葡聚糖水合二激酶的生物学功能及其应用
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摘要
葡聚糖水合二激酶(glucan-water dikinase,GWD)催化淀粉发生的磷酸化反应是植物临时淀粉降解的必要过程.该文总结了GWD的结构、生物学功能及其与其他淀粉代谢相关酶类的互作,以及GWD基因的研究进展及其在淀粉改性工艺上的应用现状,展望了GWD在淀粉磷酸化改性工艺上的应用前景,旨在为淀粉生物改性提供一种环境友好型、资源集约型的思路和方法.
Starch phosphorylation is a general phenomenon existing naturally in the plant kingdom.The reaction of phosphorylation catalyzed by the glucan-water dikinase(GWD)is a necessary process during the degradation of the transitory starch in plant.In this review,we summarized the progress made on GWD from the view of its structure,biological functions and its interactions with other enzymes involved in starch metabolism.We also discussed its potential application in starch processing industries to provide environment-friendly methods for starch modification through genetic engineering.Potato GWD gene(StGWD)encodes a protein of 1461 amino acid residues with a calculated molecular mass of about 160 ku for the entire coding sequence.From the N-terminal to C-terminal of the mature StGWD protein,there is a chloroplast transit peptide domain,two carbohydratebinding modules,aphosphohistidine domain and nucleotide binding domain in turn.GWD marks sections for glucan hydrolysis via C6 phosphorylation to initiate granule surface hydration,while phosphoglucan water dikinase(PWD),an isoform of GWD then recognizes these partially solubilized sections and catalyzes C3 phosphorylation of nascent glucans to induce steric strain that breaks the helical structure and prevents recrystallization.The starch in leaves will encounter an excess accumulation if the expression of the GWD is inhibited or decreased.And the overexpression of the enzyme may increase the phosphate content of the natural starch.In vivo,many enzymes related to starch metabolism can interact with the GWD,but the mechanisms of their real interaction are not clear yet.The growth will be profoundly inhibited if plants are encountered with environmental stress.We hypothesizethat higher phosphorylated starch may be accumulated if farmers can make good use of the adversary conditions.However,further research and experimental data are needed to support this hypothesis.
Starch phosphorylation is a general phenomenon existing naturally in the plant kingdom.The reaction of phosphorylation catalyzed by the glucan-water dikinase(GWD)is a necessary process during the degradation of the transitory starch in plant.In this review,we summarized the progress made on GWD from the view of its structure,biological functions and its interactions with other enzymes involved in starch metabolism.We also discussed its potential application in starch processing industries to provide environment-friendly methods for starch modification through genetic engineering.Potato GWD gene(StGWD)encodes a protein of 1461 amino acid residues with a calculated molecular mass of about 160 ku for the entire coding sequence.From the N-terminal to C-terminal of the mature StGWD protein,there is a chloroplast transit peptide domain,two carbohydratebinding modules,aphosphohistidine domain and nucleotide binding domain in turn.GWD marks sections for glucan hydrolysis via C6 phosphorylation to initiate granule surface hydration,while phosphoglucan water dikinase(PWD),an isoform of GWD then recognizes these partially solubilized sections and catalyzes C3 phosphorylation of nascent glucans to induce steric strain that breaks the helical structure and prevents recrystallization.The starch in leaves will encounter an excess accumulation if the expression of the GWD is inhibited or decreased.And the overexpression of the enzyme may increase the phosphate content of the natural starch.In vivo,many enzymes related to starch metabolism can interact with the GWD,but the mechanisms of their real interaction are not clear yet.The growth will be profoundly inhibited if plants are encountered with environmental stress.We hypothesizethat higher phosphorylated starch may be accumulated if farmers can make good use of the adversary conditions.However,further research and experimental data are needed to support this hypothesis.
引文
[1]Singh N,Singh J,Kaur L,et al.Morphological,thermal and rheological properties of starches from different botanical sources.Food Chemistry,2003,81(2):219-231.
[2]Fernbach A.Quelques Observations sur la Composition de l'amidon de Pommes de terre.Deslis Frères,1904,138:428-430.
[3]Lorberth R,Ritte G,Willmitzer L,et al.Inhibition of a starch-granule-bound protein leads to modified starch and repression of cold sweetening.Nature Biotechnology,1998,16(5):473-477.
[4]Ritte G,Lloyd J R,Eckermann N,et al.The starch-related R1protein is anα-glucan,water dikinase.Proceedings of the National Academy of Sciences of the United States of America,2002,99(10):7166-7171.
