchoni S O, Larrimore K E, Mukherjee M, Kempinski C F, Barth C. 2009. Alterations in the endogenous ascorbic acid content affect flowering time in Arabidopsis. Plant Physiology, 149:803-815. doi: 10.1104/pp.108.132324 pmid: 19028878
[33]	Laing W A, Wright M A, Cooney J, Bulley S M. 2007. The missing step of the L-galactose pathway of ascorbate biosynthesis in plants,an L-galactose guanyl-transferase,increases leaf ascorbate content. Proceedings of the National Academy of Sciences of the United States of America, 104 (22):9534-9539.
[34]	Laloum T, de Mita S, Gamas P, Baudin M, Niebel A. 2013. CCAAT-box binding transcription factors in plants:Y so many? Trends in Plant Science, 18 (3):157-166.
[35]	Lester G E. 2006. Environmental regulation of human health nutrients(ascorbic acid,β-carotene,and folic acid)in fruits and vegetables. HortScience, 41 (1):59-64. doi: 10.21273/HORTSCI.41.1.59 URL
[36]	Li H, Liu Z W, Wu Z J, Wang Y X, Teng R M, Zhuang J. 2018a. Differentially expressed protein and gene analysis revealed the effects of temperature on changes in ascorbic acid metabolism in harvested tea leaves. Horticulture Research, 5:65. doi: 10.1038/s41438-018-0070-x URL
[37]	Li M J, Ma F W, Liu J, Li J. 2010. Shading the whole vines during young fruit development decreases ascorbate accumulation in kiwi. Physiologia Plantarum, 140 (3):225-237.
[38]	Li M, Ma F W, Shang, P F, Zhang M, Hou C M, Liang D. 2009. Influence of light on ascorbate formation and metabolism in apple fruits. Planta, 230:39-51. doi: 10.1007/s00425-009-0925-3 pmid: 19337748
[39]	Li Qingzhu. 2011. Effect of overexpressing potato dehydroascorbate reductase(DHAR)gene in tomato[M. D. Dessertation]. Tai’an:Shangdong Agriculture University. (in Chinese)
	李青竹. 2011. 马铃薯脱氧抗坏血酸还原酶(DHAR)基因在番茄中超表达效应的应用[硕士论文]. 泰安: 山东农业大学.
[40]	Li Y, Chu Z N, Luo J Y, Zhou Y H, Cai Y J, Lu Y E, Xia J H, Kuang H H, Ye Z B, Ouyang B. 2018b. The C2H2 zinc-finger protein SlZF3 regulates AsA synthesis and salt tolerance by interacting with CSN5B. Plant Biotechnology Journal, 16 (6):1201-1213. doi: 10.1111/pbi.12863 URL
[41]	Liu Jingyu, Teng Ruimin, Li Hui, Liu Hao, Zhuang Jing. 2020. Cloning and expression analysis under abiotic stress of the DHAR gene in Camellia sinensis. Acta Horticulturae Sinica, 47 (5):983-994. (in Chinese)
	刘婧愉, 滕瑞敏, 李辉, 刘昊, 庄静. 2020. 茶树DHAR 酶基因的克隆与非生物胁迫响应分析. 园艺学报, 47 (5):983-994.
[42]	Liu Pan, Geng Xingmin, Zhao Hui. 2020. Subcellular distribution and responses of antioxidant systems in leaves of three rhododendron cultivars under alkali stress. Acta Horticulturae Sinica, 47 (5):916-926. (in Chinese)
	刘攀, 耿兴敏, 赵晖. 2020. 碱胁迫下杜鹃花抗氧化体系的响应及亚细胞分布. 园艺学报, 47 (5):916-926.
[43]	Liu R, Howkit A, Stammitti L, Teyssier E, Rolin D, Mortain-Bertrand A, Halle S, Liu M, Kong J, Wu C, Degraeve-Guibault C, Chapman N H, Maucourt M, Hodgman T C, Tost J, Bouzayen M, Hong Y, Seymour G B, Giovannoni J J, Gallusci P. 2015. A DEMETER-like DNA demethylase governs tomato fruit ripening. Proceedings of the National Academy of Sciences of the United States of America, 112 (34):10804-10809.
