植物中光合产物运输相关蛋白研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-0494

摘要/Abstract

摘要：

植物光合产物的运输是植物生长和发育中至关重要的过程，涉及多种特定的转运蛋白，这些转运蛋白在维持植物正常代谢、调节能量分配、促进不同源库组织间的营养物质交换方面发挥重要作用。近年来，关于糖转运蛋白的研究取得了一定进展，蔗糖转运蛋白（sucrose transporter, SUT）和糖输出转运蛋白（the sugars will eventually be exported transporter, SWEET）主要负责在不同植物的组织器官之间运输蔗糖，单糖转运蛋白（monosaccharide transporter, MST）则介导单糖的跨膜转运。糖转化酶（invertase, INV）和蔗糖合酶（sucrose synthase, SUS）参与蔗糖的水解与合成。这些蛋白的协同作用对于调控光合产物的分配与代谢至关重要，且影响着植物对环境变化的响应和产量优化。本文综述了植物中与光合产物运输相关的主要转运蛋白，重点介绍了不同植物中SUT、SWEET、INV等的生物学功能及其分子机制，探讨了它们在植物生长发育、抗逆性以及产量调控中的潜在应用，以期为未来提高作物产量，推动农业生产提供新的理论依据和实践策略。

关键词: 蔗糖转运蛋白, 糖输出转运蛋白, 单糖转运蛋白, 糖转化酶, 蔗糖合酶

Abstract:

The transport of photosynthetic products is a critical process in plant growth and development, involving multiple specialized transporters that play essential roles in maintaining normal plant metabolism, regulating energy allocation, and facilitating nutrient exchange between source and sink tissues. Recent years have witnessed significant advances in research on sugar transporters. Sucrose transporters (SUT) and sugars will eventually be exported transporters (SWEET) primarily mediate sucrose translocation between different plant tissues and organs, while monosaccharide transporters (MST) facilitate transmembrane monosaccharide transport. Invertases (INV) and sucrose synthases (SUS) participate in sucrose hydrolysis and synthesis, thereby regulating the transport of photosynthetic assimilates within plants. The coordinated action of these proteins is crucial for controlling the allocation and metabolism of photosynthetic products, with profound implications for plant responses to environmental changes and yield optimization. This review summarizes major transporters associated with photosynthetic product transport in plants, highlighting the biological functions and molecular mechanisms of sucrose transporters, SWEET proteins, invertases, and related proteins across diverse plant species. It further explores their potential applications in plant development, stress resistance, and yield regulation. Finally, this review aims to provide new theoretical foundations and practical strategies for enhancing crop productivity and advancing agricultural production.

Key words: SUT, SWEET, MST, INV, SUS

吕若彤, 孙晶, 李新颖, 王旭晶, 艾鹏飞, 王雁伟. 植物中光合产物运输相关蛋白研究进展[J]. 生物技术通报, 2025, 41(10): 43-53.

LYU Ruo-tong, SUN Jing, LI Xin-ying, WANG Xu-jing, AI Peng-fei, WANG Yan-wei. Progress in the Study of Proteins Related to Photosynthetic Product Transport in Plants[J]. Biotechnology Bulletin, 2025, 41(10): 43-53.

