苹果蔗糖转运蛋白MdSUT2调控花青苷积累的研究
|
摘要
从‘嘎拉’苹果(Malus×domestica Borkh.)中克隆蔗糖转运蛋白基因MdSUT2(序列号:MDP0000277235),该基因包含1 704 bp开放阅读框,编码566个氨基酸,蛋白大小为56 kD。构建MdSUT2表达载体并利用农杆菌介导法侵染拟南芥,半定量RT-PCR证明Md SUT2在拟南芥中异源表达。构建瞬时表达载体,‘红星’苹果进行Md SUT2瞬时表达,其花青苷含量增多,而沉默该基因花青苷含量明显降低。
A gene named MdSUT2(MDP0000277235)was cloned from‘Gala'apple(Malus × domestica Borkh.). Sequence analysis showed that this gene contained 1 704 bp nucleotides,and encoded a protein of 566 amino acids. Overexpression vector of MdSUT2 was constructed and transformed into Arabidopsis seedlings via Agrobacterium-mediated method. Semi RT-PCR analysis indicated that MdSUT2 was genetically transformed into Arabidopsis. In addition, the transient expression vectors were constructed and transformed into apple fruits. We found that overexpression of MdSUT2 in apple fruits significantly increased the anthocyanin content, whereas suppression of MdSUT2 decreased the anthocyanin content.
A gene named MdSUT2(MDP0000277235)was cloned from‘Gala'apple(Malus × domestica Borkh.). Sequence analysis showed that this gene contained 1 704 bp nucleotides,and encoded a protein of 566 amino acids. Overexpression vector of MdSUT2 was constructed and transformed into Arabidopsis seedlings via Agrobacterium-mediated method. Semi RT-PCR analysis indicated that MdSUT2 was genetically transformed into Arabidopsis. In addition, the transient expression vectors were constructed and transformed into apple fruits. We found that overexpression of MdSUT2 in apple fruits significantly increased the anthocyanin content, whereas suppression of MdSUT2 decreased the anthocyanin content.
引文
Arabidopsis GenomeⅠ.2000.Analysis of the genome sequence of the flowering plant Arabidopsis thaliana.Nature,408(6814):796-815.
Aoki N,Hirose T,Scofield G N,Whitfeld P R,Furbank R T.2003.The sucrose transporter gene family in rice.Plant and Cell Physiology,44(3):223-232.
Aoki N,Whitfeld P,Hoeren F,Scofield G,Newell K,Patrick J.2002.Three sucrose transporter genes are expressed in the developing grain of hexaploid wheat.Plant Molecular Biology,50(3):453-462.
An Xiu-hong.2013.Studies on the mechanism by which MdTTG1,MdMYB9 and MdMYB11 regulate anthocyanin biosynthesis in apple[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)安秀红.2013.苹果MdTTG1、MdMYB9与Md MYB11基因调控花青苷合成的作用机制研究[博士论文].泰安:山东农业大学.
Bai Xue-mei,Zhang Li-jun,Wu Xiao-dan,Hu Kai,Ruan Yan-ye.2006.Sucrose transporters in plants.Plant Physiology Communications,42(6):195-202.(in Chinese)白雪梅,张立军,吴晓丹,胡凯,阮燕晔.2006.植物蔗糖转运蛋白.植物生理学通,42(6):195-202.
Baier M,Hemmann G,Holman R,Corke F,Card R,Smith C,Rook F,Bevan M W.2004.Characterization of mutants in Arabidopsis showing increased sugar-specific gene expression,growth,and developmental responses.Plant Physiology,134(1):81-91.
Bassolino L,Zhang Y,Schoonbeek H J,Kiferle C,Perata P,Martin C.2013.Accumulation of anthocyanins in tomato skin extends shelf life.New Phytologist,200(3):650-655.
Baud S,Wuilleme S,Lemoine R.2005.The AtSUC5 sucrose transporter specifically expressed in the endosperm is involved in early seed development in Arabidopsis.Plant Journal,43:824-836.
Butelli E,Titta L,Giorgio M,Mock H P,Matros A,Peterek S,Schijlen E G,Hall R D,Bovy A G,Luo J,Martin C.2008.Enrichment of tomato fruit with health-promoting anthocyanins by expression of select transcription factors.Nature Biotechnology,26(11):1301-1308.
