柑橘果实糖积累特征及蔗糖转运机制研究进展

doi:10.16420/j.issn.0513-353x.2021-0662

摘要/Abstract

摘要：

糖是决定柑橘果实品质的重要因子，阐明柑橘果实糖积累特征及其调控机制有助于果实糖积累的精准调控。本文全面综述了柑橘果实糖累积的机制，总结了不同糖积累型柑橘果实中糖积累差异、糖随果实发育的积累特征及糖与其代谢相关酶活的关系，并整合构建了蔗糖积累型和己糖积累型的代谢差异模型。同时系统阐述了蔗糖由源叶合成到果实库器官的储存过程，主要包括韧皮部装载、卸载、运输以及蔗糖在果实液泡中的储存等过程，并绘制了蔗糖从“源”到“库”的转运模式图。

关键词: 柑橘, 糖, 蔗糖转运, 蔗糖代谢, 液泡储存

Abstract:

Sugar is an important factor that determines fruit quality of citrus. Illustrating the characteristic and regulation mechanism of citrus fruit sugar metabolism and transport will greatly contribute to regulating sugar accumulation accurately. In this paper，the mechanisms underlying sugar accumulation were comprehensively reviewed，we also summarized the differences of fruit sugar accumulation in various citrus varieties，the characteristics of sugar accumulation with fruit development and the relationship between sugar and the related enzyme activities. Subsequently，a model of metabolic differences between sucrose accumulation type and hexose accumulation type was mapped in citrus. Moreover，the storage process of sucrose from leaf（source organ）to fruit（sink organ）was described systematically，including phloem loading，unloading，transport and sucrose storage in fruit vacuole. Finally，a model of sugar transport from source to sink was constructed in citrus.

Key words: citrus, sugar, sucrose transport, sucrose metabolism, vacuolar storage

中图分类号:

S666
Q 945

吴孔杰, 胡承孝, 谭启玲, 孙学成, 赵小虎, 武松伟. 柑橘果实糖积累特征及蔗糖转运机制研究进展[J]. 园艺学报, 2022, 49(12): 2543-2558.

WU Kongjie, HU Chengxiao, TAN Qiling, SUN Xuecheng, ZHAO Xiaohu, WU Songwei. Research Advanced on Character of Sugar Accumulation and Mechanism of Sucrose Transport in Citrus Fruit[J]. Acta Horticulturae Sinica, 2022, 49(12): 2543-2558.