图/表 3

图1 植物韧皮部装卸示意图

Fig. 1 Schematic diagram of plant phloem loading and unloading

图2 MFS转运蛋白的运输机制

Fig. 2 Transport mechanism of MFS transporters

图3 糖转运蛋白调控植物的源库平衡

Fig. 3 Sugar transporter proteins regulate source-sink homeostasis in plants

参考文献 89

[1]	Ruan YL. Sucrose metabolism: gateway to diverse carbon use and sugar signaling [J]. Annu Rev Plant Biol, 2014, 65: 33-67.
[2]	Rossi M, Bermudez L, Carrari F. Crop yield: challenges from a metabolic perspective [J]. Curr Opin Plant Biol, 2015, 25: 79-89.
[3]	Miras M, Pottier M, Schladt TM, et al. Plasmodesmata and their role in assimilate translocation [J]. J Plant Physiol, 2022, 270: 153633.
[4]	Braun DM, Wang L, Ruan YL. Understanding and manipulating sucrose phloem loading, unloading, metabolism, and signalling to enhance crop yield and food security [J]. J Exp Bot, 2014, 65(7): 1713-1735.
[5]	Singh J, Das S, Jagadis Gupta K, et al. Physiological implications of SWEETs in plants and their potential applications in improving source-sink relationships for enhanced yield [J]. Plant Biotechnol J, 2023, 21(8): 1528-1541.
[6]	Niño-González M, Novo-Uzal E, Richardson DN, et al. More transporters, more substrates: the Arabidopsis major facilitator superfamily revisited [J]. Mol Plant, 2019, 12(9): 1182-1202.
[7]	邓东, 颜宁. MFS超家族转运蛋白结构基础及转运机制 [J]. 科学通报, 2015, 60(8): 720-728.
	Deng D, Yan N. Structural basis and transport mechanism of the major facility superfamily (MFS) transporter [J]. Chin Sci Bull, 2015, 60(8): 720-728.
[8]	Shi YG. Common folds and transport mechanisms of secondary active transporters [J]. Annu Rev Biophys, 2013, 42: 51-72.
[9]	Li Q, ChengWX, Jiang YD, et al. Research progress on the structural basis and transport mechanism of MFS superfamily transporters [J]. Int J Front Med, 2024, 6(11): 53-58.
[10]	Yan NE. Structural biology of the major facilitator superfamily transporters [J]. Annu Rev Biophys, 2015, 44: 257-283.
[11]	Chen LQ, Hou BH, Lalonde S, et al. Sugar transporters for intercellular exchange and nutrition of pathogens [J]. Nature, 2010, 468(7323): 527-532.
[12]	Anjali A, Fatima U, Manu MS, et al. Structure and regulation of SWEET transporters in plants: an update [J]. Plant Physiol Biochem, 2020, 156: 1-6.
[13]	Jeena GS, Kumar S, Shukla RK. Structure, evolution and diverse physiological roles of SWEET sugar transporters in plants [J]. Plant Mol Biol, 2019, 100(4-5): 351-365.
[14]	Chang YA, Dai NC, Chen HJ, et al. Regulation of rice sucrose transporter 4 gene expression in response to insect herbivore chewing [J]. J Plant Interact, 2019, 14(1): 525-532.
[15]	Gottwald JR, Krysan PJ, Young JC, et al. Genetic evidence for the in planta role of phloem-specific plasma membrane sucrose transporters [J]. Proc Natl Acad Sci USA, 2000, 97(25): 13979-13984.
[16]	Seitz J, Reimann TM, Fritz C, et al. How pollen tubes fight for food: the impact of sucrose carriers and invertases of Arabidopsis thaliana on pollen development and pollen tube growth [J]. Front Plant Sci, 2023, 14: 1063765.
[17]	Durand M, Mainson D, Porcheron B, et al. Carbon source-sink relationship in Arabidopsis thaliana: the role of sucrose transporters [J]. Planta, 2018, 247(3): 587-611.
[18]	Eom JS, Cho JI, Reinders A, et al. Impaired function of the tonoplast-localized sucrose transporter in rice, OsSUT2, limits the transport of vacuolar reserve sucrose and affects plant growth [J]. Plant Physiol, 2011, 157(1): 109-119.
