植物紫色酸性磷酸酶的研究进展
|
摘要
酸性磷酸酶(APase)是酸性条件下(pH <7.0)能催化磷酸单酯或酸酐裂解从而释放无机磷酸根离子的水解酶类。紫色酸性磷酸酶(Purple acid phosphatase,PAP)是一类特殊的酸性磷酸酶,其具有鲜明的特征,如:酶的提取液呈紫色或粉色、酶活性不受酒石酸盐抑制、氨基酸序列具有5个保守结构域和双金属离子催化中心等。已有的研究表明,紫色酸性磷酸酶在植物适应低磷胁迫过程中发挥着重要作用。本文综述了紫色酸性磷酸酶的生化特性、亚细胞定位、生物学功能以及最新研究进展。
Acid phosphatase(APase) belongs to the hydrolase enzyme, which hydrolyzes orthophosphate from phosphomonoester or anhydride with an acidic pH optimum(pH < 7.0). Among APase, purple acid phosphatase(PAP) has distinctive biochemical and molecular characteristics, such as appearing purple or pink in extracted enzyme solutions,insensitive activity to tartrate inhibition, harboring of a binuclear metal center and five conserved motif blocks in amino acid sequences. It has been documented that PAP plays an important part in plant adaptation to phosphorus deficiency.In this review, we summarized the biochemical characterization, subcellular localization, biological function, and recent research progress of PAP in plants.
Acid phosphatase(APase) belongs to the hydrolase enzyme, which hydrolyzes orthophosphate from phosphomonoester or anhydride with an acidic pH optimum(pH < 7.0). Among APase, purple acid phosphatase(PAP) has distinctive biochemical and molecular characteristics, such as appearing purple or pink in extracted enzyme solutions,insensitive activity to tartrate inhibition, harboring of a binuclear metal center and five conserved motif blocks in amino acid sequences. It has been documented that PAP plays an important part in plant adaptation to phosphorus deficiency.In this review, we summarized the biochemical characterization, subcellular localization, biological function, and recent research progress of PAP in plants.
引文
[1]梁翠月,廖红.植物根系响应低磷胁迫的机理研究[J].生命科学,2015,27(3):389-397.
[2]Vance C P,Uhde-Stone C,Allan D L.Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource[J].New Phytologist,2003,157:423-447.
[3]López-Arredondo D L,Leyva-González M A,González-Morales S I,et al.Phosphate nutrition:improving low-phosphate tolerance in crops[J].Annual Review of Plant Biology,2014,65:95-123.
[4]Shen J,Yuan L,Zhang J,et al.Phosphorus dynamics:from soil to plant[J].Plant Physiology,2011,156:997-1005.
[5]黄宇,张海伟,徐芳森.植物酸性磷酸酶的研究进展[J].华中农业大学学报,2008,27(1):148-154.
[6]Wang L,Liu D.Functions and regulation of phosphate starvation-induced secreted acid phosphatases in higher plants[J].Plant Science,2018,271:108-116.
[7]Tian J,Liao H.The role of intracellular and secreted purple acid phosphatases in plant phosphorus scavenging and recycling[M]//Plaxton W C,Lambers H.Annual Plant Reviews,Volume 48:Phosphorus metabolism in plants.Hoboken,New Jersey:Wiley-Blackwell,2015:265-287.
[8]Zhang Q,Wang C,Tian J,et al.Identification of rice purple acid phosphatases related to phosphate starvation signalling[J].Plant Biology,2011,13:7-15.
[9]George T S,Gregory P J,Hocking P,et al.Variation in root-associated phosphatase activities in wheat contributes to the utilization of organic P substrates in vitro,but does not explain differences in the P-nutrition of plants when grown in soils[J].Environmental and Experimental Botany,2008,64:239-249.
[10]González-Mu?oz E,Avenda?o-Vázquez A,Montes R A C,et al.The Maize(Zea mays ssp.mays var.B73)genome encodes 33 members of the purple acid phosphatase gene family[J].Frontiers in Plant Science,2015,6:341.
[11]Li C,Gui S,Yang T,et al.Identification of soybean purple acid phosphatase genes and their expression responses to phosphorus availability and symbiosis[J].Annals of Botany,2012,109:275-285.
