园艺作物糖转运蛋白研究进展

doi:10.16420/j.issn.0513-353x.2020-0412

摘要/Abstract

摘要：

糖转运蛋白在园艺作物生长发育、果实内糖积累以及响应逆境胁迫等方面发挥重要作用,其表达受多种因素的调节。单糖转运蛋白、蔗糖转运蛋白和SWEET（bidirectional sugar transporter,SWEET）是植物中已发现的三大类转运蛋白。就这3种转运蛋白运输活性与底物特异性、组织/器官和细胞定位进行了讨论和比较;重点介绍了园艺作物中糖转运蛋白功能和调控的最新研究进展;进一步分析糖信号、生物胁迫和非生物胁迫等对糖转运蛋白表达的调控。

关键词: 园艺作物, 单糖转运蛋白, 蔗糖转运蛋白, SWEETs

Abstract:

Sugar transporters play important roles in the growth and development,fruits sugar accumulation and response to stress in horticultural crops,and their expression is regulated by many factors. Monosaccharide transporters,sucrose transporters and SWEETs are three kinds of transporters in plants. In this study,the transport activity and substrate specificity,tissue and cell localization of the three types of transporters were discussed and compared,with emphasis on the latest progress in the function and regulation of sugar transporters in horticultural crops. What’s more,the regulation of sugar transporter expression by sugar signal,biotic stress and abiotic stress was analyzed.

Key words: horticultural crop, monosaccharide transporters, sucrose transporters, SWEETs

中图分类号:

耿艳秋, 董肖昌, 张春梅. 园艺作物糖转运蛋白研究进展[J]. 园艺学报, 2021, 48(4): 676-688.

GENG Yanqiu, DONG Xiaochang, ZHANG Chunmei. Recent Progress of Sugar Transporter in Horticultural Crops[J]. Acta Horticulturae Sinica, 2021, 48(4): 676-688.

图/表 1

图1 糖转运蛋白的表达调控途径糖转运蛋白的表达受植物自身和外界因素的共同调控,在糖、激素、低温、干旱及盐等信号刺激下,转录因子与糖转运蛋白启动子区顺式作用元件中相应信号响应因子结合后,可诱导或抑制糖转运蛋白基因的转录,mRNA经核孔进入核糖体加工后翻译为糖转运蛋白,这些糖转运蛋白可被分配至细胞质膜或液泡膜行使相应的生理功能（Conde et al.2006;Fan et al.2009;Li et al.2015;Ren et al.2018;Peng et al.2020b）。

Fig. 1 Expression regulatory pathways of sugar transporters The expression regulation of sugar transporter is affected by both plant itself and external factors. Under the stimulation of sugar,hormones,low temperature,drought or salt,the transcription factor binds to the corresponding signal response factor in the cis-acting element in the promoter region,then it will induce or inhibit the transcription of sugar transporter genes. mRNA enters the ribosome through the nuclear pore for processing and then will be translated into sugar transporters. These sugar transporters can be distributed to the plasma membrane or vacuolar membrane to perform physiological functions（Conde et al.2006;Fan et al.2009;Li et al.2015;Ren et al.2018;Peng et al.2020b）.