[5]Singh J,Kaur L,McCarthy O J.Factors influencing the physicochemical,morphological,thermal and rheological properties of some chemically modified starches for food applications:a review.Food Hydrocolloids,2007,21(1):1-22.
[6]Wei M,Liu Y,Liu B X,et al.Preparation and application of starch phosphate with a low degree of substitution.Phosphorus,Sulfur,and Silicon and the Related Elements,2011,186(4):974-982.
[7]Whistler R J,Bemiller J N,Pascal E F.Starch:Chemistry and Technology.Beijing:Chinese Food Press,1987:350-354.
[8]Blennow A,Jensen S L,Shaik S S,et al.Future cereal starch bioengineering:cereal ancestors encounter gene technology and designer enzymes.Cereal Chemistry,2013,90(4):274-287.
[9]Schewe G,Knies P,Amati S F,et al.Monocotyledon plant cells and plants which synthesise modified starch.US,6734340.2004-05-11[2014-04-30].
[10]Basu S,Lanahan M.Modified starch,uses,methods for production thereof.US,10/556276,2004-05-24[2014-04-30].
[11]Carciofi M,Shaik S S,Jensen S L,et al.Hyperphosphorylation of cereal starch.Journal of Cereal Science,2011,54(3):339-346.
[12]Frohberg C.Genetically modified plants which synthesize a starch having increased swelling power.US,8304606,2012-11-06[2014-04-30].
[13]Blennow A,Engelsen S B,Munck,L,et al.Starch molecular structure and phosphorylation investigated by a combined chromatographic and chemometric approach.Carbohydrate Polymers,2000,41(2):163-174.
[14]Blennow A,Nielsen T H,Baunsgaard L,et al.Starch phosphorylation:a new front line in starch research.Trends in Plant Science,2002,7(10):445-450.
[15]Zeeman S C,Kossmann J,Smith A M.Starch:its metabolism,evolution,and biotechnological modification in plants.Annual Review of Plant Biology,2010,61:209-234.
[16]Bay-Smidt A M,Wischmann B,Olsen C E,et al.Starch bound phosphate in potato as studied by a simple method for determination of organic phosphate and(31)P-NMR.Strke,1994,46(5):167-172.
[17]Blennow A,Bay-Smidt A M,Wischmann B,et al.The degree of starch phosphorylation is related to the chain length distribution of the neutral and the phosphorylated chains of amylopectin.Carbohydrate Research,1998,307(1):45-54.
[18]Hizukuri S,Takeda Y,Yasuda M,et al.Multi-branched nature of amylose and the action of debranching enzymes.Carbohydrate Research,1981,94(2):205-213.
[19]Mahlow S,Hejazi M,Kuhnert F,et al.Phosphorylation of transitory starch byα-glucan,water dikinase during starch turnover affects the surface properties and morphology of starch granules.New Phytologist,2014,203(2):495-507.
[20]Edner C,Li J,Albrecht T,et al.Glucan,water dikinase activity stimulates breakdown of starch granules by plastidialβ-amylases.Plant Physiology,2007,145(1):17-28.
[21]毛海锋.碎米淀粉及其磷酸酯的制备和理化性质研究.长沙:湖南农业大学,2009:17-18.Mao H F.Study on preparation and physicochemical properties of broken rice and broken rice phosphate ester.Changsha:Hunan Agricultural University,2009:17-18.(in Chinese with English abstract)
[22]Smith A M,Zeeman S C,Smith S M.Starch degradation.Annual Review of Plant Biology,2005,56:73-98.
[23]Ritte G,Heydenreich M,Mahlow S,et al.Phosphorylation of C6-and C3-positions of glucosyl residues in starch is catalysed by distinct dikinases.FEBS Letters,2006,580(20):4872-4876.
[24]Glaring M A,Zygadlo A,Thorneycroft D,et al.An extraplastidialα-glucan,water dikinase from Arabidopsis phosphorylates amylopectin in vitro and is not necessary for transient starch degradation.Journal of Experimental Botany,2007,58(14):3949-3960.
[25]Orzechowski S,Grabowska A,Sitnicka D,et al.Analysis of the expression,subcellular and tissue localisation of phosphoglucan,water dikinase(PWD/GWD3)in Solanum tuberosum L.:a bioinformatics approach for the comparative analysis of twoα-glucan,water dikinases(GWDs)from Solanum tuberosum L.Acta Physiologiae Plantarum,2013,35(2):483-500.
[26]National Center for Biotechnology Information,US.National Library of Medicine8600 Rockville Pike,Bethesda MD,20894 USA[DB/OL].http://www.ncbi.nlm.nih.gov/genbank/.