[12]Liang C,Tian J,Lam H M,et al.Biochemical and molecular characterization of PvPAP3,a novel purple acid phosphatase isolated from common bean enhancing extracellular ATPutilization[J].Plant Physiology,2010,152:854-865.
[13]刘攀道,董荣书,丁西朋,等.不同磷效率柱花草基因型对外源DNA活化利用能力的比较分析[J].分子植物育种,2018,16(4):1085-1091.
[14]Bozzo G G,Raghothama K G,Plaxton W C.Structural and kinetic properties of a novel purple acid phosphatase from phosphate-starved tomato(Lycopersicon esculentum)cell cultures[J].Biochemical Journal,2004,377:419-428.
[15]Tran H T,Qian W,Hurley B A,et al.Biochemical and molecular characterization of At PAP12 and At PAP26:the predominant purple acid phosphatase isozymes secreted by phosphate-starved Arabidopsis thaliana[J].Plant,Cell&Environment,2010,33:1789-1803.
[16]Miller S S,Liu J,Allan D L,et al.Molecular control of acid phosphatase secretion into the rhizosphere of proteoid roots from phosphorus-stressed white lupin[J].Plant Physiology,2001,127:594-606.
[17]Li D,Zhu H,Liu K,et al.Purple acid phosphatases of Arabidopsis thaliana:comparative analysis and differential regulation by phosphate deprivation[J].Journal of Biological Chemistry,2002,277:27772-27781.
[18]Bhadouria J,Singh A P,Mehra P,et al.Identification of purple acid phosphatases in chickpea and potential roles of Ca PAP7 in seed phytate accumulation[J].Scientific Reports,2017,7:11012.
[19]Beck J L,McConachie L A,Summors A C,et al.Properties of a purple phosphatase from red kidney bean:a zinc-iron metalloenzyme[J].Biochimica et Biophysica Acta(BBA)-Protein Structure and Molecular Enzymology,1986,869:61-68.
[20]Schenk G,Guddat L W,Ge Y,et al.Identification of mammalian-like purple acid phosphatases in a wide range of plants[J].Gene,2000,250:117-125.
[21]Schenk G,Korsinczky M L J,Hume D A,et al.Purple acid phosphatases from bacteria:similarities to mammalian and plant enzymes[J].Gene,2000,255:419-424.
[22]Flanagan J U,Cassady A I,Schenk G,et al.Identification and molecular modeling of a novel,plant-like,human purple acid phosphatase[J].Gene,2006,377:12-20.
[23]Schenk G,Mitic N,Hanson G R,et al.Purple acid phosphatase:a journey into the function and mechanism of a colorful enzyme[J].Coordination Chemistry Reviews,2013,257:473-482.
[24]Tran H T,Hurley B A,Plaxton W C.Feeding hungry plants:the role of purple acid phosphatases in phosphate nutrition[J].Plant Science,2010,179:14-27.
[25]Durmus A,Eicken C,Sift B H,et al.The active site of purple acid phosphatase from aweet potatoes(Ipomoea batatas):metal content and spectroscopic characterization[J].European Journal of Biochemistry,1999,260:709-716.
[26]Schenk G,Ge Y,Carrington L E,et al.Binuclear metal centers in plant purple acid phosphatases:Fe-Mn in sweet potato and Fe-Zn in soybean[J].Archives of Biochemistry and Biophysics,1999,370:183-189.
[27]Schenk G,Gahan L R,Carrington L E,et al.Phosphate forms an unusual tripodal complex with the Fe-Mn center of sweet potato purple acid phosphatase[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102:273-278.
[28]Antonyuk S V,Olczak M,Olczak T,et al.The structure of a purple acid phosphatase involved in plant growth and pathogen defence exhibits a novel immunoglobulin-like fold[J].IUCrJ,2014,1:101-109.
[29]Wang L,Li Z,Qian W,et al.The Arabidopsis purple acid phosphatase At PAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation[J].Plant Physiology,2011,157:1283-1299.
[30]Del Vecchio H A,Ying S,Park J,et al.The cell wall-targeted purple acid phosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation[J].The Plant Journal,2014,80:569-581.
