马铃薯淀粉合成与降解研究进展
|
摘要
马铃薯是我国第四大粮食作物,其总产量占全国粮食产量的20%。马铃薯是C3作物,其淀粉含量相对较低,这与其淀粉合成和降解过程有关。与马铃薯淀粉合成的相关酶包括腺苷二磷酸葡萄糖焦磷酸化酶(ADP-glucose pyrophosphorylase,AGPase)、颗粒结合性淀粉合成酶(Granule-bound starch synthase,GBSS)、可溶性淀粉合成酶(Soluble starch synthase,SSS)、淀粉分支酶(Starch branching enzyme,SBE)和淀粉去分支酶(Starch debranching enzyme,DBE)。α-淀粉酶(α-amylase,Amy)、β-淀粉酶(β-amylase,BAM)、葡聚糖水双激酶(Glucan water dikinase,GWD)、磷酸葡聚糖水双激酶(Phosphoglucan water dikinase,PWD)和酸性转化酶(Acid invertase,AI)、中/碱性转化酶(Neutral/alkaline inverstase,NI)则与马铃薯淀粉降解有关。马铃薯淀粉合成与降解基因克隆对马铃薯品种遗传改良具有重要意义,目前部分克隆的马铃薯淀粉合成与降解基因已在增加马铃薯淀粉含量、淀粉改性和改变淀粉组分方面得到广泛应用。总结了马铃薯淀粉合成与降解酶基因的研究进展,并提出今后研究建议,旨为深入开展马铃薯淀粉改良工程研究提供参考。
Potato is the fourth largest food crop in China and its total output accounts for 20% of the grain yield in the country. Potato is a C3 crop and its starch content is relatively low,which is related to its starch synthesis and degradation process. Enzymes related to potato starch synthesis include ADP-glucose pyrophosphorylase,granule-bound starch synthase,soluble starch synthase,starch branching enzyme,and starch debranching enzyme,while α-amylase,β-amylase,glucan water dikinase,phosphoglucan water dikinase,acid invertase,and neutral/alkaline inverstase are associated with the degradation of potato starch. Cloning potato starch synthesis and degradation genes is of great significance for genetic improvement of potato varieties. At present some cloned genes have been widely used in increasing starch content,modifying starch,and changing starch components in potato. This paper summarized the research progress on potato starch synthesis and degrading enzyme genes. Some suggestions have been put forward for future research,aiming at providing reference for future research on potato starch improvement engineering.
Potato is the fourth largest food crop in China and its total output accounts for 20% of the grain yield in the country. Potato is a C3 crop and its starch content is relatively low,which is related to its starch synthesis and degradation process. Enzymes related to potato starch synthesis include ADP-glucose pyrophosphorylase,granule-bound starch synthase,soluble starch synthase,starch branching enzyme,and starch debranching enzyme,while α-amylase,β-amylase,glucan water dikinase,phosphoglucan water dikinase,acid invertase,and neutral/alkaline inverstase are associated with the degradation of potato starch. Cloning potato starch synthesis and degradation genes is of great significance for genetic improvement of potato varieties. At present some cloned genes have been widely used in increasing starch content,modifying starch,and changing starch components in potato. This paper summarized the research progress on potato starch synthesis and degrading enzyme genes. Some suggestions have been put forward for future research,aiming at providing reference for future research on potato starch improvement engineering.
引文
[1]Fernie AR,Willmitzer L,Trethewey RN.Sucrose to starch:a transition in molecular plant physiology[J].Trends Plant Sci,2002,7(1):35-41.
[2]Sweetlove LJ,Müller-R?eber B,Willmitzer L,Hill SA.The contribution of adenosine 5'-diphosphoglucose pyrophosphorylase to the control of starch synthesis in potato tubers[J].Planta,1999,209:330-337.
[3]Stitt M.Progress in understanding and engineering primary plant metabolism[J].Curr Opin Biotechnol,2013,24(2):229-238.