[27]Frohberg C,Koetting O,Ritte G,et al.Methods for identifying proteins with starch phosphorylating enzymatic activity.US,8269063,2012-09-18.Washington DC:US.Patent and Trademark Office.
[28]Christiansen C,Hachem M A,Glaring M A,et al.A CBM20 low-affinity starch-binding domain from glucan,water dikinase.FEBS Letters,2009,583(7):1159-1163.
[29]Mikkelsen R,Baunsgaard L,Blennow A.Functional characterization of alpha-glucan,water dikinase,the starch phosphorylating enzyme.Biochemica Journal,2004,377:525-532.
[30]Mikkelsen R,Mutenda K E,Mant A,et al.α-Glucan,water dikinase(GWD):aplastidic enzyme with redox-regulated and coordinated catalytic activity and binding affinity.Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1785-1790.
[31]Hejazi M,Steup M,Fettke J.The plastidial glucan,water dikinase(GWD)catalyses multiple phosphotransfer reactions.FEBS Journal,2012,279(11):1953-1966.
[32]Hejazi M,Fettke J,Paris O,et al.The two plastidial starchrelated dikinases sequentially phosphorylate glucosyl residues at the surface of both the A-and B-type allomorphs of crystallized maltodextrins but the mode of action differs.Plant Physiology,2009,150(2):962-976.
[33]K?tting O,Santelia D,Edner C,et al.STARCH-EXCESS4 is a laforin-like phosphoglucan phosphatase required for starch degradation in Arabidopsis thaliana.The Plant Cell Online,2009,21(1):334-346.
[34]Blennow A,Engelsen S B.Helix-breaking news:fighting crystalline starch energy deposits in the cell.Trends in Plant Science,2010,15(4):236-240.
[35]Ritte G,Scharf A,Eckermann N,et al.Phosphorylation of transitory starch is increased during degradation.Plant Physiology,2004,135(4):2068-2077.
[36]Hejazi M,Fettke J,Steup M.Starch phosphorylation and dephosphorylation:the consecutive action of starch-related dikinases and phosphatases//Tetlow I J.Starch:Origins,Structure and Metabolism.London:Society of Experimental Biology,2012:279-310.
[37]Yu T S,Kofler H,Husler R E,et al.The Arabidopsis sex1 mutant is defective in the R1protein,ageneral regulator of starch degradation in plants,and not in the chloroplast hexose transporter.The Plant Cell Online,2001,13(8):1907-1918.
[38]Hirose T,Aoki N,Harada Y,et al.Disruption of a rice gene forα-glucan water dikinase,OsGWD1,leads to hyperaccumulation of starch in leaves but exhibits limited effects on growth.Frontiers in Plant Science,2013,4:147.
[39]Ral J P,Bowerman A F,Li Z,et al.Down-regulation of glucan,water-dikinase activity in wheat endosperm increases vegetative biomass and yield.Plant Biotechnology Journal,2012,10(7):871-882.
[40]Cenci U,Nitschke F,Steup M,et al.Transition from glycogen to starch metabolism in Archaeplastida.Trends in Plant Science,2014,19(1):18-28.
[41]Silver D M,K?tting O,Moorhead G G B.Phosphoglucan phosphatase function sheds light on starch degradation.Trends in Plant Science,2014,19(7):471-478.
[42]Roldán I,Wattebled F,Mercedes Lucas M,et al.The phenotype of soluble starch synthase Ⅳ defective mutants of Arabidopsis thaliana suggests a novel function of elongation enzymes in the control of starch granule formation.The Plant Journal,2014,49(3):492-504.
[43]Boriboonkaset T,Theerawitaya C,Yamada N,et al.Regulation of some carbohydrate metabolism-related genes,starch and soluble sugar contents,photosynthetic activities and yield attributes of two contrasting rice genotypes subjected to salt stress.Protoplasma,2013,250(5):1157-1167.
[44]Kaplan F,Guy C L.RNA interference of Arabidopsis betaamylase8prevents maltose accumulation upon cold shock and increases sensitivity of PSⅡphotochemical efficiency to freezing stress.The Plant Journal,2005,44(5):730-743.
[2]Fernbach A.Quelques Observations sur la Composition de l'amidon de Pommes de terre.Deslis Frères,1904,138:428-430.
[3]Lorberth R,Ritte G,Willmitzer L,et al.Inhibition of a starch-granule-bound protein leads to modified starch and repression of cold sweetening.Nature Biotechnology,1998,16(5):473-477.