参考文献 86

[1]	Afoufa-Bastien D, Medici A, Jeauffre J, Coutos-Thévenot P, Lemoine R, Atanassova R, Laloi M. 2010. The Vitis vinifera sugar transporter gene family:phylogenetic overview and macroarray expression profiling. BioMed Central, 10 (1):doi: 10.1186/1471-2229-10-245.
[2]	Atanassova R, Leterrier M, Gaillard C, Agasse A, Sagot E, Coutos-Thévenot P, Delrot S. 2003. Sugar-regulated expression of a putative hexose transport gene in grape. Plant Physiology, 131 (1):326-334. doi: 10.1104/pp.009522 URL
[3]	Barker L, Kühn C, Weise A, Schulz A, Gebhardt C, Hirner B, Hellmann H, Schulze W, Ward J M, Frommer W B. 2000. SUT2,a putative sucrose sensor in sieve elements. Plant Cell, 12 (7):1153-1164. pmid: 10899981
[4]	Burkle L, Hibberd J M, Quick W P, Kuhn C, Hirner B, Frommer W B. 1998. The H⁺-sucrose cotransporter NtSUT 1 is essential for sugar export from tobacco leaves. Plant Physiology, 118 (1):59-68. pmid: 9733526
[5]	Büttner M. 2007. The monosaccharide transporter(-like)gene family in Arabidopsis. Federation of European Biochemical Societies, 581:2318-2324. doi: 10.1016/j.febslet.2007.03.016 URL
[6]	Büttner M. 2010. The Arabidopsis sugar transporter(AtSTP)family:an update. Plant Biology, 12:35-41.
[7]	Cakir B, Agasse A, Gaillard C, Saumonneau A, Delrot S, Atanassova R. 2003. A grape ASR protein involved in sugar and abscisic acid signaling. The Plant Cell, 15 (9):2165-2180. doi: 10.1105/tpc.013854 URL
[8]	Chandran D, Reinders A, Ward J M. 2003. Substrate specificity of the Arabidopsis thaliana sucrose transporter AtSUC2. The Journal of Biological Chemistry, 278 (45):44320-44325. doi: 10.1074/jbc.M308490200 URL
[9]	Chardon F, Bedu M, Calenge F, Klemens P A W, Spinner L, Clement G, Chietera G, Léran S, Ferrand M, Lacombe B, Loudet O, Dinant S, Bellini C, Neuhaus H E, Daniel-Vedele F, Krapp A. 2013. Leaf fructose content is controlled by the vacuolar transporter SWEET 17 in Arabidopsis. Current Biology, 23 (8):697-702. doi: 10.1016/j.cub.2013.03.021 URL
[10]	Chen Jun-wei, Zhang Shang-long, Zhang Liang-cheng. 2004. Transport,metabolism and accumulation of sugar in fruit and its regulation. Journal of Plant Physiology and Molecular Biology, 30:1-10. (in Chinese)
	陈俊伟, 张上隆, 张良诚. 2004. 果实中糖的运输、代谢与积累及其调控. 植物生理与分子生物学学报, 30:1-10.
[11]	Chen L Q. 2014. SWEET sugar transporters for phloem transport and pathogen nutrition. New phytologist, 201 (4):1150-1155. doi: 10.1111/nph.2014.201.issue-4 URL
[12]	Chen L Q, Qu X Q, Hou B H, Sosso D, Osorio S, Fernie A R, Frommer W B. 2012. Sucrose efflux mediated by SWEET proteins as a key step for phloem transport. Science, 335 (6065):207-211. doi: 10.1126/science.1213351 URL
[13]	Cheng R, Cheng Y S, Lü J H, Chen J Q, Wang Y Z, Zhang S L, Zhang H P. 2018a. The gene PbTMT4from pear( Pyrus bretschneideri) mediates vacuolar sugar transport and strongly affects sugar accumulation in fruit. Physiologia Plantarum,doi: 10.1111/ppl.12742.
[14]	Cheng J T, Wen S Y, Xiao S, Lu B Y, Ma M, Bie Z. 2018b. Overexpression of the tonoplast sugar transporter CmTST 2 in melon fruit increases sugar accumulation. Journal of Experimental Botany, 69 (3):511-523. doi: 10.1093/jxb/erx440 URL