[44]	Liu X Y, Wu R M, Bulley S M, Zhong C H, Li D W. 2022. Kiwifruit MYBS1-like and GBF 3 transcription factors influence L-ascorbic acid biosynthesis by activating transcription of GDP-L-galactose phosphorylase. New Phytologist, 234 (5):1782-1800. doi: 10.1111/nph.18097 URL
[45]	Loewus F A. 1999. Biosynthesis and metabolism of ascorbic acid in plants and of analogs of ascorbic acid in fungi. Phytochemistry, 52 (2):193-210. doi: 10.1016/S0031-9422(99)00145-4 URL
[46]	Lorence A, Chevone B I, Mendes P, Nessler C L. 2004. Myo-Inositol oxygenase offers a possible entry point into plant ascorbate biosynthesis. Plant Physiology, 134 (3):1200-1205. doi: 10.1104/pp.103.033936 pmid: 14976233
[47]	Ma G, Zhang L C, Setiawan C K, Yamawaki K, Asaia T, Nishikawa F, Maezawa C, Sato H, Kanemitsue N, Kato M. 2014. Effect of red and blue LED light irradiation on ascorbate content and expression of genes related to ascorbate metabolism in postharvest broccoli. Postharvest Biology and Technology, 94:97-103. doi: 10.1016/j.postharvbio.2014.03.010 URL
[48]	Ma S Y, Li H X, Wang L, Li B Y, Wang Z Y, Ma B Q, Ma F W, Li M J. 2021. F-box protein MdAMR1L 1 regulates ascorbate biosynthesis in apple by modulating GDP-mannose pyrophosphorylase. Plant Physiology, 188 (1):653-669. doi: 10.1093/plphys/kiab427 URL
[49]	Mackenzie S, Mcintosh L. 1999. Higher plant mitochondria. The Plant Cell, 11 (4):571-585. doi: 10.1105/tpc.11.4.571 pmid: 10213779
[50]	Maharaj R, Arul J, Nadeau P. 2014. UV-C irradiation effects on levels of enzymic and non-enzymic phytochemicals in tomato. Innovative Food Science and Emerging Technologies, 21:99-106. doi: 10.1016/j.ifset.2013.10.001 URL
[51]	Maruta T, Yonemitsu M, Yabuta Y, Tamoi M, Ishikawa T, Shigeoka S. 2008. Arabidopsis phosphomannose isomerase 1,but not phosphomannose isomerase 2,is essential for ascorbic acid biosynthesis. Journal of Biological Chemistry, 283 (43):28842-28851. doi: 10.1074/jbc.M805538200 URL
[52]	Massot C, Génard M, Stevens R, Gautier H. 2010. Fluctuations in sugar content are not determinant in explaining variations in vitamin C in tomato fruit. Plant Physiology and Biochemistry, 48 (9):751-757. doi: 10.1016/j.plaphy.2010.06.001 pmid: 20621498
[53]	Massot C, Stevens R, Génard M, Longuenesse J, Gautier H. 2012. Light affects ascorbate content and ascorbate-related gene expression in tomato leaves more than in fruits. Planta, 235:153-163. doi: 10.1007/s00425-011-1493-x pmid: 21861113
[54]	May M J, Vernoux T, Leaver C, Montagu M V, Inze´ D. 1998. Glutathione homeostasis in plants:implications for environmental sensing and plant development. Journal of Experimental Botany, 49 (321):649-667.