[19]	Siahpoosh MR, Sanchez DH, Schlereth A, et al. Modification of OsSUT1 gene expression modulates the salt response of rice Oryza sativa cv. Taipei 309 [J]. Plant Sci, 2012, 182: 101-111.
[20]	Wang XW, Liu XL, Hu Z, et al. Essentiality for rice fertility and alternative splicing of OsSUT1 [J]. Plant Sci, 2022, 314: 111065.
[21]	Wei SA, Chen JY, Hsiao HH, et al. Characterization of OsSUT2 expression and regulation in germinating embryos of rice seeds [J]. Rice, 2011, 4(2): 39-49.
[22]	Prasad D, Jung WJ, Seo YW. Identification and molecular characterization of novel sucrose transporters in the hexaploid wheat (Triticum aestivum L.) [J]. Gene, 2023, 860: 147245.
[23]	Li DD, Xu RC, Lv D, et al. Identification of the core pollen-specific regulation in the rice OsSUT3 promoter [J]. Int J Mol Sci, 2020, 21(6): 1909.
[24]	李孟珠, 王高鹏, 巫月, 等. 水稻蔗糖转运蛋白OsSUT4参与蔗糖转运的功能研究 [J].中国水稻科学, 2020, 34(6): 491-498.
	Li MZ, Wang GP, Wu Y, et al. Function analysis of sucrose transporter OsSUT4 in sucrosetransport in rice [J]. Chin J Rice Sci,2020, 34(6): 491-498.
[25]	张雅文, 包淑慧, 唐振家, 等. 蔗糖转运蛋白OsSUT5在水稻花粉发育及结实中的作用 [J]. 中国农业科学, 2021, 54(16): 3369-3385.
	Zhang YW, Bao SH, Tang ZJ, et al. Function of sucrose transporter OsSUT5 in rice pollen development and seed setting [J]. Sci Agric Sin, 2021, 54(16): 3369-3385.
[26]	Xu QY, Chen SY, Ren YJ, et al. Regulation of sucrose transporters and phloem loading in response to environmental cues [J]. Plant Physiol, 2018, 176(1): 930-945.
[27]	Takahashi S, Meguro-Maoka A, Yoshida M. Analysis of sugar content and expression of sucrose transporter genes (OsSUTs) in rice tissues in response to a chilling temperature [J]. Jpn Agric Res Q, 2017, 51(2): 137-146.
[28]	Peng CC, Xu YH, Xi RC, et al. Expression, subcellular localization and phytohormone stimulation of a functional sucrose transporter (MdSUT1) in apple fruit [J]. Sci Hortic, 2011, 128(3): 206-212.
[29]	Zhang B, Li YN, Wu BH, et al. Plasma membrane-localized transporter MdSWEET12 is involved in sucrose unloading in apple fruit [J]. J Agric Food Chem, 2022, 70(49): 15517-15530.
[30]	许海峰, 曲常志, 刘静轩, 等. 苹果液泡膜蔗糖转运蛋白基因MdSUT4的表达分析与功能鉴定 [J]. 园艺学报, 2017, 44(7): 1235-1243.
	Xu HF, Qu CZ, Liu JX, et al. Expression analysis and functional identification of a vacuolar sucrose transporter gene MdSUT4 in apple [J]. Acta Hortic Sin, 2017, 44(7): 1235-1243.
[31]	Li W, Sun K, Ren ZY, et al. Molecular evolution and stress and phytohormone responsiveness of SUT genes in Gossypium hirsutum [J]. Front Genet, 2018, 9: 494.
[32]	Iftikhar J, Lyu ML, Liu ZY, et al. Sugar and hormone dynamics and the expression profiles of SUT/SUC and SWEET sweet sugar transporters during flower development in Petunia axillaris [J]. Plants, 2020, 9(12): 1770.
[33]	Abelenda JA, Bergonzi S, Oortwijn M, et al. Source-sink regulation is mediated by interaction of an FT homolog with a SWEET protein in potato [J]. Curr Biol, 2019, 29(7): 1178-1186.e6.
[34]	汪洋一舟, 郭尽新, 乔凯彬, 等. SWEET蛋白在植物与病原物互作中的功能研究进展 [J]. 吉林大学学报: 理学版, 2025, 63(1): 241-252.
	Wang Y, Guo JX, Qiao KB, et al. Research advances on function of SWEET protein in plant-pathogen interactions [J]. J Jilin Univ Sci Ed, 2025, 63(1): 241-252.
[35]	Wang J, Yu YC, Li Y, et al. Hexose transporter SWEET5 confers galactose sensitivity to Arabidopsis pollen germination via a galactokinase [J]. Plant Physiol, 2022, 189(1): 388-401.