[15]	Cheng Yinsheng, Chen Jianqiu, Chen Dan, Lü Jiahong, Zhang Jun, Zhang Shaoling, Wu Tao, Zhang Huping. 2019. Cloning and functional analysis of the promoter of PbTMT4 gene related sugar transport in pear. Acta Horticulturae Sinica, 46 (1):25-36. (in Chinese)
	程寅胜, 陈健秋, 陈丹, 吕佳红, 张俊, 张绍铃, 伍涛, 张虎平. 2019. 梨糖转运相关基因 PbTMT4启动子克隆及功能分析. 园艺学报, 46 (1):25-36.
[16]	Chong J L, Piron M C, Meyer S, Merdinoglu D, Bertsch C, Mestre P. 2014. The SWEET family of sugar transporters in grapevine:VvSWEET 4 is involved in the interaction with Botrytis cinerea. Journal of Experimental Botany, 65 (22):6589-6601. doi: 10.1093/jxb/eru375 URL
[17]	Christina K, Christopher P G. 2010. Sucrose transporters of higher plants. Plant Biology, 13 (3):287-297.
[18]	Conde C, Agasse A, Glissant D, Tavares R, Gerós H, Delrot S. 2006. Pathways of glucose regulation of monosaccharide transport in grape cells. Plant Physiology, 141 (4):1563-1577. doi: 10.1104/pp.106.080804 URL
[19]	Dai Mei-song, Xu Fei, Shi Ze-bin, Xu Chang-jie. 2015. Preliminary study on expression characteristics of sorbitol transporter(SOT)gene family and the role in sugar accumulation in Pyrus pyrifolia fruits. Acta Horticulturae Sinica, 42 (8):1457-1466. (in Chinese)
	戴美松, 徐飞, 施泽彬, 徐昌杰. 2015. 砂梨山梨醇转运蛋白(SOT)基因家族成员表达特性及在果实糖积累中的作用初探. 园艺学报, 42 (8):1457-1466.
[20]	Engel M L, Holmes-Davis R, McCormick S G. 2005. Identification of male gamete promoters in Arabidopsis. Plant Physiol, 138:2124-2133. doi: 10.1104/pp.104.054213 URL
[21]	Fan R C, Peng C C, Xu Y H, Wang X F, Li Y, Shang Y, Du S Y, Zhao R, Zhang X Y, Zhang L Y, Zhang D P. 2009. Apple sucrose transporter SUT1 and sorbitol transporter SOT 6 interact with cytochrome b5 to regulate their affinity for substrate sugars. Plant Physiology, 150 (4):1880-1901. doi: 10.1104/pp.109.141374 URL
[22]	Fang T, Peng Y, Rao Y, Li S H, Zeng L H. 2020. Genome-wide identification and expression analysis of Sugar Transporter(ST)gene family in longan( Dimocarpus longan L.). Plants,doi: 10.3390/plants9030342.
[23]	Fei Guang-xue. 2018. Studies on sugar accumulation and activities of related metabolic enzymes in Junzao fruit. Xinjiang Land Reclamation Science and Technology, 41:28-31. (in Chinese)
	费光雪. 2018. 骏枣果实糖分积累及相关代谢酶活性变化的研究. 新疆农垦科技, 41:28-31.
[24]	Gao Z. 2003. Cloning,expression,and characterization of sorbitol transporters from developing sour cherry fruit and leaf sink tissues. Plant Physiology, 131 (4):1566-1575. doi: 10.1104/pp.102.016725 URL
[25]	Gebauer P, Korn M, Engelsdorf T, Sonnewald U, Koch C, Voll L M. 2017. Sugar accumulation in Leaves of Arabidopsis SWEET11/SWEET 12 double mutants enhances priming of the salicylic acid-mediated defense response. Frontiers in Plant Science,doi: 10.3389/fpls.2017.01378.
[26]	Geng Y, Wu M, Zhang C. 2020. Sugar transporter ZjSWEET2.2 mediates sugar loading in leaves of Ziziphus jujuba Mill. Frontiers in Plant Science, 11:1081. doi: 10.3389/fpls.2020.01081 URL