图/表 4

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品种 Variety	可溶性固形物/% Soluble solid	总糖/% Total sugar	调查年限 Investigation period	产区 Producing area	参考文献 Reference
砂糖橘Citrus reticulata‘Shatangju’	12.60 ~ 17.90	8.36 ~ 14.03	2012、2013、2015—2018	广东 Guangdong	刘翔宇等，2015;刘振等，2016;吴文等，2017，2020
南丰蜜橘C. reticulata ‘Nanfeng Tangerine’	12.50 ~ 13.90	11.70 ~ 12.40	2015、2017、2019	江西 Jiangxi	成传香等，2019;汪勇等，2021;付学琴等，2020
冰糖橙C. sinensis ‘Bingtangcheng’	12.50 ~ 15.60	11.20 ~ 12.30	2002、2004、2017、2018	湖南 Hunan	廖振坤等，2006;朱丽莎等，2018;李大志等，2019;曹胜等，2021
宫川温州蜜柑C. unshiu‘Miyagawa-wase’	10.70 ~ 13.30	8.40 ~ 11.70	2001、2002、2011、2017	浙江、湖北、湖南 Zhejiang，Hunan and Hubei	廖振坤等，2006;谢亚超等，2012;林媚和吴韶辉，2019;成传香等，2019
沃柑C. reticulata‘Orah’	10.80 ~ 13.30	7.80 ~ 11.30	2017、2019、2020、2021	浙江、四川、广西 Zhejiang，Sichuan and Guangxi	林媚和吴韶辉，2019;蒲成伟等，2020;梁芳等，2021
椪柑C. reticulata ‘Ponkan’	11.40 ~ 13.00	8.00 ~ 11.00	2017—2019	浙江、湖南 Zhejiang and Hunan	成传香等，2019;林媚和吴韶辉，2019;侯海军等，2020
伦晚脐橙C. sinensis‘Lunwan Navel’	10.60 ~ 13.60	7.80 ~ 9.76	2013、2016、2018	重庆、湖北 Chongqing and Hubei	叶荫民，2010;吴黎明等，2015;朱丽莎等，2018;姚京磊等，2020
塔罗科血橙C. sinensis‘Tarocco’	10.40 ~ 12.70	7.90 ~ 9.10	2018、2019	重庆 Chongqing	叶荫民，2010;朱丽莎等，2018;刘雪峰等，2021
不知火C. reticulata ‘Shiranuhi Tangor’	11.50 ~ 13.80	7.70 ~ 10.40	2017、2020	浙江、四川 Zhejiang and Sichuan	林媚和吴韶辉，2019;蒲成伟等，2020;
葡萄柚C. paradisi	9.80 ~ 12.50	6.70 ~ 10.20	2017、2019、2020	浙江 Zhejiang	林媚和吴韶辉，2019;刘春荣等，2020
三红蜜柚C. grandis ‘Sanhong Miyou’	10.50 ~ 12.00	8.90 ~ 9.50	2018	福建、广西 Fujian and Guangxi	吴世涛等，2020;汪妮娜等，2021
纽荷尔脐橙C. sinensis‘Navel Orange’	10.30 ~ 13.60	5.60 ~ 8.80	2001、2015、2017、2018	四川、湖南 Sichuan and Hunan	廖振坤等，2006;陈品文等，2017;李大志等，2019;朱丽莎等，2018
本地早C. reticulata‘Succosa’	9.80 ~ 11.90	6.80 ~ 9.20	2016—2018	浙江、湖北 Zhejiang and Hubei	林媚和吴韶辉，2019;王蓉等，2019
龙安柚C. grandis ‘Longanyou’	10.20 ~ 10.90	6.20 ~ 8.30	2016—2017	四川 Sichuan	李荣飞等，2017，2020
琯溪蜜柚C. grandis‘Guanxi Miyou’	9.70 ~ 11.60	5.60 ~ 6.90	2004、2017、2018	福建 Fujian	廖振坤等，2006;晏幸等，2019;吴世涛等，2020;刘怀伟等，2021
柠檬C. limon	6.10 ~ 10.30	0.20 ~ 1.20	2008—2010、2016—2018	重庆、云南 Chongqing and Yunnan	洪林和谢永红，2012;杜玉霞等，2021

品种 Variety	可溶性固形物/% Soluble solid	总糖/% Total sugar	调查年限 Investigation period	产区 Producing area	参考文献 Reference
砂糖橘Citrus reticulata‘Shatangju’	12.60 ~ 17.90	8.36 ~ 14.03	2012、2013、2015—2018	广东 Guangdong	刘翔宇等，2015;刘振等，2016;吴文等，2017，2020
南丰蜜橘C. reticulata ‘Nanfeng Tangerine’	12.50 ~ 13.90	11.70 ~ 12.40	2015、2017、2019	江西 Jiangxi	成传香等，2019;汪勇等，2021;付学琴等，2020
冰糖橙C. sinensis ‘Bingtangcheng’	12.50 ~ 15.60	11.20 ~ 12.30	2002、2004、2017、2018	湖南 Hunan	廖振坤等，2006;朱丽莎等，2018;李大志等，2019;曹胜等，2021
宫川温州蜜柑C. unshiu‘Miyagawa-wase’	10.70 ~ 13.30	8.40 ~ 11.70	2001、2002、2011、2017	浙江、湖北、湖南 Zhejiang，Hunan and Hubei	廖振坤等，2006;谢亚超等，2012;林媚和吴韶辉，2019;成传香等，2019
沃柑C. reticulata‘Orah’	10.80 ~ 13.30	7.80 ~ 11.30	2017、2019、2020、2021	浙江、四川、广西 Zhejiang，Sichuan and Guangxi	林媚和吴韶辉，2019;蒲成伟等，2020;梁芳等，2021
椪柑C. reticulata ‘Ponkan’	11.40 ~ 13.00	8.00 ~ 11.00	2017—2019	浙江、湖南 Zhejiang and Hunan	成传香等，2019;林媚和吴韶辉，2019;侯海军等，2020
伦晚脐橙C. sinensis‘Lunwan Navel’	10.60 ~ 13.60	7.80 ~ 9.76	2013、2016、2018	重庆、湖北 Chongqing and Hubei	叶荫民，2010;吴黎明等，2015;朱丽莎等，2018;姚京磊等，2020
塔罗科血橙C. sinensis‘Tarocco’	10.40 ~ 12.70	7.90 ~ 9.10	2018、2019	重庆 Chongqing	叶荫民，2010;朱丽莎等，2018;刘雪峰等，2021
不知火C. reticulata ‘Shiranuhi Tangor’	11.50 ~ 13.80	7.70 ~ 10.40	2017、2020	浙江、四川 Zhejiang and Sichuan	林媚和吴韶辉，2019;蒲成伟等，2020;
葡萄柚C. paradisi	9.80 ~ 12.50	6.70 ~ 10.20	2017、2019、2020	浙江 Zhejiang	林媚和吴韶辉，2019;刘春荣等，2020
三红蜜柚C. grandis ‘Sanhong Miyou’	10.50 ~ 12.00	8.90 ~ 9.50	2018	福建、广西 Fujian and Guangxi	吴世涛等，2020;汪妮娜等，2021
纽荷尔脐橙C. sinensis‘Navel Orange’	10.30 ~ 13.60	5.60 ~ 8.80	2001、2015、2017、2018	四川、湖南 Sichuan and Hunan	廖振坤等，2006;陈品文等，2017;李大志等，2019;朱丽莎等，2018
本地早C. reticulata‘Succosa’	9.80 ~ 11.90	6.80 ~ 9.20	2016—2018	浙江、湖北 Zhejiang and Hubei	林媚和吴韶辉，2019;王蓉等，2019
龙安柚C. grandis ‘Longanyou’	10.20 ~ 10.90	6.20 ~ 8.30	2016—2017	四川 Sichuan	李荣飞等，2017，2020
琯溪蜜柚C. grandis‘Guanxi Miyou’	9.70 ~ 11.60	5.60 ~ 6.90	2004、2017、2018	福建 Fujian	廖振坤等，2006;晏幸等，2019;吴世涛等，2020;刘怀伟等，2021
柠檬C. limon	6.10 ~ 10.30	0.20 ~ 1.20	2008—2010、2016—2018	重庆、云南 Chongqing and Yunnan	洪林和谢永红，2012;杜玉霞等，2021