[4]白桦,崔雪琼,白少星,姚新灵.马铃薯AGPase活力反馈调控光合速率定量分析[J].生物技术通报,2014(11):125-129.
[5]成善汉,苏振洪,谢从华,柳俊.淀粉-糖代谢酶活性变化对马铃薯块茎还原糖积累及加工品质的影响[J].中国农业科学,2004,37(12):1904-1910.
[6]陈国梁,张金文,王蒂.马铃薯gbss、ssò和ssó基因片段的融合及其RNAi载体的构建[J].中国生物工程杂志,2008,28(8):51-56.
[7]戴卫列,邓炜,崔伟英,等.马铃薯GBSS基因的克隆与DNA顺序分析[J].植物学报,1996,38(10):777-784.
[8]李淑洁,张金文,王煜,等.一个新的马铃薯GBSS基因5'侧翼序列克隆及调控活性研究[J].中国马铃薯,2005,19(3):129-133.
[9]宋东光,孙国枫,单海燕,等.马铃薯GBSS基因5'侧翼区调控作用的研究[J].植物学报,1998,40(9):796-802.
[10]Edwards A,Fulton DC,Hylton CM,et al.A combined reduction in activity of starch synthases II and III of potato has novel effects on the starch of tubers[J].Plant J,1999,17(3):251-261.
[11]杨涛,张宁,栗亮,等.马铃薯可溶性淀粉合成酶SSIII基因克隆及生物信息学分析[J].分子植物育种,2009,7(3):545-549.
[12]Kristensen M,Lok F,Planchot V,et al.Isolation and characterisation of the gene encoding the starch debranching enzyme limit dextrinase from germinating barley[J].Biochim Biophys Acta,1999,1431:538-546.
[13]Kim KN,Fisher DK,Gao M,Guiltinan MJ.Genome organization and promoter activity of the maize starch branching enzyme Igene[J].Gene,1998,216:233-243.
[14]Bae JM,Liu JR.Molecular cloning and characterization of two novel iosoforms of the small subunit of ADPglucose pyrophosphorylase from sweet potato[J].Mol Gen Genet,1997(2):179-185.
[15]Harn CH,Bae JM,Lee SS,et al.Presence of multiple cDNAs encoding an isoform of ADP-glucose pyrophosphorylase large subunit from sweet potato and characterization of expression levels[J].Plant Cell Physiol,2000,41(11):1235-1242.
[16]Dry I,Smith A,Edward A,et al.Characterization of cDNAs encoding two isoforms of granule-bound starch synthase which show differential expression in developing storage organs of pea and potato[J].Plant J,1992,2(2):193-202.
[17]Kossmann J,Abel GJW,Springer F,Lloyd JR,Willmitzer L.Cloning and functional analysis of a cDNA encoding a starch synthase from potato(Solanum tuberosum L.)that is predominantly expressed in leaf tissue[J].Planta,1999,208:503-511.
[18]Larsson CT,Khoshnoodi J,Ek B,et al.Molecular cloning and characterization of starch-branching enzyme II from potato[J].Plant Mol Biol,1998,37(3):505-511.
[19]Abel GJW,Springer F,Willmitzer L,Kossmann J.Cloning and fuctional analysis of a cDNA encoding a novel 139 kDa starch synthase from potato(Solanum tuberosum L.)[J].Plant J,1996,10(6):981-991.
[20]Marshall J,Sidebottom C,Debet M,et al.Identification of the major starch synthase in the soluble fraction of potato tubers[J].Plant Cell,1996,8(7):1121-1135.
[21]Yang T,Zhang N,Li L,et al.Cloning and bioinformatics analysis of soluble starch synthase SSIII gene in potato[J].Mol Plant Breed,2009,7:545-549.
[22]Jobling SA,Westcott RJ,Tayal A,Jeffcoat R,Schwall GP.Production of a freeze-thaw-stable potato starch by antisense inhibition of three starch synthase genes[J].Nature Biotechnol,2002,20(3):295-299.