[27]	Gottwald J R, Krysan P J, Young J C, Evert R F, Sussman M R. 2000. Genetic evidence for the in planta role of phloem-specific plasma membrane sucrose transporters. Proceedings of the National Academy of Sciences of the United States of America, 97 (25):13979-13984.
[28]	Guan Y F, Huang X Y, Zhu J, Gao J F, Zhang H X, Yang Z N. 2008. Ruptured pollen erain1,a member of the MtN3/saliva gene family,is crucial for exine pattern formation and cell integrity of microspores in Arabidopsis. Plant Physiology, 147 (2):852-863. doi: 10.1104/pp.108.118026 URL
[29]	Hackel A, Schauer N, Carrari F, Fernie A, Grimm B, Kühn C. 2006. Sucrose transporter LeSUT1 and LeSUT 2 inhibition affects tomato fruit development in different ways. The Plant Journal, 45 (2):180-192. doi: 10.1111/tpj.2006.45.issue-2 URL
[30]	Hayes M A, Davies C, Dry I B. 2007. Isolation,functional characterization,and expression analysis of grapevine( Vitis vinifera L.)hexose transporters:differential roles in sink and source tissues. Journal of Experimental Botany, 58 (8):1985-1997. doi: 10.1093/jxb/erm061 URL
[31]	Hayes M A, Feechan A, Dry I B. 2010. Involvement of abscisic acid in the coordinated regulation of a stress-inducible hexose transporter(VvHT5) and a cell wall invertase in grapevine in response to biotrophic fungal infection. Plant Physiology, 153 (1):211-221. doi: 10.1104/pp.110.154765 URL
[32]	Ho L H, Klemens P A W, Neuhaus H E, Ko H Y, Hsieh S Y, Guo W J. 2019. SlSWEET1a is involved in glucose import to young leaves in tomato plants. Journal of Experimental Botany,doi: 10.1093/jxb/erz154.
[33]	Huang W F, Hu B, Liu J L, Zhou Y, Liu S Q. 2020. Identification and characterization of tonoplast sugar transporter(TST) gene family in cucumber. Horticultural Plant Journal, 6 (3):145-157. doi: 10.1016/j.hpj.2020.03.005 URL
[34]	Jia H F, Wang Y H, Sun M Z, Li B B, Han Y. 2013. Sucrose functions as a signal involved in the regulation of strawberry fruit development and ripening. New Phytologist, 198 (2):453-465. doi: 10.1111/nph.2013.198.issue-2 URL
[35]	Jia Hai-feng. 2013. Role and mechanism of sucrose and jasmonic acid signals in the regulation of strawberry fruit development [Ph. D. Dissertation]. Beijing:China Agricultural University. (in Chinese)
	贾海锋. 2013. 蔗糖及茉莉酸信号在草莓果实发育中的作用及其机理分析[博士论文]. 北京:中国农业大学.
[36]	Jiang W Z, Zhou H B, Bi H H, Fromm M, Yang B, Weeks D P. 2013. Demonstration of CRISPR/Cas9/sgRNA-mediated targeted gene modification in Arabidopsis,tobacco,sorghum and rice. Nucleic Acids Research,doi: 10.1093/nar/gkt780.
[37]	Jung B, Ludewig F, Schulz A, Meißner G, Wöstefeld N, Flügge U, Pommerrenig B, Wirsching P, Sauer No, Koch W, Sommer F, Mühlhaus T, Schroda M, Cuin T A, Graus D, Marten I, Hedrich R, Neuhaus H E. 2015. Identification of the transporter responsible for sucrose accumulation in sugar beet taproots. Nature Plants,doi: 10.1038/nplants.2014.1.
[38]	Kafkas E. 2006. Analysis of sugars,organic acids and vitamin C contents of blackberry genotypes from Turkey. Food Chemistry, 97 (4):732-736. doi: 10.1016/j.foodchem.2005.09.023 URL