[4]Ritte G,Lloyd J R,Eckermann N,et al.The starch-related R1protein is anα-glucan,water dikinase.Proceedings of the National Academy of Sciences of the United States of America,2002,99(10):7166-7171.
[5]Singh J,Kaur L,McCarthy O J.Factors influencing the physicochemical,morphological,thermal and rheological properties of some chemically modified starches for food applications:a review.Food Hydrocolloids,2007,21(1):1-22.
[6]Wei M,Liu Y,Liu B X,et al.Preparation and application of starch phosphate with a low degree of substitution.Phosphorus,Sulfur,and Silicon and the Related Elements,2011,186(4):974-982.
[7]Whistler R J,Bemiller J N,Pascal E F.Starch:Chemistry and Technology.Beijing:Chinese Food Press,1987:350-354.
[8]Blennow A,Jensen S L,Shaik S S,et al.Future cereal starch bioengineering:cereal ancestors encounter gene technology and designer enzymes.Cereal Chemistry,2013,90(4):274-287.
[9]Schewe G,Knies P,Amati S F,et al.Monocotyledon plant cells and plants which synthesise modified starch.US,6734340.2004-05-11[2014-04-30].
[10]Basu S,Lanahan M.Modified starch,uses,methods for production thereof.US,10/556276,2004-05-24[2014-04-30].
[11]Carciofi M,Shaik S S,Jensen S L,et al.Hyperphosphorylation of cereal starch.Journal of Cereal Science,2011,54(3):339-346.
[12]Frohberg C.Genetically modified plants which synthesize a starch having increased swelling power.US,8304606,2012-11-06[2014-04-30].
[13]Blennow A,Engelsen S B,Munck,L,et al.Starch molecular structure and phosphorylation investigated by a combined chromatographic and chemometric approach.Carbohydrate Polymers,2000,41(2):163-174.
[14]Blennow A,Nielsen T H,Baunsgaard L,et al.Starch phosphorylation:a new front line in starch research.Trends in Plant Science,2002,7(10):445-450.
[15]Zeeman S C,Kossmann J,Smith A M.Starch:its metabolism,evolution,and biotechnological modification in plants.Annual Review of Plant Biology,2010,61:209-234.
[16]Bay-Smidt A M,Wischmann B,Olsen C E,et al.Starch bound phosphate in potato as studied by a simple method for determination of organic phosphate and(31)P-NMR.Strke,1994,46(5):167-172.
[17]Blennow A,Bay-Smidt A M,Wischmann B,et al.The degree of starch phosphorylation is related to the chain length distribution of the neutral and the phosphorylated chains of amylopectin.Carbohydrate Research,1998,307(1):45-54.
[18]Hizukuri S,Takeda Y,Yasuda M,et al.Multi-branched nature of amylose and the action of debranching enzymes.Carbohydrate Research,1981,94(2):205-213.
[19]Mahlow S,Hejazi M,Kuhnert F,et al.Phosphorylation of transitory starch byα-glucan,water dikinase during starch turnover affects the surface properties and morphology of starch granules.New Phytologist,2014,203(2):495-507.
[20]Edner C,Li J,Albrecht T,et al.Glucan,water dikinase activity stimulates breakdown of starch granules by plastidialβ-amylases.Plant Physiology,2007,145(1):17-28.
[21]毛海锋.碎米淀粉及其磷酸酯的制备和理化性质研究.长沙:湖南农业大学,2009:17-18.Mao H F.Study on preparation and physicochemical properties of broken rice and broken rice phosphate ester.Changsha:Hunan Agricultural University,2009:17-18.(in Chinese with English abstract)
[22]Smith A M,Zeeman S C,Smith S M.Starch degradation.Annual Review of Plant Biology,2005,56:73-98.
[23]Ritte G,Heydenreich M,Mahlow S,et al.Phosphorylation of C6-and C3-positions of glucosyl residues in starch is catalysed by distinct dikinases.FEBS Letters,2006,580(20):4872-4876.
[24]Glaring M A,Zygadlo A,Thorneycroft D,et al.An extraplastidialα-glucan,water dikinase from Arabidopsis phosphorylates amylopectin in vitro and is not necessary for transient starch degradation.Journal of Experimental Botany,2007,58(14):3949-3960.
[25]Orzechowski S,Grabowska A,Sitnicka D,et al.Analysis of the expression,subcellular and tissue localisation of phosphoglucan,water dikinase(PWD/GWD3)in Solanum tuberosum L.:a bioinformatics approach for the comparative analysis of twoα-glucan,water dikinases(GWDs)from Solanum tuberosum L.Acta Physiologiae Plantarum,2013,35(2):483-500.