Chen H J,Wang S J.2008.Molecular regulation of sink-source transition in rice leaf sheaths during the heading period.Acta Physiologiae Plantarum,30(5):639-649.
Chen J Y,Liu S L,Siao W,Wang S J.2010.Hormone and sugar effects on rice sucrose transporter OsSUT1 expression in germinating embryos.Acta Physiologiae Plantarum,32(4):749-756.
Chincinska I,Gier K,Krugel U,Liesche J,He H,Grimm B.2013.Photoperiodic regulation of the sucrose transporter StSUT4 affects the expression of circadian-regulated genes and ethylene production.Frontiers in Plant Science,4(2):26.
Chincinska I A,Liesche J,Krugel U,Michalska J,Geigenberger P,Grimm B.2008.Sucrose transporter StSUT4 from potato affects flowering,tuberization,and shade avoidance response.Plant Physiology,146(2):515-528.
Chun O K,Kim D,Lee C Y.2003.Superoxide radical scavenging activity of the major polyphenols in fresh plums.Journal of Agricultural and Food Chemistry,51(27):8067-8072.
Cominelli E,Gusmaroli G,Allegra D,Galbiati M,Wade H K,Jenkins G I,Tonelli C.2008.Expression analysis of anthocyanin regulatory genes in response to different light qualities in Arabidopsis thaliana.Journal of Plant Physiology,165(8):886-894.
Das P K,Shin D H,Choi S B,Park Y I.2012.Sugar-hormone cross-talk in anthocyanin biosynthesis.Molecules and Cells.34(6):501-507.
Deng En-xin,Wang Wen-zhong,Wang Zhi-xin.1992.A simple device that injected solution into the plant’s thin blade.Plant Physiology Communications,28(4):296.(in Chinese)邓恩新,王文忠,王智炘.1992.一种简易的向植物薄叶片内注射溶液的装置.植物生理学通讯,28(4):296.
Gyula P,Schafer E,Nagy F.2003.Light perception and signaling in higher plants,Current Opinion Plant Biology,6:446-452.
Hackel A,Schauer N,Carrari F,Fernie A R,Grimm B,Kuhn C.2006.Sucrose transporter LeSUT1 and LeSUT2 inhibition affects tomato fruit development in different ways.The Plant Journal,45(2):180-192.
He J,Giusti M M.2010.Anthocyanins:natural colorants with health-promoting properties.Annual Review of Food Science Technology,1(1):163-187.
Hiratsuka S,Onodera H,Kawai Y,Kubo T,Itoh H,Wada R.2001.ABA and sugar effects on anthocyanin formation in grapeberry cultured in vitro.Scientia Horticulturae,90(1-2):121-130.
Hirose T,Imaizumi N,Scofield G N,Furbank R T,Ohsugi R.1997.cDNA cloning and tissue specific expression of a gene for sucrose transporter from rice(Oryza sativa L.).Plant Cell Physiology,38(12):1389-1396.
Jeong S W,Das P K,Jeoung S C,Song J Y,Lee H K,Kim Y K,Kim W J,Park Y I,Yoo S D,Choi S B.2010.Ethylene suppression of sugar-induced anthocyanin pigmentation in Arabidopsis thaliana.Plant Physiology,154(3):1515-1531.
Lloyd J C,Zakhleniuk O V.2004.Responses of primary and secondary metabolism to sugar accumulation revealed by microarray expression analysis of the Arabidopsis mutant,pho3.Journal of Experimental Botany,55(400):1221-1230.
Loreti E,Povero G,Novi G,Solfanelli C,Alpi A,Perata P.2008.Gibberellins,jasmonate and abscisic acid modulate the sucrose-induced expression of anthocyanin biosynthetic genes in Arabidopsis.New Phytologist,179(4):1004-1016.
Liu Xiao-bin,Liu Na,Li Fu-kuan,Wu Li-zhu,Zhang Jie,Wang Dong-mei.2015.Establishment of TRV-mediated transient gene-silencing system in soybean.Scientia Agricultura Sinica,48(12):2479-2486.(in Chinese)刘晓彬,刘娜,李福宽,吴立柱,张洁,王冬梅.2015.TRV介导的大豆基因瞬时沉默体系的建立.中国农业科学,48(12):2479-2486.