[55]	Millar A H, Day D A. 1997. Alternative solutions to radical problems. Trends in Plant Science, 2 (8):288-290. doi: 10.1016/S1360-1385(97)89948-7 URL
[56]	Millar A H, Mittova V, Kiddle G, Heazlewood J L, Bartoli C G, Theodoulou F L, Foyer C H. 2003. Control of Ascorbate synthesis by respiration and its implications for stress responses. Plant Physiology, 133 (2):443-447. pmid: 14555771
[57]	Monteiro J A, Nell T A, Barrett J E. 2002. Effects of exogenous sucrose on carbohydrate levels,flower respiration and longevity of potted miniature rose(Rosa hybrida)flowers during postproduction. Postharvest Biology and Technology, 26:221-229. doi: 10.1016/S0925-5214(02)00010-8 URL
[58]	Munir S, Mumtaz M A, Ahiakpa J K, Liu G, Chen W, Zhou G, Zheng W, Ye Z, Zhang Y. 2020. Genome-wide analysis of Myo-inositol oxygenase gene family in tomato reveals their involvement in ascorbic acid accumulation. BMC Genomics, 21 (1):284. doi: 10.1186/s12864-020-6708-8 pmid: 32252624
[59]	Nishikawa F, Kato M, Hyodo H, Ikoma Y, Sugiura M, Yano M. 2005. Effect of sucrose on ascorbate level and expression of genes involved in the ascorbate biosynthesis and recycling pathway in harvested broccoli florets. Journal of Experimental Botany, 56 (409):65-72. pmid: 15520028
[60]	Noctor G, de Paepe R, Foyer C H. 2007. Mitochondrial redox biology and homeostasis in plants. Trends in Plant Science, 12 (9):125-134. doi: 10.1016/j.tplants.2007.01.005 URL
[61]	Noguchi K, Yoshida K. 2008. Interaction between photosynthesis and respiration in illuminated leaves. Mitochondrion, 8 (1):87-99. pmid: 18024239
[62]	Noshi M, Hatanaka R, Tanabe N, Terai Y, Maruta T, Shigeoka S. 2016. Redox regulation of ascorbate and glutathione by a chloroplastic dehydroascorbate reductase is required for high-light stress tolerance in Arabidopsis. Bioscience,Biotechnology,and Biochemistry, 80 (5):1-8. doi: 10.1080/09168451.2015.1056512 URL
[63]	Ntagkasa N, Woltering E J, Marcelis L F M. 2018. Light regulates ascorbate in plants:an integrated view on physiology and biochemistry. Environmental and Experimental Botany 147 (1):271-280. doi: 10.1016/j.envexpbot.2017.10.009 URL
[64]	Ntagkas N, Woltering E, Nicole C, Labried C, Marcelis L F M. 2019. Light regulation of vitamin C in tomato fruit is mediated through photosynthesis. Environmental and Experimental Botany, 158:180-188. doi: 10.1016/j.envexpbot.2018.12.002 URL
[65]	Nunes-Nesi A, Carrari F, Lytovchenko A, Smith A M O, Loureiro M E, Ratcliffe R G, Sweetlove L J, Fernie A R. 2005. Enhanced photosynthetic performance and growth as a consequence of decreasing mitochondrial malate dehydrogenase activity in transgenic tomato plants. Plant Physiology, 137 (2):611-622. doi: 10.1104/pp.104.055566 pmid: 15665243
[66]	Ögren E, Nilsson T, Sundblad L G. 1997. Relationship between respiratory depletion of sugars and loss of cold hardiness in coniferous seedlings over-wintering at raised temperatures:indications of different sensitivities of spruce and pine. Plant,Cell and Environment, 20 (2):247-453. doi: 10.1046/j.1365-3040.1997.d01-56.x URL
[67]	Paul M J, Foyer C H. 2001. Sink regulation of photosynthesis. Journal of Experimental Botany, 52 (360):1383-1400. doi: 10.1093/jexbot/52.360.1383 pmid: 11457898
[68]	Pellny T K, Locato V, Vivancos P D, Markovic J, De Gara L, Pallardó F V, Foyer C H. 2009. Pyridine nucleotide cycling and control of intracellular redox state in relation to poly(ADP-ribose)polymerase activity and nuclear localization of glutathione during exponential growth of Arabidopsis cells in culture. Molecular Plant, 2 (3):442-456. doi: 10.1093/mp/ssp008 pmid: 19825628
[69]	Proietti S, Moscatello S, Colla G, Battistelli Y. 2004. The effect of growing spinach(Spinacia oleracea L.)at two light intensities on the amounts of oxalate,ascorbate and nitrate in their leaves. The Journal of Horticultural Science and Biotechnology, 79 (4):606-609. doi: 10.1080/14620316.2004.11511814 URL