[23]Lee SS,Bae JM,Oh MS,et al.Isolation and characterization of polymorphic cDNAs parially encoding ADP-glucose pyrophosphorylase(AGPase)large subunit from sweet potato[J].Mol Cells,2000,10(1):108-112.
[24]Silver DM,Kotting O,Moorhead GBG.Phosphoglucan phosphatase function sheds light on starch degradation[J].Trends Plant Sci,2014,19(7):471-478.
[25]Preiss J.Regulation of the biosynthesis and degradation of starch[J].Annu Rev Plant Physiol,1982,33:431-454.
[26]Streb S,Zeeman SC.Starch metabolism in Arabidopis[J].The Arabidopsis Book,2012,e0160.
[27]Monroe JD,Storm AR,Badley EM,et al.β-amylase 1 andβ-amylase3 are plastidic starch hydrolases in Arabidopsis that seem to be adapted for different thermal,pH,and stress conditions[J].Plant Physiol,2014,166:1748-1763.
[28]Bagnaresi P,Moschella A,Beretta O,et al.Heterologous microarray experiments allow the identification of the early events associated with potato tuber cold sweetening[J].BMC Genomics,2008,9:176-198.
[29]Zhang H,Hou J,Liu J,et al.Amylase anlaysis in potato starch degradation during cold storage and sprouting[J].Potato Res,2014,57:47-58.
[30]Ritte G,Heydenreich M,Mahlow S,et al.Phosphorylation of C6-and C3-position of glycosyl residues in starch is catalysed by distinct dikinases[J].FEBS Lett,2006,580:4872-4876.
[31]Liu X,Zhang C,Ou YB,et al.Systematic analysis of potato acid invertase genes reveals that a cold-responsive member,StvacINV1,regulates cold-induced sweetening of tubers[J].Mol Genet Genomics,2011,286:109-118.
[32]Sweetlove LJ,Burrell MM,Rees T.Characterization of transgenic potato(Solanum tuberosum)tubers with increased ADP-Glucose pyrophosphorylase[J].Biochemistry J,1996,320:478-492.
[33]Tjaden J,Mohlmann T,Kampfenkel K.Altered plastidic ATP/ADP-transporter activity influence potato(Solanum tuberosum L.)tuber morphology,yield and composition of tuber starch[J].Plant J,1998,16:531-540.
[34]宋波涛,谢从华,柳俊.马铃薯s AGP基因表达对块茎淀粉和还原糖含量的影响[J].中国农业科学,2005,38(7):1439-1446.
[35]刘廷国,李斌,谢笔钧.转AGPase基因马铃薯淀粉溶液行为及热特性比较研究[J].作物学报,2006,32(2):310-312.
[36]Lorbeth R,Ritte G,Willmitzer L,et al.Inhibition of a starch-granule-bound protein leads to modified starch and repression of cold sweetening[J].Nat Biotechnol,1998,16:473-477.
[37]Zhang H,Liu J,Hou J,et al.The potato amylase inhibitor gene SbAIregulates cold-induced sweetening in potato tubers by modulating amylase activity[J].Plant Biotechnol J,2014,12:984-993.
[38]崔雪琼.重组glgC基因调控块茎淀粉生物合成研究[M].银川:宁夏大学,2012.
[39]Vardy KA,Emes MJ,Burrell MM.Starch synthesis in potato tubers transformed with the wheat genes for ADPglucose pyrophosphorylase[J].Functional Plant Biol,2002,29(8):975-985.
[40]Muller-Rober B,Sonnewald U,Willmitzer L.Inhibition of AGPase in transgenic potatoes leads to sugar-storing tubers and influences tuber formation and expression of tuber-storage protein genes[J].EMBO J,1992,11:1229-1238.