参与过程Participate in the process	物种 Species	编码基因 Encoding gene	亚细胞定位 Subcellular localization	主要生理功能 Main physiological function	转运方向 Transport direction	参考文献Reference
韧皮部装载Phloem loading	葡萄 Vitis vinifera	VvSWEET4	质膜 Plasma membrane	葡萄糖转运载体 Glucose transporter	流出 Efflux	Julie et al.，2014
可能参与韧皮部装载 Probably be invovled in phloem loading	甜橙 C. sinensis	CsSWEETs	可能位于韧皮部薄壁细胞膜 Probably located on phloem parenchyma cell membrane	蔗糖异化扩散转运体 Facilitated diffusion transporter of sucrose	流出 Efflux	Zheng et al.，2014
	脐橙 C. sinensis	CsSWEET10	韧皮部薄壁细胞膜 Phloem parenchyma cell membrane	蔗糖转运载体 Sucrose transporter	流出 Efflux	Feng et al.，2021
	脐橙 C. sinensis	CsSWEET15	韧皮部薄壁细胞膜 Phloem parenchyma cell membrane	蔗糖转运载体 Sucrose transporter	流出 Efflux	Feng et al.，2021
	温州蜜柑 Satsuma Mandarin	CitSUT1，CitSUT2，CitSUT3	可能定位于质膜 Probably located on plasma membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	可能流出Probably efflux	Islam et al.，2015
	默科特 Murcott Tangerine	CitSUT1	可能位于韧皮部薄壁细胞膜 Probably located on phloem parenchyma cell membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Li et al.，2003
	甜橙 C. sinensis	CsSUT1，CsSUT2	可能定位于质膜 Probably located on plasma membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zheng et al.，2014
液泡糖流出 Vacuole sugar efflux	甜橙 C. sinensis	CsSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zheng et al.，2014
	甜橙 C. sinensis	CsERD6L	液泡膜 Tonoplast	葡萄糖易化扩散转运体 Facilitated diffusion transporter of glucose	流出 Efflux	Zheng et al.，2014
	脐橙 C. sinensis	CsSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Feng et al.，2021
	桃 Prunus persica	PpSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zanon et al.，2015
	苹果 Malus × domestica	MdERD6L	液泡膜 Tonoplast	葡萄糖/H⁺同向转运载体 Glucose/H⁺symporter	流出 Efflux	Zhu et al.，2021
液泡糖储存 Vacuole sugar storage	甜橙 C. sinensis	CsTMT1，CsTMT2	液泡膜 Tonoplast	单糖转运体Monosaccharides transporter	流入 Influx	Zheng et al.，2014
	甜橙 C. sinensis	CsVGT1，CsVGT2	液泡膜 Tonoplast	葡萄糖转运载体 Glucose transporter	流入 Influx	Zheng et al.，2014
	苹果 Malus × domestica	MdVGT1	液泡膜 Tonoplast	葡萄糖转运载体 Glucose transporter	流入 Influx	许海峰等，2016，2017
	苹果 Malus × domestica	MdTST1，MdTST2	液泡膜 Tonoplast	糖/H⁺反向转运载体 Sugar/H⁺ antiporters	流入 Influx	Zhu et al.，2021
	桃 Prunus persica	PpTST1	液泡膜 Tonoplast	蔗糖/H⁺反向转运载体 Sucrose/H⁺ antiporters	流入 Influx	Peng et al.，2020