[36]	Talbot NJ. Cell biology: raiding the sweet shop [J]. Nature, 2010, 468(7323): 510-511.
[37]	Lin IW, Sosso D, Chen LQ, et al. Nectar secretion requires sucrose phosphate synthases and the sugar transporter SWEET9 [J]. Nature, 2014, 508(7497): 546-549.
[38]	Chen LQ, Qu XQ, Hou BH, et al. Sucrose efflux mediated by SWEET proteins as a key step for phloem transport [J]. Science, 2012, 335(6065): 207-211.
[39]	Sun MX, Huang XY, Yang J, et al. Arabidopsis RPG1 is important for primexine deposition and functions redundantly with RPG2 for plant fertility at the late reproductive stage [J]. Plant Reprod, 2013, 26(2): 83-91.
[40]	Klemens PAW, Patzke K, Deitmer J, et al. Overexpression of the vacuolar sugar carrier AtSWEET16 modifies germination, growth, and stress tolerance in Arabidopsis [J]. Plant Physiol, 2013, 163(3): 1338-1352.
[41]	Valifard M, Le Hir R, Müller J, et al. Vacuolar fructose transporter SWEET17 is critical for root development and drought tolerance [J]. Plant Physiol, 2021, 187(4): 2716-2730.
[42]	Morii M, Sugihara A, Takehara S, et al. The dual function of OsSWEET3a as a gibberellin and glucose transporter is important for young shoot development in rice [J]. Plant Cell Physiol, 2020, 61(11): 1935-1945.
[43]	Zhou Y, Liu L, Huang WF, et al. Overexpression of OsSWEET5 in rice causes growth retardation and precocious senescence [J]. PLoS One, 2014, 9(4): e94210.
[44]	Li J, Chen D, Jiang GL, et al. Molecular cloning and expression analysis of EjSWEET15, enconding for a sugar transporter from loquat [J]. Sci Hortic, 2020, 272: 109552.
[45]	Fei HH, Yang ZP, Lu QT, et al. OsSWEET14 cooperates with OsSWEET11 to contribute to grain filling in rice [J]. Plant Sci, 2021, 306: 110851.
[46]	Li GH, Zhou CY, Yang ZJ, et al. Low nitrogen enhances apoplastic phloem loading and improves the translocation of photoassimilates in rice leaves and stems [J]. Plant Cell Physiol, 2022, 63(7): 991-1007.
[47]	Gupta PK, Balyan HS, Gautam T. SWEET genes and TAL effectors for disease resistance in plants: Present status and future prospects [J]. Mol Plant Pathol, 2021, 22(8): 1014-1026.
[48]	Wang SD, Liu SL, Wang J, et al. Simultaneous changes in seed size, oil content and protein content driven by selection of SWEET homologues during soybean domestication [J]. Natl Sci Rev, 2020, 7(11): 1776-1786.
[49]	Bezrutczyk M, Hartwig T, Horschman M, et al. Impaired phloem loading in zmsweet13a, b, c sucrose transporter triple knock-out mutants in Zea mays [J]. New Phytol, 2018, 218(2): 594-603.
[50]	Sosso D, van der Linde K, Bezrutczyk M, et al. Sugar partitioning between Ustilago maydis and its host Zea mays L during infection [J]. Plant Physiol, 2019, 179(4): 1373-1385.
[51]	Wang SD, Yokosho K, Guo RZ, et al. The soybean sugar transporter GmSWEET15 mediates sucrose export from endosperm to early embryo [J]. Plant Physiol, 2019, 180(4): 2133-2141.
[52]	Zhai ZY, Liu H, Xu CC, et al. Sugar potentiation of fatty acid and triacylglycerol accumulation [J]. Plant Physiol, 2017, 175(2): 696-707.
[53]	Ho LH, Klemens PAW, Neuhaus HE, et al. SlSWEET1a is involved in glucose import to young leaves in tomato plants [J]. J Exp Bot, 2019, 70(12): 3241-3254.
[54]	Ko HY, Tseng HW, Ho LH, et al. Hexose translocation mediated by SlSWEET5b is required for pollen maturation in Solanum lycopersicum [J]. Plant Physiol, 2022, 189(1): 344-359.