[41]Scheidig A,Frohlich A,Schulze S,et al.Downregulation of a chloroplast-targetedβ-amylase leads to a starch-excess phenotype in leaves[J].Plant J,2002,30:581-591.
[42]Bustos R,Fahy B,Hylton CM,et al.Starch granule initiation is controlled by a heteromultimeric isoamylase in potato tubers[J].Proc Natl Acad Sci USA,2004,101:2215-2220.
[43]Ferreira SJ.Transcriptome based analysis of starch metabolism in Solanum tuberosum[D].Friedrich-Alexander-Universitat Erlangen-Nurnberg,2011.
[44]Holen T,Amarzguioui M,Wiiger MT,et al.Positional effects of short interfering RNAs targeting the human coagulation trigger tissue factor[J].Nucleic Acids Res,2002,30(8):1757-1766.
[45]Heilersig HJB,Loonen A,Bergervoet M,et al.Post-transcriptional gene silencing of GBSSI in potato:effects of size and sequence of the inverted repeats[J].Plant Mol Biol,2006,60:647-662.
[46]Baba T.Identification,cDNA cloning,and gene expression of soluable starch synthase in rice(Oryza sativa L.)immature seeds[J].Plant Physiol,1993,103:565-573.
[47]杜宏辉.可溶性淀粉合成酶SSIII基因对马铃薯的遗传转化[D].兰州:甘肃农业大学,2011:7-10.
[48]Abel GJW,Springer F,Willmitzer L,et al.Cloning and functional analysis of a cDNA encoding a novel 139 kDa starch synthase from potato(Solanum tuberosum L.)[J].Plant J,1996,10(6):981-991.
[49]Blauth SL,Yao Y,Klucinec JD,et al.Identification of mutator insertional mutants of starch-branching enzymes in corn[J].Plant Physiol,2001,125(3):1396-1405.
[50]Wolters AMA,Janssen EM,Rozeboom-Schippers MGM,et al.Composition of endogenous alleles can influence the level of antisense inhibition of granule-bound starch synthase gene expression in tetraploid potato plants[J].Mol Breeding,1998,4(4):343-358.
[51]蔺琰东.马铃薯GBSSI基因的ihpRNAi载体转化试管薯片及转基因块茎相关生理分析[D].兰州:甘肃农业大学,2011.
[52]杜宏辉,文义凯,张宁,等.可溶性淀粉合成酶SSIII基因对马铃薯的遗传转化[J].基因组学与应用生物学,2011,30(3):303-307.
[53]Andersson M,Melander M,Pojmark P,et al.Targeted gene suppression by RNA interference:An efficient method for production of highamylose potato lines[J].J Biotechnol 2006,123(2):137-148.
[54]Xu X,Dees D,Dechesne A,et al.Starch phosphorylation plays an important role in starch biosynthesis[J].Carbohyd Polym,2017,157:1628-1637.
[55]Zhang H,Hou J,Liu J,et al.The roles of starch metabolic pathways in the cold-induced sweetening process in potatoes[J].StarchStarke,2016,doi:10.1002/star.201600194.
[56]Liu XY,Rocha-Sosa M,Hummel S,et al.A detailed study of the regulation and evolution of the two classes of patatin genes in Solanum tuberosum L.[J].Plant Mol Biol,1991,17(6):1139-1154.
[57]Clasen BM,Stoddard TJ,Luo S,et al.Improving cold storage and processing traits in potato through targeted gene knockout[J].Plant Biotechnol J,2016,14:169-176.
[58]Carpenter MA,Joyce NI,Genet RA,et al.Starch phosphorylation in potato tubers is influenced by allelic variation in the genes encoding glucan water dikinase,starch branching enzymes I and II,and starch synthase III[J].Front Plant Sci,2015,6:143.
[2]Sweetlove LJ,Müller-R?eber B,Willmitzer L,Hill SA.The contribution of adenosine 5'-diphosphoglucose pyrophosphorylase to the control of starch synthesis in potato tubers[J].Planta,1999,209:330-337.