参与过程Participate in the process	物种 Species	编码基因 Encoding gene	亚细胞定位 Subcellular localization	主要生理功能 Main physiological function	转运方向 Transport direction	参考文献Reference
韧皮部装载Phloem loading	葡萄 Vitis vinifera	VvSWEET4	质膜 Plasma membrane	葡萄糖转运载体 Glucose transporter	流出 Efflux	Julie et al.，2014
可能参与韧皮部装载 Probably be invovled in phloem loading	甜橙 C. sinensis	CsSWEETs	可能位于韧皮部薄壁细胞膜 Probably located on phloem parenchyma cell membrane	蔗糖异化扩散转运体 Facilitated diffusion transporter of sucrose	流出 Efflux	Zheng et al.，2014
	脐橙 C. sinensis	CsSWEET10	韧皮部薄壁细胞膜 Phloem parenchyma cell membrane	蔗糖转运载体 Sucrose transporter	流出 Efflux	Feng et al.，2021
	脐橙 C. sinensis	CsSWEET15	韧皮部薄壁细胞膜 Phloem parenchyma cell membrane	蔗糖转运载体 Sucrose transporter	流出 Efflux	Feng et al.，2021
	温州蜜柑 Satsuma Mandarin	CitSUT1，CitSUT2，CitSUT3	可能定位于质膜 Probably located on plasma membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	可能流出Probably efflux	Islam et al.，2015
	默科特 Murcott Tangerine	CitSUT1	可能位于韧皮部薄壁细胞膜 Probably located on phloem parenchyma cell membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Li et al.，2003
	甜橙 C. sinensis	CsSUT1，CsSUT2	可能定位于质膜 Probably located on plasma membrane	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zheng et al.，2014
液泡糖流出 Vacuole sugar efflux	甜橙 C. sinensis	CsSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zheng et al.，2014
	甜橙 C. sinensis	CsERD6L	液泡膜 Tonoplast	葡萄糖易化扩散转运体 Facilitated diffusion transporter of glucose	流出 Efflux	Zheng et al.，2014
	脐橙 C. sinensis	CsSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Feng et al.，2021
	桃 Prunus persica	PpSUT4	液泡膜 Tonoplast	蔗糖/H⁺同向转运载体 Sucrose/H⁺ symporter	流出 Efflux	Zanon et al.，2015
	苹果 Malus × domestica	MdERD6L	液泡膜 Tonoplast	葡萄糖/H⁺同向转运载体 Glucose/H⁺symporter	流出 Efflux	Zhu et al.，2021
液泡糖储存 Vacuole sugar storage	甜橙 C. sinensis	CsTMT1，CsTMT2	液泡膜 Tonoplast	单糖转运体Monosaccharides transporter	流入 Influx	Zheng et al.，2014
	甜橙 C. sinensis	CsVGT1，CsVGT2	液泡膜 Tonoplast	葡萄糖转运载体 Glucose transporter	流入 Influx	Zheng et al.，2014
	苹果 Malus × domestica	MdVGT1	液泡膜 Tonoplast	葡萄糖转运载体 Glucose transporter	流入 Influx	许海峰等，2016，2017
	苹果 Malus × domestica	MdTST1，MdTST2	液泡膜 Tonoplast	糖/H⁺反向转运载体 Sugar/H⁺ antiporters	流入 Influx	Zhu et al.，2021
	桃 Prunus persica	PpTST1	液泡膜 Tonoplast	蔗糖/H⁺反向转运载体 Sucrose/H⁺ antiporters	流入 Influx	Peng et al.，2020