[26]National Center for Biotechnology Information,US.National Library of Medicine8600 Rockville Pike,Bethesda MD,20894 USA[DB/OL].http://www.ncbi.nlm.nih.gov/genbank/.
[27]Frohberg C,Koetting O,Ritte G,et al.Methods for identifying proteins with starch phosphorylating enzymatic activity.US,8269063,2012-09-18.Washington DC:US.Patent and Trademark Office.
[28]Christiansen C,Hachem M A,Glaring M A,et al.A CBM20 low-affinity starch-binding domain from glucan,water dikinase.FEBS Letters,2009,583(7):1159-1163.
[29]Mikkelsen R,Baunsgaard L,Blennow A.Functional characterization of alpha-glucan,water dikinase,the starch phosphorylating enzyme.Biochemica Journal,2004,377:525-532.
[30]Mikkelsen R,Mutenda K E,Mant A,et al.α-Glucan,water dikinase(GWD):aplastidic enzyme with redox-regulated and coordinated catalytic activity and binding affinity.Proceedings of the National Academy of Sciences of the United States of America,2005,102(5):1785-1790.
[31]Hejazi M,Steup M,Fettke J.The plastidial glucan,water dikinase(GWD)catalyses multiple phosphotransfer reactions.FEBS Journal,2012,279(11):1953-1966.
[32]Hejazi M,Fettke J,Paris O,et al.The two plastidial starchrelated dikinases sequentially phosphorylate glucosyl residues at the surface of both the A-and B-type allomorphs of crystallized maltodextrins but the mode of action differs.Plant Physiology,2009,150(2):962-976.
[33]K?tting O,Santelia D,Edner C,et al.STARCH-EXCESS4 is a laforin-like phosphoglucan phosphatase required for starch degradation in Arabidopsis thaliana.The Plant Cell Online,2009,21(1):334-346.
[34]Blennow A,Engelsen S B.Helix-breaking news:fighting crystalline starch energy deposits in the cell.Trends in Plant Science,2010,15(4):236-240.
[35]Ritte G,Scharf A,Eckermann N,et al.Phosphorylation of transitory starch is increased during degradation.Plant Physiology,2004,135(4):2068-2077.
[36]Hejazi M,Fettke J,Steup M.Starch phosphorylation and dephosphorylation:the consecutive action of starch-related dikinases and phosphatases//Tetlow I J.Starch:Origins,Structure and Metabolism.London:Society of Experimental Biology,2012:279-310.
[37]Yu T S,Kofler H,Husler R E,et al.The Arabidopsis sex1 mutant is defective in the R1protein,ageneral regulator of starch degradation in plants,and not in the chloroplast hexose transporter.The Plant Cell Online,2001,13(8):1907-1918.
[38]Hirose T,Aoki N,Harada Y,et al.Disruption of a rice gene forα-glucan water dikinase,OsGWD1,leads to hyperaccumulation of starch in leaves but exhibits limited effects on growth.Frontiers in Plant Science,2013,4:147.
[39]Ral J P,Bowerman A F,Li Z,et al.Down-regulation of glucan,water-dikinase activity in wheat endosperm increases vegetative biomass and yield.Plant Biotechnology Journal,2012,10(7):871-882.
[40]Cenci U,Nitschke F,Steup M,et al.Transition from glycogen to starch metabolism in Archaeplastida.Trends in Plant Science,2014,19(1):18-28.
[41]Silver D M,K?tting O,Moorhead G G B.Phosphoglucan phosphatase function sheds light on starch degradation.Trends in Plant Science,2014,19(7):471-478.
[42]Roldán I,Wattebled F,Mercedes Lucas M,et al.The phenotype of soluble starch synthase Ⅳ defective mutants of Arabidopsis thaliana suggests a novel function of elongation enzymes in the control of starch granule formation.The Plant Journal,2014,49(3):492-504.
[43]Boriboonkaset T,Theerawitaya C,Yamada N,et al.Regulation of some carbohydrate metabolism-related genes,starch and soluble sugar contents,photosynthetic activities and yield attributes of two contrasting rice genotypes subjected to salt stress.Protoplasma,2013,250(5):1157-1167.
[44]Kaplan F,Guy C L.RNA interference of Arabidopsis betaamylase8prevents maltose accumulation upon cold shock and increases sensitivity of PSⅡphotochemical efficiency to freezing stress.The Plant Journal,2005,44(5):730-743.