[3]Stitt M.Progress in understanding and engineering primary plant metabolism[J].Curr Opin Biotechnol,2013,24(2):229-238.
[4]白桦,崔雪琼,白少星,姚新灵.马铃薯AGPase活力反馈调控光合速率定量分析[J].生物技术通报,2014(11):125-129.
[5]成善汉,苏振洪,谢从华,柳俊.淀粉-糖代谢酶活性变化对马铃薯块茎还原糖积累及加工品质的影响[J].中国农业科学,2004,37(12):1904-1910.
[6]陈国梁,张金文,王蒂.马铃薯gbss、ssò和ssó基因片段的融合及其RNAi载体的构建[J].中国生物工程杂志,2008,28(8):51-56.
[7]戴卫列,邓炜,崔伟英,等.马铃薯GBSS基因的克隆与DNA顺序分析[J].植物学报,1996,38(10):777-784.
[8]李淑洁,张金文,王煜,等.一个新的马铃薯GBSS基因5'侧翼序列克隆及调控活性研究[J].中国马铃薯,2005,19(3):129-133.
[9]宋东光,孙国枫,单海燕,等.马铃薯GBSS基因5'侧翼区调控作用的研究[J].植物学报,1998,40(9):796-802.
[10]Edwards A,Fulton DC,Hylton CM,et al.A combined reduction in activity of starch synthases II and III of potato has novel effects on the starch of tubers[J].Plant J,1999,17(3):251-261.
[11]杨涛,张宁,栗亮,等.马铃薯可溶性淀粉合成酶SSIII基因克隆及生物信息学分析[J].分子植物育种,2009,7(3):545-549.
[12]Kristensen M,Lok F,Planchot V,et al.Isolation and characterisation of the gene encoding the starch debranching enzyme limit dextrinase from germinating barley[J].Biochim Biophys Acta,1999,1431:538-546.
[13]Kim KN,Fisher DK,Gao M,Guiltinan MJ.Genome organization and promoter activity of the maize starch branching enzyme Igene[J].Gene,1998,216:233-243.
[14]Bae JM,Liu JR.Molecular cloning and characterization of two novel iosoforms of the small subunit of ADPglucose pyrophosphorylase from sweet potato[J].Mol Gen Genet,1997(2):179-185.
[15]Harn CH,Bae JM,Lee SS,et al.Presence of multiple cDNAs encoding an isoform of ADP-glucose pyrophosphorylase large subunit from sweet potato and characterization of expression levels[J].Plant Cell Physiol,2000,41(11):1235-1242.
[16]Dry I,Smith A,Edward A,et al.Characterization of cDNAs encoding two isoforms of granule-bound starch synthase which show differential expression in developing storage organs of pea and potato[J].Plant J,1992,2(2):193-202.
[17]Kossmann J,Abel GJW,Springer F,Lloyd JR,Willmitzer L.Cloning and functional analysis of a cDNA encoding a starch synthase from potato(Solanum tuberosum L.)that is predominantly expressed in leaf tissue[J].Planta,1999,208:503-511.
[18]Larsson CT,Khoshnoodi J,Ek B,et al.Molecular cloning and characterization of starch-branching enzyme II from potato[J].Plant Mol Biol,1998,37(3):505-511.
[19]Abel GJW,Springer F,Willmitzer L,Kossmann J.Cloning and fuctional analysis of a cDNA encoding a novel 139 kDa starch synthase from potato(Solanum tuberosum L.)[J].Plant J,1996,10(6):981-991.
[20]Marshall J,Sidebottom C,Debet M,et al.Identification of the major starch synthase in the soluble fraction of potato tubers[J].Plant Cell,1996,8(7):1121-1135.
[21]Yang T,Zhang N,Li L,et al.Cloning and bioinformatics analysis of soluble starch synthase SSIII gene in potato[J].Mol Plant Breed,2009,7:545-549.