[70]	Reid M E. 1938. The effect of light on the accumulation of ascorbic acid in young cowpea plants. American Journal of Botany, 25 (9):701-711. doi: 10.1002/j.1537-2197.1938.tb12839.x URL
[71]	Sawake S, Tajima N, Mortimer J C, Lao J, Ishikawa I, Yu X, Yamanashi Y, Yoshimi Y, Kawai-Yamada M, Dupree P, Tsumuraya Y, Kotake T. 2015. KONJAC1 and 2 are key factors for GDP-mannose generation and affect L-ascorbic acid and glucomannan biosynthesis in Arabidopsis. The Plant Cell, 27 (12):3397-3409. doi: 10.1105/tpc.15.00379 pmid: 26672069
[72]	Schimmeyer J, Bock R, Meyer E H. 2016. L-Galactono-1,4-lactone dehydrogenase is an assembly factor of the membrane arm of mitochondrial complex I in Arabidopsis. Plant Molecular Biology, 90 (1-2):117-126. doi: 10.1007/s11103-015-0400-4 pmid: 26520835
[73]	Shiroma S, Tanaka M, Sasaki T, Ogawa T, Yoshimura K, Sawa Y, Maruta T, Ishikawa T. 2019. Chloroplast development activates the expression of ascorbate biosynthesis-associated genes in Arabidopsis roots. Plant Science, 284:185-191. doi: S0168-9452(19)30026-3 pmid: 31084871
[74]	Smirnoff N. 2011. Vitamin C:the metabolism and functions of Ascorbic acid in plants. Advances in Botanical Research, 59:109-155.
[75]	Smirnoff N. 2018. Ascorbic acid metabolism and functions:a comparison of plants and mammals. Free Radical Biology & Medicine, 122:116-129. doi: 10.1016/j.freeradbiomed.2018.03.033 URL
[76]	Smirnoff N, Wheeler G L. 2000. Ascorbic acid in plants:biosynthesis and function. Critical Reviews in Biochemistry and Molecular Biology, 35 (4):291-314. doi: 10.1080/10409230008984166 pmid: 11005203
[77]	Sun X, Wang Y, Sui N. 2018. Transcriptional regulation of bHLH during plant response to stress. Biochemical and Biophysical Research Communications, 503 (2):397-401. doi: S0006-291X(18)31625-5 pmid: 30057319
[78]	Tamim S A, Li F, Wang Y, Shang L, Zhang X, Tao J, Wang Y, Gai W, Dong H, Ahiakpa J K, Mumtaz M A, Zhang Y Y. 2022. Effect of shading on ascorbic acid accumulation and biosynthetic gene expression during tomato fruit development and ripening. Vegetable Research, 2:1.
[79]	Tokunaga T, Miyahara K, Tabata K, Esaka M. 2005. Generation and properties of ascorbic acid-overproducing transgenic tobacco cells expressing sense RNA for L-galactono-1,4-lactone dehydrogenase. Planta, 220 (6):854-863. pmid: 15549373
[80]	Torabinejad J, Donahue J L, Gunesekera B N, Allen-Daniels M J, Gillaspy G E. 2009. VTC 4 is a bifunctional enzyme that affects myoinositol and ascorbate biosynthesis in plants. Plant Physiology, 150 (2):951-961. doi: 10.1104/pp.108.135129 pmid: 19339506
[81]	Wang F, Wang X J, Zhang Y, Yan J R, Ahammed G J, Bu X, Sun X, Liu Y F, Xu T, Qi H Y, Qi M F, Li T L. 2022. SlFHY3 and SlHY5 act compliantly to enhance cold tolerance through the integration of myo-inositol and light signaling in tomato. New Phytologist, 233:2127-2143. doi: 10.1111/nph.17934 URL
[82]	Wang F, Wu N, Zhang L Y, Ahammed G L, Chen X X, Xiang X, Zhou J, Xia X J, Shi K, Yu J Q, Foyer C H, Zhou Y H. 2018. Light signaling-dependent regulation of photoinhibition and photoprotection in tomato. Plant Physiology, 176:1311-1326. doi: 10.1104/pp.17.01143 pmid: 29146776
[83]	Wang J, Yu Y W, Zhang Z, Quan R D, Zhang H W, Ma L G, Deng X W, Huang R F. 2013. Arabidopsis CSN5B interacts with VTC1 and modulates ascorbic acid synthesis. The Plant Cell, 25 (2):625-636. doi: 10.1105/tpc.112.106880 URL
[84]	Wei Shenjun, Geng Dianxiang, Wu Qi, Zhang Xiaoyan, Su Nana, Cui Jin. 2015. Effect of UV-A irradiation on ascorbic acid content in soybean sprout. Soybean Science, 34 (3):420-426. (in Chinese)
	魏圣军, 耿殿祥, 邬奇, 张晓燕, 苏娜娜, 崔瑾. 2015. UV-A辐射对大豆芽苗菜中抗坏血酸含量的影响. 大豆科学, 34 (3):420-426.