Martens H J,Roberts A G,Oparka K J.2006.Schulz A.Quantification of plasmodesmatal endoplasmic reticulum coupling between sieve elements and companion cells using fluorescence redistribution after photobleaching.Plant Physiology,142(2):471-480.
Mita S,Murano N,Akaike M,Nakamura K.1997.Mutants of Arabidopsis thaliana with pleiotropic effects on the expression of the gene for betaamylase and on the accumulation of anthocyanin that are inducible by sugars.The Plant Journal,11(4):841-851.
Ohto M,Onai K,Furukawa Y,Aoki E,Araki T,Nakamura K.2001.Effects of sugar on vegetative development and floral transition in Arabidopsis.Plant Physiology,127(1):252-261.
Payyavula R S,Tay K H,Tsai C J,Harding S A.2011.The sucrose transporter family in Populus:the importance of a tonoplast PtaSUT4 to biomass and carbon partitioning.The Plant Journal,65(5):757-770.
Rinaldo A R,Cavallini E,Jia Y,Moss S M A,Mcdavid D A J,Hooper L C,Robinson S P,Tornielli G B,Zenoni S,Ford C M,Boss P K,Walker A R.2015.A grapevine anthocyanin acyltransferase,transcriptionally regulated by VvMYBA,can produce most acylated anthocyanins present in grape skins.Plant Physiology,169(3):1897-1916.
Riesmeier J W,Willmitzer L,Frommer W B.1992.Isolation and characterization of a sucrose carrier cDNA from spinach by functional expression in yeast.Embo Journal,11(13):4705-13.
Rosa M,Prado C,Podazza G,Interdonato R,González J A,Hilal M,Prado F E.2009.Soluble sugars:metabolism,sensing and abiotic stress:a complex network in the life of plants.Plant Signaling Behavior,4(5):388-393.
Sauer N.2007.Molecular physiology of higher plant sucrose transporters.FEBS Letters,581(12):2309-2317.
Schmitz J,Heinrichs L,Scossa F,Fernie A R,Oelze M L,Dietz K J.2014.The essential role of sugar metabolism in the acclimation response of Arabidopsis thaliana to high light intensities.Journal of Experimental Botany,65(6):1619-1636.
Scofield G N,Aoki N,Hirose T,Takano M,Jenkins C L,Furbank R T.2007.The role of the sucrose transporter,Os SUT1,in germination and early seedling growth and development of rice plants.Journal of Experimental Botany,58(3):483-495.
Sivitz A B,Reinders A,Ward J M.2008.Arabidopsis sucrose transporter AtSUC1 is important for pollen germination and sucrose-induced anthocyanin accumulation.Plant Physiology,147(1):92-100.
Solfanelli C,Poggi A,Loreti E,Alpi A,Perata P.2006.Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis.Plant Physiology,140(2):637-646.
Stadler R,Sauer N.1996.The Arabidopsis thaliana AtSUC2 gene is specifically expressed in companion cells.Plant Biology,109(4):299-306.
Télef N,Stammitti-Bert L,Mortain-Bertrand A,Maucourt M,Carde J P,Rolin D.2006.Sucrose deficiency delays lycopene accumulation in tomato fruit pericarp discs.Plant Molecular Biology,62(3):453-469.
Teng S,Keurentjes J,Bentsink L,Koornneef M,Smeekens S.2005.Sucrose-specific induction of anthocyanin biosynthesis in Arabidopsis requires the MYB75/PAP1 gene.Plant Physiology,139(4):1840-1852.
Tsuda T.2012.Dietary anthocyanin-rich plants:biochemical basis and recent progress in health benefits studies.Molecular Nutrition&Food Research,56:159-170.
Wade H K,Bibikova T N,Valentine W J,Jenkins G I.2001.Interactions within a network of phytochrome,cryptochrome and UV-B phototransduction pathways regulate chalcone synthase gene expression in Arabidopsis leaf tissue.The Plant Journal,25(6):675-685.
Weise A,Barker L,Kuhn C,Lalonde S,Buschmann H,Frommer W B.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.
Wheeler S,Loveys B,Ford C,Davies C.2009.The relationship between the expression of abscisic acid biosynthesis genes,accumulation of abscisic acid and the promotion of Vitis vinifera L.berry ripening by abscisic acid.Australian Journal of Grape and Wine Research,15(3):195-204.