[22]Jobling SA,Westcott RJ,Tayal A,Jeffcoat R,Schwall GP.Production of a freeze-thaw-stable potato starch by antisense inhibition of three starch synthase genes[J].Nature Biotechnol,2002,20(3):295-299.
[23]Lee SS,Bae JM,Oh MS,et al.Isolation and characterization of polymorphic cDNAs parially encoding ADP-glucose pyrophosphorylase(AGPase)large subunit from sweet potato[J].Mol Cells,2000,10(1):108-112.
[24]Silver DM,Kotting O,Moorhead GBG.Phosphoglucan phosphatase function sheds light on starch degradation[J].Trends Plant Sci,2014,19(7):471-478.
[25]Preiss J.Regulation of the biosynthesis and degradation of starch[J].Annu Rev Plant Physiol,1982,33:431-454.
[26]Streb S,Zeeman SC.Starch metabolism in Arabidopis[J].The Arabidopsis Book,2012,e0160.
[27]Monroe JD,Storm AR,Badley EM,et al.β-amylase 1 andβ-amylase3 are plastidic starch hydrolases in Arabidopsis that seem to be adapted for different thermal,pH,and stress conditions[J].Plant Physiol,2014,166:1748-1763.
[28]Bagnaresi P,Moschella A,Beretta O,et al.Heterologous microarray experiments allow the identification of the early events associated with potato tuber cold sweetening[J].BMC Genomics,2008,9:176-198.
[29]Zhang H,Hou J,Liu J,et al.Amylase anlaysis in potato starch degradation during cold storage and sprouting[J].Potato Res,2014,57:47-58.
[30]Ritte G,Heydenreich M,Mahlow S,et al.Phosphorylation of C6-and C3-position of glycosyl residues in starch is catalysed by distinct dikinases[J].FEBS Lett,2006,580:4872-4876.
[31]Liu X,Zhang C,Ou YB,et al.Systematic analysis of potato acid invertase genes reveals that a cold-responsive member,StvacINV1,regulates cold-induced sweetening of tubers[J].Mol Genet Genomics,2011,286:109-118.
[32]Sweetlove LJ,Burrell MM,Rees T.Characterization of transgenic potato(Solanum tuberosum)tubers with increased ADP-Glucose pyrophosphorylase[J].Biochemistry J,1996,320:478-492.
[33]Tjaden J,Mohlmann T,Kampfenkel K.Altered plastidic ATP/ADP-transporter activity influence potato(Solanum tuberosum L.)tuber morphology,yield and composition of tuber starch[J].Plant J,1998,16:531-540.
[34]宋波涛,谢从华,柳俊.马铃薯s AGP基因表达对块茎淀粉和还原糖含量的影响[J].中国农业科学,2005,38(7):1439-1446.
[35]刘廷国,李斌,谢笔钧.转AGPase基因马铃薯淀粉溶液行为及热特性比较研究[J].作物学报,2006,32(2):310-312.
[36]Lorbeth R,Ritte G,Willmitzer L,et al.Inhibition of a starch-granule-bound protein leads to modified starch and repression of cold sweetening[J].Nat Biotechnol,1998,16:473-477.
[37]Zhang H,Liu J,Hou J,et al.The potato amylase inhibitor gene SbAIregulates cold-induced sweetening in potato tubers by modulating amylase activity[J].Plant Biotechnol J,2014,12:984-993.
[38]崔雪琼.重组glgC基因调控块茎淀粉生物合成研究[M].银川:宁夏大学,2012.
[39]Vardy KA,Emes MJ,Burrell MM.Starch synthesis in potato tubers transformed with the wheat genes for ADPglucose pyrophosphorylase[J].Functional Plant Biol,2002,29(8):975-985.