[39]	Klemens P A W, Patzke K, Deitmer J, Spinner L, Le H R, Bellini C, Bedu M, Chardon F, Krapp A, Neuhaus H E. 2013. Overexpression of the vacuolar sugar carrier AtSWEET 16 modifies germination,growth,and stress tolerance in Arabidopsis. Plant Physiology, 163 (3):1338-1352. doi: 10.1104/pp.113.224972 URL
[40]	Koch K E, Avigne W T. 1990. Postphloem,nonvascular transfer in citrus:kinetic,metabolism,and sugar gradients. Plant Physiology, 93 (4):1405-1416. doi: 10.1104/pp.93.4.1405 URL
[41]	Kryvoruchko I S, Sinharoy S, Torres-Jerez I, Sosso D, Pislariu C I, Guan D, Murray J, Benedito V A, Frommer W B, Udvardi M K. 2016. MtSWEET11,a nodule-specific sucrose transporter of Medicago truncatula. Plant Physiology, 171 (1):554-565. doi: 10.1104/pp.15.01910 pmid: 27021190
[42]	Kühn C, Grof C P L. 2010. Sucrose transporters of higher plants. Current Opinion in Plant Biology, 13 (3):287-297. doi: 10.1016/j.pbi.2010.02.001 URL
[43]	Lalonde S, Wipf D, Frommer W B. 2004. Transport mechanisms for organic forms of carbon and nitrogen between source and sink. Annual Review of Plant Biology, 55:341-372. pmid: 15377224
[44]	Lemonnier P, Gaillard C, Veillet F, Verbeke J, Lemoine R, Coutos-Thévenot Pierre, La Camera S. 2014. Expression of Arabidopsis sugar transport protein STP 13 differentially affects glucose transport activity and basal resistance to Botrytis cinerea. Plant Molecular Biology, 85 (4-5):473-484.
[45]	Leterrier M, Atanassova R, Laquitaine L, Gaillard C, Coutos-Thévenot P, Delrot S. 2003. Expression of a putative grapevine hexose transporter in tobacco alters morphogenesis and assimilate partitioning. Pubmed, 54 (385):1193-1204.
[46]	Li C, Meng D, Piñeros M A, Mao Y, Dandekar A M, Cheng L. 2020a. A sugar transporter takes up both hexose and sucrose for sorbitol-modulated in vitro pollen tube growth in apple. The Plant Cell, 32 (2):449-469. doi: 10.1105/tpc.19.00638 URL
[47]	Li J M, Qin M F, Qiao X, Cheng Y S, Li X L, Zhang H P, Wu J. 2017. A new insight into the evolution and functional divergence of SWEET transporters in Chinese white pear (Pyrus bretschneideri). Plant & Cell Physiology, 58 (4):839-850.
[48]	Li J M, Zheng D M, Li L T, Qiao X, Wei S W, Bai B, Zhang S L, Wu J. 2015. Genome-wide function,evolutionary characterization and expression analysis of sugar transporter family genes in pear( Pyrus bretschneideri Rehd). Plant & Cell Physiology, 56 (9):1721-1737.
[49]	Li X Y, Guo W, Li J C, Yue P T, Bu H D, Jiang J, Liu W T, Xu Y X, Yuan H, Li T, Wang A. 2020b. Histone acetylation at the promoter for the transcription factor PuWRKY 31 affects sucrose accumulation in pear fruit. Plant Physiology,doi: 10.1104/pp.20.00002.
[50]	Lin I W, Sosso D, Chen L Q, Gase K, Kim S, Kessler D, Klinkenberg P, Gorder M K, Hou B H, Qu X Q, Carter C J, Baldwin I T, Frommer W B. 2014. Nectar secretion requires sucrose phosphate synthases and the sugar transporter SWEET9. Nature, 508 (7497):546-549. doi: 10.1038/nature13082 URL
[51]	Lu J, Sun M H, Ma Q J, Kang H, Liu Y J, Hao Y J, You C X. 2019. MdSWEET17,a sugar transporter in apple,enhances drought tolerance in tomato. Journal of Integrative Agriculture, 18 (9):2041-2051. doi: 10.1016/S2095-3119(19)62695-X URL