[31]Yoneyama T,Shiozawa M,Nakamura M,et al.Characterization of a novel acid phosphatase from embryonic axes of kidney bean exhibiting vanadate-dependent chloroperoxidase activity[J].Journal of Biological Chemistry,2004,279:37477-37484.
[32]Cashikar A G,Kumaresan R,Rao N M.Biochemical characterization and subcellular localization of the red kidney bean purple acid phosphatase[J].Plant Physiology,1997,114:907-915.
[33]Pintus F,Spano D,Corongiu S,et al.Purification,primary structure,and properties of Euphorbia characias latex purple acid phosphatase[J].Biochemistry(Moscow),2011,76:694-701.
[34]Shinano T,Yonetani R,Ushihara N,et al.Characteristics of phosphoenolpyruvate phosphatase purified from Allium cepa[J].Plant Science,2001,161:861-869.
[35]Zhang W,Gruszewski H A,Chevone B I,et al.An Arabidopsis purple acid phosphatase with phytase activity increases foliar ascorbate[J].Plant Physiology,2008,146:431-440.
[36]Zhu H F,Qian W Q,Lu X Z,et al.Expression patterns of purple acid phosphatase genes in Arabidopsis organs and functional analysis of AtPAP23 predominantly transcribed in flower[J].Plant Molecular Biology,2005,59:581-594.
[37]Dionisio G,Madsen C K,Holm P B,et al.Cloning and characterization of purple acid phosphatase phytases from wheat,barley,maize,and rice[J].Plant Physiology,2011,156:1087-1100.
[38]Lung S C,Leung A,Kuang R,et al.Phytase activity in tobacco(Nicotiana tabacum)root exudates is exhibited by a purple acid phosphatase[J].Phytochemistry,2008,69:365-373.
[39]Hegeman C E,Grabau E A.A novel phytase with sequence similarity to purple acid phosphatases is expressed in cotyledons of germinating soybean seedlings[J].Plant Physiology,2001,126:1598-1608.
[40]Wang J,Si Z,Li F,et al.A purple acid phosphatase plays a role in nodule formation and nitrogen fixation in Astragalus sinicus[J].Plant Molecular Biology,2015,88:515-529.
[41]Kaneko K,Inomata T,Masui T,et al.Nucleotide Pyrophosphatase/Phosphodiesterase 1 exerts a negative effect on starch accumulation and growth in rice seedlings under high temperature and CO2 concentration conditions[J].Plant&Cell Physiology,2014,55:320-332.
[42]Lebansky B R,Mc Knight T D,Griffing L R.Purification and characterization of a secreted purple phosphatase from soybean suspension cultures[J].Plant Physiology,1992,99:391-395.
[43]Wu W,Lin Y,Liu P,et al.Association of extracellular d NTP utilization with a GmPAP1-like protein identified in cell wall proteomic analysis of soybean roots[J].Journal of Experimental Botany,2018,69:603-617.
[44]Lu L,Qiu W,Gao W,et al.OsPAP10c,a novel secreted acid phosphatase in rice,plays an important role in the utilization of external organic phosphorus[J].Plant,Cell&Environment,2016,39:2247-2259.
[45]Mehra P,Pandey B K,Giri J.Improvement in phosphate acquisition and utilization by a secretory purple acid phosphatase(OsPAP21b)in rice[J].Plant Biotechnology Journal,2017,15:1054-1067.
[46]Nishikoori M,Washio K,Hase A,et al.Cloning and characterization of c DNA of the GPI-Anchored purple acid phosphatase and its root tissue distribution in Spirodela Oligorrhiza[J].Physiologia Plantarum,2001,113:241-248.
[47]Liang C,Sun L,Yao Z,et al.Comparative analysis of Pv PAP gene family and their functions in response to phosphorus deficiency in common bean[J].PLo S One,2012,7:65-65.
[48]Liu P D,Xue Y B,Chen Z J,et al.Characterization of purple acid phosphatases involved in extracellular dNTP utilization in Stylosanthes[J].Journal of Experimental Botany,2016,67:4141-4154.
[49]Hurley B A,Tran H T,Marty N J,et al.The dual-targeted purple acid phosphatase isozyme At PAP26 is essential for efficient acclimation of Arabidopsis to nutritional phosphate deprivation[J].Plant Physiology,2010,153:1112-1122.