[40]Muller-Rober B,Sonnewald U,Willmitzer L.Inhibition of AGPase in transgenic potatoes leads to sugar-storing tubers and influences tuber formation and expression of tuber-storage protein genes[J].EMBO J,1992,11:1229-1238.
[41]Scheidig A,Frohlich A,Schulze S,et al.Downregulation of a chloroplast-targetedβ-amylase leads to a starch-excess phenotype in leaves[J].Plant J,2002,30:581-591.
[42]Bustos R,Fahy B,Hylton CM,et al.Starch granule initiation is controlled by a heteromultimeric isoamylase in potato tubers[J].Proc Natl Acad Sci USA,2004,101:2215-2220.
[43]Ferreira SJ.Transcriptome based analysis of starch metabolism in Solanum tuberosum[D].Friedrich-Alexander-Universitat Erlangen-Nurnberg,2011.
[44]Holen T,Amarzguioui M,Wiiger MT,et al.Positional effects of short interfering RNAs targeting the human coagulation trigger tissue factor[J].Nucleic Acids Res,2002,30(8):1757-1766.
[45]Heilersig HJB,Loonen A,Bergervoet M,et al.Post-transcriptional gene silencing of GBSSI in potato:effects of size and sequence of the inverted repeats[J].Plant Mol Biol,2006,60:647-662.
[46]Baba T.Identification,cDNA cloning,and gene expression of soluable starch synthase in rice(Oryza sativa L.)immature seeds[J].Plant Physiol,1993,103:565-573.
[47]杜宏辉.可溶性淀粉合成酶SSIII基因对马铃薯的遗传转化[D].兰州:甘肃农业大学,2011:7-10.
[48]Abel GJW,Springer F,Willmitzer L,et al.Cloning and functional analysis of a cDNA encoding a novel 139 kDa starch synthase from potato(Solanum tuberosum L.)[J].Plant J,1996,10(6):981-991.
[49]Blauth SL,Yao Y,Klucinec JD,et al.Identification of mutator insertional mutants of starch-branching enzymes in corn[J].Plant Physiol,2001,125(3):1396-1405.
[50]Wolters AMA,Janssen EM,Rozeboom-Schippers MGM,et al.Composition of endogenous alleles can influence the level of antisense inhibition of granule-bound starch synthase gene expression in tetraploid potato plants[J].Mol Breeding,1998,4(4):343-358.
[51]蔺琰东.马铃薯GBSSI基因的ihpRNAi载体转化试管薯片及转基因块茎相关生理分析[D].兰州:甘肃农业大学,2011.
[52]杜宏辉,文义凯,张宁,等.可溶性淀粉合成酶SSIII基因对马铃薯的遗传转化[J].基因组学与应用生物学,2011,30(3):303-307.
[53]Andersson M,Melander M,Pojmark P,et al.Targeted gene suppression by RNA interference:An efficient method for production of highamylose potato lines[J].J Biotechnol 2006,123(2):137-148.
[54]Xu X,Dees D,Dechesne A,et al.Starch phosphorylation plays an important role in starch biosynthesis[J].Carbohyd Polym,2017,157:1628-1637.
[55]Zhang H,Hou J,Liu J,et al.The roles of starch metabolic pathways in the cold-induced sweetening process in potatoes[J].StarchStarke,2016,doi:10.1002/star.201600194.
[56]Liu XY,Rocha-Sosa M,Hummel S,et al.A detailed study of the regulation and evolution of the two classes of patatin genes in Solanum tuberosum L.[J].Plant Mol Biol,1991,17(6):1139-1154.
[57]Clasen BM,Stoddard TJ,Luo S,et al.Improving cold storage and processing traits in potato through targeted gene knockout[J].Plant Biotechnol J,2016,14:169-176.
[58]Carpenter MA,Joyce NI,Genet RA,et al.Starch phosphorylation in potato tubers is influenced by allelic variation in the genes encoding glucan water dikinase,starch branching enzymes I and II,and starch synthase III[J].Front Plant Sci,2015,6:143.