[52]	Lu Jing, Ma Qijun, Kang Hui, Li Wenhao, Liu Yajing, Hao Yujin, You Chunxiang. 2019. Ectopic expressing MdSWEET1 in tomato enhanced salt tolerance. Acta Horticulturae Sinica, 46 (3):433-443. (in Chinese)
	路静, 马齐军, 康慧, 李文浩, 刘亚静, 郝玉金, 由春香. 2019. 苹果糖转运蛋白基因 MdSWEET1在番茄中异源表达提高其耐盐性. 园艺学报, 46 (3):433-443.
[53]	Lu M Z, Snyder R, Grant J, Tegeder M. 2020. Manipulation of sucrose phloem and embryo loading affects pea leaf metabolism,carbon and nitrogen partitioning to sinks as well as seed storage pools. The Plant Journal:for Cell and Molecular Biology,doi: 10.1111/tpj.14533.
[54]	Ma Li-xin, Qin Yuan, Wei Xiao-yu, Ma Feng-wang, Li Ming-jun. 2014. Expression of apple sugar transporter TMT gene and its relationship with sugar accumulation. Acta Horticulturae Sinica, 41 (7):1317-1325. (in Chinese)
	马新立, 秦源, 魏晓钰, 马锋旺, 李明军. 2014. 苹果糖转运蛋白TMT基因的表达及其与糖积累的关系. 园艺学报, 41 (7):1317-1325.
[55]	Ma Q J, Sun M H, Kang H, Lu J, You C X, Hao Y J. 2019. A CIPK protein kinase targets sucrose transporter MdSUT2.2 at Ser for phosphorylation to enhance salt tolerance. Plant Cell and Environment, 42 (3):918-930. doi: 10.1111/pce.v42.3 URL
[56]	Ma Q J, Sun M H, Lu J, Liu Y J, Hu D G, Hao Y J. 2017. Transcription factor AREB 2 is involved in soluble sugar accumulation by activating sugar transporter and amylase genes. Plant Physiology, 174 (4):2348-2362. doi: 10.1104/pp.17.00502 URL
[57]	Moriyama E N, Strope P K, Opiyo S O, Chen Z Y, Jones A M. 2006. Mining the Arabidopsis thaliana genome for highly-divergent seven transmembrane receptors. Genome Biology,doi: 10.1186/gb-2006-7-10-r96.
[58]	Ni J P, Li J M, Zhu R X, Zhang M Y, Qi K J, Zhang S L, Wu J. 2020. Overexpression of sugar transporter gene PbSWEET4 of pear causes sugar reduce and early senescence in leaves. Gene,743. doi: 10.1016/j.gene.2020.144582.
[59]	Nie Pei-xian, Li Chen, Gao Yan, Du Guo-dong, Lü De-guo, Guan Qiu-zhu. 2019. Ultrastructural observation and functional analysis of phloem and parenchyma cells around the fruit of Ziziphus jujuba. Journal of Chinese Electron Microscopy Society, 55 (6):697-702. (in Chinese)
	聂佩显, 李晨, 高艳, 杜国栋, 吕德国, 关秋竹. 2009. 光合同化物韧皮部卸载途径的研究进展. 植物生理学报, 55 (6):697-702.
[60]	Peng C C, Xu Y H, Xi R C, Zhao X L. 2011. Expression,subcellular localization and phytohormone stimulation of a functional sucrose transporter (MdSUT1)in apple fruit. Scientia Horticulturae, 128 (3):206-212. doi: 10.1016/j.scienta.2011.01.019 URL
[61]	Peng Chang-cao, Xiao Wen-fang, Li Wen-yan, Zhao Xiao-lan. 2011. Polyclonal antibody preparing and specific expression analyzing of sucrose transporter MdSUT 1 in apple. Journal of Plant Physiology, 47 (3):256-262. (in Chinese)
	彭昌操, 肖文芳, 李文燕, 赵小兰. 2011. 苹果蔗糖载体蛋白MdSUT1多克隆抗体的制备及MdSUT1蛋白表达特异性分析. 植物生理学报, 47 (3):256-262.
[62]	Peng Q, Cai Y M, Lai E H, Nakamura M, Liao L, Zheng B B, Ogutu C, Cherono S, Han Y P. 2020a. The sucrose transporter MdSUT4.1 participates in the regulation of fruit sugar accumulation in apple. BMC Plant Biology,doi: 10.1186/s12870-020-02406-3.