[85]	Wheeler G L, Jones M A, Smirnoff N. 1998. The biosynthetic pathway of vitamin C in higher plants. Nature, 393 (6683):365-369. doi: 10.1038/30728 URL
[86]	Wolucka B,van Montagu M. 2003. GDP-mannose 3’,5’-epimerase forms GDP-L-gulose,a putative intermediate for the de novo biosynthesis of vitamin C in plants. Journal of Biological Chemistry, 278 (48):47483. doi: 10.1074/jbc.M309135200 pmid: 12954627
[87]	Xiang L, Li Y, Rolland F, Ende W V D. 2011. Neutral invertase,hexokinase and mitochondrial ROS homeostasis:emerging links between sugar metabolism,sugar signaling and Ascorbate synthesis. Plant Signaling & Behavior, 6 (10):1567-1573.
[88]	Xu M J, Dong J F, Zhu M Y. 2005. Effects of germination conditions on ascorbic acid level and yield of soybean sprouts. Journal of the Science of Food and Agriculture, 85 (6):943-947. doi: 10.1002/jsfa.2050 URL
[89]	Yabuta Y, Mieda T, Rapolu M, Nakamura A, Motoki T, Maruta T, Yoshimura K, Ishikawa T, Shigeoka S. 2007. Light regulation of ascorbate biosynthesis is dependent on the photosynthetic electron transport chain but independent of sugars in Arabidopsis. Journal of Experimental Botany, 58 (10):2661-2671. doi: 10.1093/jxb/erm124 URL
[90]	Yang C, Huang S, Zeng Y, Liu C, Ma Q, Pruneda-Paz J, Kay S A, Li L. 2021. Two bHLH transcription factors,bHLH48 and bHLH60,associate with phytochrome interacting factor 7 to regulate hypocotyl elongation in Arabidopsis. Cell Reports, 35 (5):109054. doi: 10.1016/j.celrep.2021.109054 URL
[91]	Yang Tao. 2017. Study on effect of genetically modified polymerization on ascorbic acid content in tomato[M. D. Dessertation]. Wuhan:Huazhong Agriculture University. (in Chinese)
	杨涛. 2017. 转基因聚合影响番茄抗坏血酸含量的研究[硕士论文]. 武汉: 华中农业大学.
[92]	Ye J, Li W, Ai G, Li C X, Liu G Z, Chen W F, Wang B, Wang W Q, Lu Y G, Zhang J H, Li H X, Ouyang B, Zhang H Y, Fei Z J, Giovannoni J J, Ye Z B, Zhang Y Y. 2019. Genome-wide association analysis identifies a natural variation in basic helix-loop-helix transcription factor regulating ascorbate biosynthesis via D-mannose/L-galactose pathway in tomato. PLoS Genetics, 15 (5):e1008149. doi: 10.1371/journal.pgen.1008149 URL
[93]	Yoshimura K, Yabuta Y, Ishikawa T, Shigeoka S. 2000. Expression of spinach ascorbate peroxidase isoenzymes in response to oxidative stresses. Plant Physiology, 123 (1):223-234. pmid: 10806239
[94]	Yu Yi, Liu Wenke. 2015. Influence of light quality and photoperiod on growth and nutritional quality of three leaf-color lettuce cultivars under weak light. Chinese Journal of Agrometeorology, 36 (6):739-745. (in Chinese)
	余意, 刘文科. 2015. 弱光条件下光质和光周期对水培生菜生长与品质的影响. 中国农业气象, 36 (6):739-745.