[50]Kataya A R A,Schei E,Lillo C.Towards understanding peroxisomal phosphoregulation in Arabidopsis thaliana[J].Planta,2016,243:699-717.
[51]Sun F,Suen P K,Zhang Y,et al.A dual-targeted purple acid phosphatase in Arabidopsis thaliana moderates carbon metabolism and its overexpression leads to faster plant growth and higher seed yield[J].New Phytologist,2012,194:206-219.
[52]Sun Q,Li J,Cheng W,et al.At PAP2,a unique member of the PAP family,functions in the plasma membrane[J].Genes.2018,9(5).
[53]Robinson W D,Park J,Tran H T,et al.The secreted purple acid phosphatase isozymes At PAP12 and At PAP26 play a pivotal role in extracellular phosphate-scavenging by Arabidopsis thaliana[J].Journal of Experimental Botany,2012,63:6531-6542.
[54]Tian J,Wang C,Zhang Q,et al.Overexpression of Os PAP10a,a root-associated acid phosphatase,increased extracellular organic phosphorus utilization in rice[J].Journal of Integrative Plant Biology,2012,54:631-639.
[55]Xiao K,Harrison M J,Wang Z Y.Transgenic expression of a novel M.truncatula phytase gene results in improved acquisition of organic phosphorus by Arabidopsis[J].Planta,2005,222:27-36.
[56]Kong Y,Li X,Ma J,et al.GmPAP4,a novel purple acid phosphatase gene isolated from soybean(Glycine max),enhanced extracellular phytate utilization in Arabidopsis thaliana[J].Plant Cell Reports,2014,33:655-667.
[57]Kong Y,Li X,Wang B,et al.The soybean purple acid phosphatase Gm PAP14 predominantly enhances external phytate utilization in plants[J].Frontiers in Plant Science,2018,9:292.
[58]Robinson W D,Carson I,Ying S,et al.Eliminating the purple acid phosphatase At PAP26 in Arabidopsis thaliana delays leaf senescence and impairs phosphorus remobilization[J].New Phytologist,2012,196:1024-1029.
[59]Kaida R,Satoh Y,Bulone V,et al.Activation ofβ-glucan synthases by wall-bound purple acid phosphatase in tobacco cells[J].Plant Physiology,2009,150:1822-1830.
[60]Ravichandran S,Stone S L,Benkel B,et al.Purple acid phosphatase 5 is required for maintaining basal resistance against pseudomonas syringae in Arabidopsis[J].BMC Plant Biology,2013,13:1-12.
[61]Li W F,Shao G,Lam H M.Ectopic expression of GmPAP3alleviates oxidative damage caused by salinity and osmotic stresses[J].New Phytologist,2008,178:80-91.
[62]Kuang R,Chan K,Yeung E,et al.Molecular and biochemical characterization of At PAP15,a purple acid phosphatase with phytase activity,in Arabidopsis[J].Plant Physiology,2009,151:199-209.
[63]Zhang Y,Yu L,Yung K F,et al.Over-expression of AtPAP2 in Camelina sativa leads to faster plant growth and higher seed yield[J].Biotechnology for Biofuels,2012,5:19.
[64]Zhang Y,Sun F,Fettke J,et al.Heterologous expression of AtPAP2 in transgenic potato influences carbon metabolism and tuber development[J].FEBS Letters,2014,588:3726-3731.
[65]Sun L,Song L,Zhang Y,et al.Arabidopsis PHL2 and PHR1act redundantly as the key components of the central regulatory system controlling transcriptional responses to phosphate starvation[J].Plant Physiology,2016,170:499-514.
[66]Del Pozo J C,Allona I,Rubio V,et al.A type 5 acid phosphatase gene from Arabidopsis thaliana is induced by phosphate starvation and by some other types of phosphate mobilising/oxidative stress conditions[J].The Plant Journal,1999,19:579-589.
[2]Vance C P,Uhde-Stone C,Allan D L.Phosphorus acquisition and use:critical adaptations by plants for securing a nonrenewable resource[J].New Phytologist,2003,157:423-447.
[3]López-Arredondo D L,Leyva-González M A,González-Morales S I,et al.Phosphate nutrition:improving low-phosphate tolerance in crops[J].Annual Review of Plant Biology,2014,65:95-123.