Xu Haifeng,Qu Changzhi,Liu Jingxuan,Wang Yicheng,Wang Deyun,Zuo Weifang,Jiang Shenghui,Wang Nan,Zhang Zongying,Chen Xuesen.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.
Zafra-Stone S,Yasmin T,Bagchi M,Chatterjee A,Vinson J A,Bagchi D.2007.Berry anthocyanins as novel antioxidants in human health and disease prevention.Molecular Nutrition&Food Research,51(6):675-683.
Zhang Q,Hu W C,Zhu F,Wang L,Yu Q Y,Ming R,Zhang J.2016.Structure,phylogeny,allelic haplotypes and expression of sucrose transporter gene families in Saccharum.BMC Genomics,17(1):88.
Zhang Weitong.2018.The mechanism of Arabidopsis VAC14 mediate pollen development[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)张威通.2018.拟南芥VAC14调控花粉发育的机理研究[博士论文].泰安:山东农业大学.
Aoki N,Hirose T,Scofield G N,Whitfeld P R,Furbank R T.2003.The sucrose transporter gene family in rice.Plant and Cell Physiology,44(3):223-232.
Aoki N,Whitfeld P,Hoeren F,Scofield G,Newell K,Patrick J.2002.Three sucrose transporter genes are expressed in the developing grain of hexaploid wheat.Plant Molecular Biology,50(3):453-462.
An Xiu-hong.2013.Studies on the mechanism by which MdTTG1,MdMYB9 and MdMYB11 regulate anthocyanin biosynthesis in apple[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)安秀红.2013.苹果MdTTG1、MdMYB9与Md MYB11基因调控花青苷合成的作用机制研究[博士论文].泰安:山东农业大学.
Bai Xue-mei,Zhang Li-jun,Wu Xiao-dan,Hu Kai,Ruan Yan-ye.2006.Sucrose transporters in plants.Plant Physiology Communications,42(6):195-202.(in Chinese)白雪梅,张立军,吴晓丹,胡凯,阮燕晔.2006.植物蔗糖转运蛋白.植物生理学通,42(6):195-202.
Baier M,Hemmann G,Holman R,Corke F,Card R,Smith C,Rook F,Bevan M W.2004.Characterization of mutants in Arabidopsis showing increased sugar-specific gene expression,growth,and developmental responses.Plant Physiology,134(1):81-91.
Bassolino L,Zhang Y,Schoonbeek H J,Kiferle C,Perata P,Martin C.2013.Accumulation of anthocyanins in tomato skin extends shelf life.New Phytologist,200(3):650-655.
Baud S,Wuilleme S,Lemoine R.2005.The AtSUC5 sucrose transporter specifically expressed in the endosperm is involved in early seed development in Arabidopsis.Plant Journal,43:824-836.
Butelli E,Titta L,Giorgio M,Mock H P,Matros A,Peterek S,Schijlen E G,Hall R D,Bovy A G,Luo J,Martin C.2008.Enrichment of tomato fruit with health-promoting anthocyanins by expression of select transcription factors.Nature Biotechnology,26(11):1301-1308.
Chen H J,Wang S J.2008.Molecular regulation of sink-source transition in rice leaf sheaths during the heading period.Acta Physiologiae Plantarum,30(5):639-649.
Chen J Y,Liu S L,Siao W,Wang S J.2010.Hormone and sugar effects on rice sucrose transporter OsSUT1 expression in germinating embryos.Acta Physiologiae Plantarum,32(4):749-756.
Chincinska I,Gier K,Krugel U,Liesche J,He H,Grimm B.2013.Photoperiodic regulation of the sucrose transporter StSUT4 affects the expression of circadian-regulated genes and ethylene production.Frontiers in Plant Science,4(2):26.
Chincinska I A,Liesche J,Krugel U,Michalska J,Geigenberger P,Grimm B.2008.Sucrose transporter StSUT4 from potato affects flowering,tuberization,and shade avoidance response.Plant Physiology,146(2):515-528.
Chun O K,Kim D,Lee C Y.2003.Superoxide radical scavenging activity of the major polyphenols in fresh plums.Journal of Agricultural and Food Chemistry,51(27):8067-8072.