[95]	Yu Y W, Wang J, Li S H, Kakan X, Zhou Y, Miao Y C, Wang F F, Qin H, Huang R F. 2019. Ascorbic acid integrates the antagonistic modulation of ethylene and abscisic acid in the accumulation of reactive oxygen species. Plant Physiology, 179:1861-1875. doi: 10.1104/pp.18.01250 pmid: 30723177
[96]	Zha L Y, Zhang Y B, Liu W K. 2019. Dynamic responses of ascorbate pool and metabolism in lettuce to longterm continuous light provided by red and blue LEDs. Environmental and Experimental Botany, 163:15-23. doi: 10.1016/j.envexpbot.2019.04.003 URL
[97]	Zhang C J, Liu J X, Zhang Y Y, Cai X F, Gong P J, Zhang J H, Wang T T, Li H X, Ye Z B. 2010. Overexpression of SlGMEs leads to ascorbate accumulation with enhanced oxidative stress,cold,and salt tolerance in tomato. Plant Cell Reports, 30 (3):389-398. doi: 10.1007/s00299-010-0939-0 URL
[98]	Zhang Huan, Zhang Lili, Li Wei, Xin Zenan, Zhang Dan, Cui Jin. 2012. Effects of photoperiod under red led on growth and quality of sunflower sprouts. Acta Horticulturae Sinica, 39 (2):297-304. (in Chinese)
	张欢, 章丽丽, 李薇, 邢泽南, 张丹, 崔瑾. 2012. 不同光周期红光对油葵芽苗菜生长和品质的影响. 园艺学报, 39 (2):297-304.
[99]	Zhang H W, Si X M, Ji X, Fan R, Liu J X, Chen K L, Wang D W, Gao C X. 2018. Genome editing of upstream open reading frames enables translational control in plants. Nature Biotechnology, 36:894-894. doi: 10.1038/nbt.4202 pmid: 30080209
[100]	Zhang L C, Ma G, Yamawaki K, Ikoma Y, Matsumoto H, Yoshioka T, Ohta S, Kato M. 2015. Regulation of ascorbic acid metabolism by blue LED light irradiation in citrus juice sacs. Plant Science, 233:134-142. doi: S0168-9452(15)00029-1 pmid: 25711821
[101]	Zhang W, Lorence A, Gruszewski H A, Chevone B I, Nessler C L. 2009. AMR1,an Arabidopsis gene that coordinately and negatively regulates the mannose/L-galactose ascorbic acid biosynthetic pathway. Plant Physiology, 150 (2):942-950. doi: 10.1104/pp.109.138453 URL
[102]	Zhang Y. 2012. Ascorbic acid in plants:biosynthesis,regulation and enhancement. Berlin: Springer Science & Business Media.
[103]	Zhang Y Y, Li H X, Shu W B, Zhang C J, Zhang W, Ye Z B. 2011. Suppressed expression of ascorbate oxidase gene promotes ascorbic acid accumulation in tomato fruit. Plant Molecular Biology Reporter, 29 (3):638-645. doi: 10.1007/s11105-010-0271-4 URL
[104]	Zhang Z J, Wang J, Zhang R X, Huang R F. 2012. ERF protein AtERF 98 enhances tolerance to salt through the transcriptional activation of ascorbic acid synthesis in Arabidopsis. The Plant Journal, 71 (2):273-287. doi: 10.1111/j.1365-313X.2012.04996.x URL
[105]	Zou L P, Li H X, Ouyang B, Zhang J H, Ye Z B. 2006. Cloning and mapping of genes involved in tomato ascorbic acid biosynthesis and metabolism. Plant Science, 170:120-127. doi: 10.1016/j.plantsci.2005.08.009 URL

园艺作物抗坏血酸生物合成中光的调控作用研究综述

Light Regulation of Ascorbic Acid Biosynthesis in Horticultural Crops

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园艺作物抗坏血酸生物合成中光的调控作用研究综述

Light Regulation of Ascorbic Acid Biosynthesis in Horticultural Crops

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