[55]	Zhang XS, Feng CY, Wang MN, et al. Plasma membrane-localized SlSWEET7a and SlSWEET14 regulate sugar transport and storage in tomato fruits [J]. Hortic Res, 2021, 8(1): 186.
[56]	Ko HY, Ho LH, Neuhaus HE, et al. Transporter SlSWEET15 unloads sucrose from phloem and seed coat for fruit and seed development in tomato [J]. Plant Physiol, 2021, 187(4): 2230-2245.
[57]	付欢欢, 朱言宇, 舒鑫, 等. 草莓糖转运蛋白SWEET基因家族的筛选及表达分析 [J]. 合肥师范学院学报, 2024, 42(6): 34-39.
	Fu HH, Zhu YY, Shu X, et al. Screening and expression analysis of SWEET gene family of strawberry sugar transporter [J]. J Hefei Norm Univ, 2024, 42(6): 34-39.
[58]	Tian RR, Xu JY, Xu ZC, et al. Genome-wide identification and expression analysis of SWEET gene family in strawberry [J]. Horticulturae, 2024, 10(2): 191.
[59]	胡圣磊, 刘冬, 郭宝, 等. OsSTP1介导蔗糖分配调控水稻氮响应 [J]. 生物工程学报, 2024, 40(10): 3500-3514.
	Hu SL, Liu D, Guo B, et al. OsSTP1 mediates sucrose partitioning to regulate nitrogen response in rice [J]. Chin J Biotechnol, 2024, 40(10): 3500-3514.
[60]	黄子洋, 刘洁, 康婕, 等. 观赏植物糖转运蛋白研究进展 [J]. 浙江大学学报: 农业与生命科学版, 2024, 50(1): 12-24.
	Huang ZY, Liu J, Kang J, et al. Advances in studies on sugar transporters in ornamental plants [J]. Journal of Zhejiang University: Agriculture & Life Sciences, 2024, 50(1): 12-24.
[61]	Büttner M. The monosaccharide transporter (-like) gene family in Arabidopsis [J]. FEBS Lett, 2007, 581(12): 2318-2324.
[62]	Truernit E, Schmid J, Epple P, et al. The sink-specific and stress-regulated Arabidopsis STP4 gene: enhanced expression of a gene encoding a monosaccharide transporter by wounding, elicitors, and pathogen challenge [J]. Plant Cell, 1996, 8(12): 2169-2182.
[63]	Geiger D. Plant glucose transporter structure and function [J]. Pflugers Arch, 2020, 472(9): 1111-1128.
[64]	Büttner M. The Arabidopsis sugar transporter (AtSTP) family: an update [J]. Plant Biol, 2010, 12(): 35-41.
[65]	Schneidereit A, Scholz-Starke J, Büttner M. Functional characterization and expression analyses of the glucose-specific AtSTP9 monosaccharide transporter in pollen of Arabidopsis [J]. Plant Physiol, 2003, 133(1): 182-190.
[66]	Schneidereit A, Scholz-Starke J, Sauer N, et al. AtSTP11, a pollen tube-specific monosaccharide transporter in Arabidopsis [J]. Planta, 2005, 221(1): 48-55.
[67]	Nørholm MHH, Nour-Eldin HH, Brodersen P, et al. Expression of the Arabidopsis high-affinity hexose transporter STP13 correlates with programmed cell death [J]. FEBS Lett, 2006, 580(9): 2381-2387.
[68]	Lemonnier P, Gaillard C, Veillet F, et al. Expression of Arabidopsis sugar transport protein STP13 differentially affects glucose transport activity and basal resistance to Botrytis cinerea [J]. Plant Mol Biol, 2014, 85(4-5): 473-484.
[69]	Schneider S, Schneidereit A, Konrad KR, et al. Arabidopsis INOSITOL TRANSPORTER4 mediates high-affinity H⁺ symport of myoinositol across the plasma membrane [J]. Plant Physiol, 2006, 141(2): 565-577.
[70]	Schulz A, Beyhl D, Marten I, et al. Proton-driven sucrose symport and antiport are provided by the vacuolar transporters SUC4 and TMT1/2 [J]. Plant J, 2011, 68(1): 129-136.
[71]	Toyofuku K, Kasahara M, Yamaguchi J. Characterization and expression of monosaccharide transporters (OsMSTs) in rice [J]. Plant Cell Physiol, 2000, 41(8): 940-947.