[4]Shen J,Yuan L,Zhang J,et al.Phosphorus dynamics:from soil to plant[J].Plant Physiology,2011,156:997-1005.
[5]黄宇,张海伟,徐芳森.植物酸性磷酸酶的研究进展[J].华中农业大学学报,2008,27(1):148-154.
[6]Wang L,Liu D.Functions and regulation of phosphate starvation-induced secreted acid phosphatases in higher plants[J].Plant Science,2018,271:108-116.
[7]Tian J,Liao H.The role of intracellular and secreted purple acid phosphatases in plant phosphorus scavenging and recycling[M]//Plaxton W C,Lambers H.Annual Plant Reviews,Volume 48:Phosphorus metabolism in plants.Hoboken,New Jersey:Wiley-Blackwell,2015:265-287.
[8]Zhang Q,Wang C,Tian J,et al.Identification of rice purple acid phosphatases related to phosphate starvation signalling[J].Plant Biology,2011,13:7-15.
[9]George T S,Gregory P J,Hocking P,et al.Variation in root-associated phosphatase activities in wheat contributes to the utilization of organic P substrates in vitro,but does not explain differences in the P-nutrition of plants when grown in soils[J].Environmental and Experimental Botany,2008,64:239-249.
[10]González-Mu?oz E,Avenda?o-Vázquez A,Montes R A C,et al.The Maize(Zea mays ssp.mays var.B73)genome encodes 33 members of the purple acid phosphatase gene family[J].Frontiers in Plant Science,2015,6:341.
[11]Li C,Gui S,Yang T,et al.Identification of soybean purple acid phosphatase genes and their expression responses to phosphorus availability and symbiosis[J].Annals of Botany,2012,109:275-285.
[12]Liang C,Tian J,Lam H M,et al.Biochemical and molecular characterization of PvPAP3,a novel purple acid phosphatase isolated from common bean enhancing extracellular ATPutilization[J].Plant Physiology,2010,152:854-865.
[13]刘攀道,董荣书,丁西朋,等.不同磷效率柱花草基因型对外源DNA活化利用能力的比较分析[J].分子植物育种,2018,16(4):1085-1091.
[14]Bozzo G G,Raghothama K G,Plaxton W C.Structural and kinetic properties of a novel purple acid phosphatase from phosphate-starved tomato(Lycopersicon esculentum)cell cultures[J].Biochemical Journal,2004,377:419-428.
[15]Tran H T,Qian W,Hurley B A,et al.Biochemical and molecular characterization of At PAP12 and At PAP26:the predominant purple acid phosphatase isozymes secreted by phosphate-starved Arabidopsis thaliana[J].Plant,Cell&Environment,2010,33:1789-1803.
[16]Miller S S,Liu J,Allan D L,et al.Molecular control of acid phosphatase secretion into the rhizosphere of proteoid roots from phosphorus-stressed white lupin[J].Plant Physiology,2001,127:594-606.
[17]Li D,Zhu H,Liu K,et al.Purple acid phosphatases of Arabidopsis thaliana:comparative analysis and differential regulation by phosphate deprivation[J].Journal of Biological Chemistry,2002,277:27772-27781.
[18]Bhadouria J,Singh A P,Mehra P,et al.Identification of purple acid phosphatases in chickpea and potential roles of Ca PAP7 in seed phytate accumulation[J].Scientific Reports,2017,7:11012.
[19]Beck J L,McConachie L A,Summors A C,et al.Properties of a purple phosphatase from red kidney bean:a zinc-iron metalloenzyme[J].Biochimica et Biophysica Acta(BBA)-Protein Structure and Molecular Enzymology,1986,869:61-68.
[20]Schenk G,Guddat L W,Ge Y,et al.Identification of mammalian-like purple acid phosphatases in a wide range of plants[J].Gene,2000,250:117-125.
[21]Schenk G,Korsinczky M L J,Hume D A,et al.Purple acid phosphatases from bacteria:similarities to mammalian and plant enzymes[J].Gene,2000,255:419-424.
[22]Flanagan J U,Cassady A I,Schenk G,et al.Identification and molecular modeling of a novel,plant-like,human purple acid phosphatase[J].Gene,2006,377:12-20.