[63]	Peng Q, Wang L, Ogutu C, Liu J J, Liu L, Mollah M D A, Han Y. 2020b. Functional analysis reveals the regulatory role of PpTST 1 encoding tonoplast sugar transporter in sugar accumulation of peach fruit. International Journal of Molecular Sciences,doi: 10.3390/ijms21031112.
[64]	Pommerrenig B, Müdsam C, Kischka D, Neuhaus H E. 2020. Treat and trick:common regulation and manipulation of sugar transporters during sink establishment by the plant and the invader. Journal of Experimental Botany,doi: 10.1093/jxb/eraa168.
[65]	Ren Y, Guo S, Zhang J, He H, Sun H, Tian S, Gong G, Zhang H, Levi A, Tadmor Y, Xu Y. 2018. A tonoplast sugar transporter underlies a sugar accumulation QTL in watermelon. Plant Physiology, 176 (1):836-850. doi: 10.1104/pp.17.01290 pmid: 29118248
[66]	Riesmeier J W, Willmitzer L, Frommer W B. 1992. Isolation and characterization of a sucrose carrier cDNA from spinach by functional expression in yeast. The EMBO Journal, 11 (13):4705-4713. doi: 10.1002/embj.1992.11.issue-13 URL
[67]	Ruan Y L, Patrick J W, Brady C. 1997. Protoplast hexose carrier activity is a determinate of genotypic difference in hexose storage in tomato fruit. Plant,Cell & Environment, 20 (3):341-349.
[68]	Schulz A, Beyhl D, Marten I, Wormit A, Neuhaus E, Poschet G, Büttner M, Schneider S, Sauer N, Hedrich R. 2011. Proton-driven sucrose symport and antiport are provided by the vacuolar transporters SUC4 and TMT1/2. The Plant Journal, 68 (1):129-136. doi: 10.1111/j.1365-313X.2011.04672.x URL
[69]	Seo P J, Park J M, Kang S K, Kim S G, Park C M. 2011. An Arabidopsis senescence-associated protein SAG 29 regulates cell viability under high salinity. Planta, 233 (1):189-200. doi: 10.1007/s00425-010-1293-8 URL
[70]	Slewinski T L. 2011. Diverse functional roles of monosaccharide transporters and their homologs in vascular plants:a physiological perspective. Molecular Plant, 4 (4):641-662. doi: 10.1093/mp/ssr051 URL pmid: 21746702
[71]	Sonnewald U. 2011. Sweets-the missing sugar efflux carriers. Frontiers in Plant Science,doi: 10.3389/fpls.2011.00007.
[72]	Sun M X, Huang X Y, Yang J, Guan Y F, Yang Z N. 2013. Arabidopsis RPG 1 is important for primexine deposition and functions redundantly with RPG2 for plant fertility at the late reproductive stage. Plant Reproduction, 26 (2):83-91. doi: 10.1007/s00497-012-0208-1 URL
[73]	Tang Juan. 2013. Cloning,characteristics of study and sequence analysis of sucrose and phosphatidylcholine transporter protein genes in pear[M. D. Dissertation]. Nanjing:Nanjing Agricultural University. (in Chinese)
	唐娟. 2013. 梨蔗糖与磷脂酰胆碱转运蛋白基因的克隆、序列分析及特性研究[硕士论文]. 南京:南京农业大学.
[74]	Watari J, Yoshihiro K, Shohei Y, Kunio Y, Kyoko T, Tabuchi T, Katsuhiro S. 2004. Identification of sorbitol transporters expressed in the phloem of apple source leaves. Plant Cell Physiology, 45 (8):1032-1041. doi: 10.1093/pcp/pch121 URL
[75]	Wang L F, Qi X X, Huang X S, Xu L L, Jin C, Wu J, Zhang S L. 2016. Overexpression of sucrose transporter gene PbSUT2 from P yrus bretschneideri,enhances sucrose content in Solanum lycopersicum fruit. Plant Physiology and Biochemistry, 105:150-161. doi: 10.1016/j.plaphy.2016.04.019 URL