[72]	Poschet G, Hannich B, Büttner M. Identification and characterization of AtSTP14, a novel galactose transporter from Arabidopsis [J]. Plant Cell Physiol, 2010, 51(9): 1571-1580.
[73]	Mamun EA, Alfred S, Cantrill LC, et al. Effects of chilling on male gametophyte development in rice [J]. Cell Biol Int, 2006, 30(7): 583-591.
[74]	Büttner M, Truernit E, Baier K, et al. AtSTP3, a green leaf-specific, low affinity monosaccharide-H ⁺ symporter of Arabidopsis thaliana [J]. Plant Cell Environ, 2000, 23(2): 175-184.
[75]	Cho JI, Burla B, Lee DW, et al. Expression analysis and functional characterization of the monosaccharide transporters, OsTMTs, involving vacuolar sugar transport in rice (Oryza sativa) [J]. New Phytol, 2010, 186(3): 657-668.
[76]	Cao H, Guo SY, Xu YY, et al. Reduced expression of a gene encoding a Golgi localized monosaccharide transporter (OsGMST1) confers hypersensitivity to salt in rice (Oryza sativa) [J]. J Exp Bot, 2011, 62(13): 4595-4604.
[77]	Li JM, Zheng DM, Li LT, et al. Genome-wide function, evolutionary characterization and expression analysis of sugar transporter family genes in pear (Pyrus bretschneideri Rehd) [J]. Plant Cell Physiol, 2015, 56(9): 1721-1737.
[78]	王丹琪, 张皓, 王鹏, 等. 梨STP基因家族鉴定及PbrSTP11调控梨花粉管生长的功能分析 [J].南京农业大学学报, 2022, 45(3): 493-502.
	Wang DQ, Zhang H, Wang P, et al. Identification of STP gene family in pear and functional analysis of PbrSTP11 in pollen tube growth regulation [J]. Journal of Nanjing Agricultural University, 2022, 45(3): 493-502.
[79]	Reuscher S, Akiyama M, Yasuda T, et al. The sugar transporter inventory of tomato: genome-wide identification and expression analysis [J]. Plant Cell Physiol, 2014, 55(6): 1123-1141.
[80]	Afoufa-Bastien D, Medici A, Jeauffre J, et al. The Vitis vinifera sugar transporter gene family: phylogenetic overview and macroarray expression profiling [J]. BMC Plant Biol, 2010, 10: 245.
[81]	Ruan YL, Jin Y, Yang YJ, et al. Sugar input, metabolism, and signaling mediated by invertase: roles in development, yield potential, and response to drought and heat [J]. Mol Plant, 2010, 3(6): 942-955.
[82]	Wu WQ, Du K, Kang XY, et al. The diverse roles of cytokinins in regulating leaf development [J]. Hortic Res, 2021, 8(1): 118.
[83]	Li B, Liu H, Zhang Y, et al. Constitutive expression of cell wall invertase genes increases grain yield and starch content in maize [J]. Plant Biotechnol J, 2013, 11(9): 1080-1091.
[84]	Wu Y, Chen HN, Wu MB, et al. A vacuolar invertase gene SlVI modulates sugar metabolism and postharvest fruit quality and stress resistance in tomato [J]. Hortic Res, 2024, 12(1): uhae283.
[85]	Ruan YL. CWIN-sugar transporter nexus is a key component for reproductive success [J]. J Plant Physiol, 2022, 268: 153572.
[86]	Paul MJ, Primavesi LF, Jhurreea D, et al. Trehalose metabolism and signaling [J]. Annu Rev Plant Biol, 2008, 59: 417-441.
[87]	Baud S, Vaultier MN, Rochat C. Structure and expression profile of the sucrose synthase multigene family in Arabidopsis [J]. J Exp Bot, 2004, 55(396): 397-409.
[88]	Wang YT, Wang XW, Xie J, et al. Identification and gene mapping of an early senescent leaf mutant esl11 of rice [J]. Crop Sci, 2018, 58(5): 1932-1941.
[89]	Li ZW, Zhao Q, Cheng FM. Sugar starvation enhances leaf senescence and genes involved in sugar signaling pathways regulate early leaf senescence in mutant rice [J]. Rice Sci, 2020, 27(3): 201-214.