Cominelli E,Gusmaroli G,Allegra D,Galbiati M,Wade H K,Jenkins G I,Tonelli C.2008.Expression analysis of anthocyanin regulatory genes in response to different light qualities in Arabidopsis thaliana.Journal of Plant Physiology,165(8):886-894.
Das P K,Shin D H,Choi S B,Park Y I.2012.Sugar-hormone cross-talk in anthocyanin biosynthesis.Molecules and Cells.34(6):501-507.
Deng En-xin,Wang Wen-zhong,Wang Zhi-xin.1992.A simple device that injected solution into the plant’s thin blade.Plant Physiology Communications,28(4):296.(in Chinese)邓恩新,王文忠,王智炘.1992.一种简易的向植物薄叶片内注射溶液的装置.植物生理学通讯,28(4):296.
Gyula P,Schafer E,Nagy F.2003.Light perception and signaling in higher plants,Current Opinion Plant Biology,6:446-452.
Hackel A,Schauer N,Carrari F,Fernie A R,Grimm B,Kuhn C.2006.Sucrose transporter LeSUT1 and LeSUT2 inhibition affects tomato fruit development in different ways.The Plant Journal,45(2):180-192.
He J,Giusti M M.2010.Anthocyanins:natural colorants with health-promoting properties.Annual Review of Food Science Technology,1(1):163-187.
Hiratsuka S,Onodera H,Kawai Y,Kubo T,Itoh H,Wada R.2001.ABA and sugar effects on anthocyanin formation in grapeberry cultured in vitro.Scientia Horticulturae,90(1-2):121-130.
Hirose T,Imaizumi N,Scofield G N,Furbank R T,Ohsugi R.1997.cDNA cloning and tissue specific expression of a gene for sucrose transporter from rice(Oryza sativa L.).Plant Cell Physiology,38(12):1389-1396.
Jeong S W,Das P K,Jeoung S C,Song J Y,Lee H K,Kim Y K,Kim W J,Park Y I,Yoo S D,Choi S B.2010.Ethylene suppression of sugar-induced anthocyanin pigmentation in Arabidopsis thaliana.Plant Physiology,154(3):1515-1531.
Lloyd J C,Zakhleniuk O V.2004.Responses of primary and secondary metabolism to sugar accumulation revealed by microarray expression analysis of the Arabidopsis mutant,pho3.Journal of Experimental Botany,55(400):1221-1230.
Loreti E,Povero G,Novi G,Solfanelli C,Alpi A,Perata P.2008.Gibberellins,jasmonate and abscisic acid modulate the sucrose-induced expression of anthocyanin biosynthetic genes in Arabidopsis.New Phytologist,179(4):1004-1016.
Liu Xiao-bin,Liu Na,Li Fu-kuan,Wu Li-zhu,Zhang Jie,Wang Dong-mei.2015.Establishment of TRV-mediated transient gene-silencing system in soybean.Scientia Agricultura Sinica,48(12):2479-2486.(in Chinese)刘晓彬,刘娜,李福宽,吴立柱,张洁,王冬梅.2015.TRV介导的大豆基因瞬时沉默体系的建立.中国农业科学,48(12):2479-2486.
Martens H J,Roberts A G,Oparka K J.2006.Schulz A.Quantification of plasmodesmatal endoplasmic reticulum coupling between sieve elements and companion cells using fluorescence redistribution after photobleaching.Plant Physiology,142(2):471-480.
Mita S,Murano N,Akaike M,Nakamura K.1997.Mutants of Arabidopsis thaliana with pleiotropic effects on the expression of the gene for betaamylase and on the accumulation of anthocyanin that are inducible by sugars.The Plant Journal,11(4):841-851.
Ohto M,Onai K,Furukawa Y,Aoki E,Araki T,Nakamura K.2001.Effects of sugar on vegetative development and floral transition in Arabidopsis.Plant Physiology,127(1):252-261.
Payyavula R S,Tay K H,Tsai C J,Harding S A.2011.The sucrose transporter family in Populus:the importance of a tonoplast PtaSUT4 to biomass and carbon partitioning.The Plant Journal,65(5):757-770.