[76]	Wang Jun-gang, Zhao Ting-ting, Yang Ben-peng, Huang Cai-xia. 2007. Monosaccharide transporters in plants. Plant Physiology Communications, 43 (6):1195-1201. (in Chinese)
	王俊刚, 赵婷婷, 张树珍, 杨本鹏, 黄彩霞. 2007. 植物单糖转运蛋白. 植物生理学通讯, 43 (6):1195-1201.
[77]	Wang Man-man, Qu Shu-ping, Li Jun-xing, Wu Yan-quan, Huang He-xun, Zhong Yu-juan. 2019. Research progress on sugar accumulation and regulation of Cucurbitaceae crops. Journal of Plant Physiology, 55 (7):941-948. (in Chinese)
	王曼曼, 屈淑平, 李俊星, 吴廷全, 黄河勋, 钟玉娟. 2019. 葫芦科作物果实糖积累及其调控研究进展. 植物生理学报, 55 (7):941-948.
[78]	Weise A, Barker L, Kühn C, Lalonde S, Buschmann H, Frommer W B, Ward J M. 2000. A new subfamily of sucrose transporters,SUT4,with low affinity/high capacity localized in enucleate sieve elements of plants. The Plant Cell, 12 (8):1345-1355. doi: 10.1105/tpc.12.8.1345 URL
[79]	Wei Xin-na. 2016. Functional indentification of sucrose transporters and protein expression analysis of PpSUT2 from peach[Ph. D. Dissertation]. Zhengzhou:Henan Agricultural University. (in Chinese)
	魏新娜. 2016. 桃蔗糖转运蛋白基因的功能验证与PpSUT2蛋白表达分析[硕士论文]. 郑州:河南农业大学.
[80]	Xu Hai-feng, Qu Chang-zhi, Liu Jing-xuan, Wang Yi-cheng, Wang De-yun, Zuo Wei-fang, Jiang Sheng-hui, Wang Nan, Zhang Zong-ying, Chen Xue-sen. 2017. Expression analysis and functional identification of a vacuolar sucrose transporter gene MdSUT4 in apple. Acta Horticulturae Sinica, 44 (7):1235-1243. (in Chinese)
	许海峰, 曲常志, 刘静轩, 王意程, 王得云, 左卫芳, 姜生辉, 王楠, 张宗营, 陈学森. 2017. 苹果液泡膜蔗糖转运蛋白基因 MdSUT4的表达分析与功能鉴定. 园艺学报, 44 (7):1235-1243.
[81]	Yang Guan-xian, Xu Hai-feng, Zhang Jing, Wang Nan, Fang Hong-cheng, Zou Qi, Jiang Yong-sheng, Chen Xue-sen. 2018. Functional identification of a sugar transporter gene MdSWEET17 in apple. Journal of Plant Physiology, 54 (11):1737-1745. (in Chinese)
	杨官显, 许海峰, 张静, 王楠, 房鸿成, 邹琦, 王意程, 姜生辉, 陈学森. 2018. 苹果糖转运蛋白基因 MdSWEET17的功能鉴定. 植物生理学报, 54 (11):1737-1745.
[82]	Yao L C, Shi J X, Weiss D, Goldschmidt E E. 2003. Sugars regulate sucrose transporter gene expression in citrus. Biochemical and Biophysical Research Communications, 306 (2):402-407. doi: 10.1016/S0006-291X(03)00978-1 URL
[83]	Zhang H, Zhang S, Qin G, Wang L, Wu T, Qi K, Zhang S. 2013. Molecular cloning and expression analysis of a gene for sucrose transporter from pear( Pyrus bretschneideri)fruit. Plant Physiol Biochem, 73:63-69. doi: 10.1016/j.plaphy.2013.08.009 URL
[84]	Zhang L Y, Peng Y B, Pelleschi T S, Fan Y, Lu Y M, Lu Y F, Gao X, Shen Y, Delort S, Zhang D. 2004. Evidence for apoplasmic phloem unloading in developing apple fruit. Plant Physiology, 135 (1):574-586. doi: 10.1104/pp.103.036632 URL
[85]	Zheng Q M, Tang Z, Xu Q, Deng X X. 2014. Isolation,phylogenetic relationship and expression profiling of sugar transporter genes in sweet orange (Citrus sinensis). Plant Cell Tissue Organ Culture, 119 (3):609-624. doi: 10.1007/s11240-014-0560-y URL
[86]	Zhou Y, Liu L, Huang W F, Yuan M, Zhou F, Li X G, Lin Y J. 2014. Overexpression of OsSWEET5 in rice causes growth retardation and precocious senescence. PLoS ONE,doi: 10.1371/journal.pone.0094210.