Rinaldo A R,Cavallini E,Jia Y,Moss S M A,Mcdavid D A J,Hooper L C,Robinson S P,Tornielli G B,Zenoni S,Ford C M,Boss P K,Walker A R.2015.A grapevine anthocyanin acyltransferase,transcriptionally regulated by VvMYBA,can produce most acylated anthocyanins present in grape skins.Plant Physiology,169(3):1897-1916.
Riesmeier J W,Willmitzer L,Frommer W B.1992.Isolation and characterization of a sucrose carrier cDNA from spinach by functional expression in yeast.Embo Journal,11(13):4705-13.
Rosa M,Prado C,Podazza G,Interdonato R,González J A,Hilal M,Prado F E.2009.Soluble sugars:metabolism,sensing and abiotic stress:a complex network in the life of plants.Plant Signaling Behavior,4(5):388-393.
Sauer N.2007.Molecular physiology of higher plant sucrose transporters.FEBS Letters,581(12):2309-2317.
Schmitz J,Heinrichs L,Scossa F,Fernie A R,Oelze M L,Dietz K J.2014.The essential role of sugar metabolism in the acclimation response of Arabidopsis thaliana to high light intensities.Journal of Experimental Botany,65(6):1619-1636.
Scofield G N,Aoki N,Hirose T,Takano M,Jenkins C L,Furbank R T.2007.The role of the sucrose transporter,Os SUT1,in germination and early seedling growth and development of rice plants.Journal of Experimental Botany,58(3):483-495.
Sivitz A B,Reinders A,Ward J M.2008.Arabidopsis sucrose transporter AtSUC1 is important for pollen germination and sucrose-induced anthocyanin accumulation.Plant Physiology,147(1):92-100.
Solfanelli C,Poggi A,Loreti E,Alpi A,Perata P.2006.Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis.Plant Physiology,140(2):637-646.
Stadler R,Sauer N.1996.The Arabidopsis thaliana AtSUC2 gene is specifically expressed in companion cells.Plant Biology,109(4):299-306.
Télef N,Stammitti-Bert L,Mortain-Bertrand A,Maucourt M,Carde J P,Rolin D.2006.Sucrose deficiency delays lycopene accumulation in tomato fruit pericarp discs.Plant Molecular Biology,62(3):453-469.
Teng S,Keurentjes J,Bentsink L,Koornneef M,Smeekens S.2005.Sucrose-specific induction of anthocyanin biosynthesis in Arabidopsis requires the MYB75/PAP1 gene.Plant Physiology,139(4):1840-1852.
Tsuda T.2012.Dietary anthocyanin-rich plants:biochemical basis and recent progress in health benefits studies.Molecular Nutrition&Food Research,56:159-170.
Wade H K,Bibikova T N,Valentine W J,Jenkins G I.2001.Interactions within a network of phytochrome,cryptochrome and UV-B phototransduction pathways regulate chalcone synthase gene expression in Arabidopsis leaf tissue.The Plant Journal,25(6):675-685.
Weise A,Barker L,Kuhn C,Lalonde S,Buschmann H,Frommer W B.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.
Wheeler S,Loveys B,Ford C,Davies C.2009.The relationship between the expression of abscisic acid biosynthesis genes,accumulation of abscisic acid and the promotion of Vitis vinifera L.berry ripening by abscisic acid.Australian Journal of Grape and Wine Research,15(3):195-204.
Xu Haifeng,Qu Changzhi,Liu Jingxuan,Wang Yicheng,Wang Deyun,Zuo Weifang,Jiang Shenghui,Wang Nan,Zhang Zongying,Chen Xuesen.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.
Zafra-Stone S,Yasmin T,Bagchi M,Chatterjee A,Vinson J A,Bagchi D.2007.Berry anthocyanins as novel antioxidants in human health and disease prevention.Molecular Nutrition&Food Research,51(6):675-683.
Zhang Q,Hu W C,Zhu F,Wang L,Yu Q Y,Ming R,Zhang J.2016.Structure,phylogeny,allelic haplotypes and expression of sucrose transporter gene families in Saccharum.BMC Genomics,17(1):88.
Zhang Weitong.2018.The mechanism of Arabidopsis VAC14 mediate pollen development[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)张威通.2018.拟南芥VAC14调控花粉发育的机理研究[博士论文].泰安:山东农业大学.