[23]Schenk G,Mitic N,Hanson G R,et al.Purple acid phosphatase:a journey into the function and mechanism of a colorful enzyme[J].Coordination Chemistry Reviews,2013,257:473-482.
[24]Tran H T,Hurley B A,Plaxton W C.Feeding hungry plants:the role of purple acid phosphatases in phosphate nutrition[J].Plant Science,2010,179:14-27.
[25]Durmus A,Eicken C,Sift B H,et al.The active site of purple acid phosphatase from aweet potatoes(Ipomoea batatas):metal content and spectroscopic characterization[J].European Journal of Biochemistry,1999,260:709-716.
[26]Schenk G,Ge Y,Carrington L E,et al.Binuclear metal centers in plant purple acid phosphatases:Fe-Mn in sweet potato and Fe-Zn in soybean[J].Archives of Biochemistry and Biophysics,1999,370:183-189.
[27]Schenk G,Gahan L R,Carrington L E,et al.Phosphate forms an unusual tripodal complex with the Fe-Mn center of sweet potato purple acid phosphatase[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102:273-278.
[28]Antonyuk S V,Olczak M,Olczak T,et al.The structure of a purple acid phosphatase involved in plant growth and pathogen defence exhibits a novel immunoglobulin-like fold[J].IUCrJ,2014,1:101-109.
[29]Wang L,Li Z,Qian W,et al.The Arabidopsis purple acid phosphatase At PAP10 is predominantly associated with the root surface and plays an important role in plant tolerance to phosphate limitation[J].Plant Physiology,2011,157:1283-1299.
[30]Del Vecchio H A,Ying S,Park J,et al.The cell wall-targeted purple acid phosphatase AtPAP25 is critical for acclimation of Arabidopsis thaliana to nutritional phosphorus deprivation[J].The Plant Journal,2014,80:569-581.
[31]Yoneyama T,Shiozawa M,Nakamura M,et al.Characterization of a novel acid phosphatase from embryonic axes of kidney bean exhibiting vanadate-dependent chloroperoxidase activity[J].Journal of Biological Chemistry,2004,279:37477-37484.
[32]Cashikar A G,Kumaresan R,Rao N M.Biochemical characterization and subcellular localization of the red kidney bean purple acid phosphatase[J].Plant Physiology,1997,114:907-915.
[33]Pintus F,Spano D,Corongiu S,et al.Purification,primary structure,and properties of Euphorbia characias latex purple acid phosphatase[J].Biochemistry(Moscow),2011,76:694-701.
[34]Shinano T,Yonetani R,Ushihara N,et al.Characteristics of phosphoenolpyruvate phosphatase purified from Allium cepa[J].Plant Science,2001,161:861-869.
[35]Zhang W,Gruszewski H A,Chevone B I,et al.An Arabidopsis purple acid phosphatase with phytase activity increases foliar ascorbate[J].Plant Physiology,2008,146:431-440.
[36]Zhu H F,Qian W Q,Lu X Z,et al.Expression patterns of purple acid phosphatase genes in Arabidopsis organs and functional analysis of AtPAP23 predominantly transcribed in flower[J].Plant Molecular Biology,2005,59:581-594.
[37]Dionisio G,Madsen C K,Holm P B,et al.Cloning and characterization of purple acid phosphatase phytases from wheat,barley,maize,and rice[J].Plant Physiology,2011,156:1087-1100.
[38]Lung S C,Leung A,Kuang R,et al.Phytase activity in tobacco(Nicotiana tabacum)root exudates is exhibited by a purple acid phosphatase[J].Phytochemistry,2008,69:365-373.
[39]Hegeman C E,Grabau E A.A novel phytase with sequence similarity to purple acid phosphatases is expressed in cotyledons of germinating soybean seedlings[J].Plant Physiology,2001,126:1598-1608.
[40]Wang J,Si Z,Li F,et al.A purple acid phosphatase plays a role in nodule formation and nitrogen fixation in Astragalus sinicus[J].Plant Molecular Biology,2015,88:515-529.
[41]Kaneko K,Inomata T,Masui T,et al.Nucleotide Pyrophosphatase/Phosphodiesterase 1 exerts a negative effect on starch accumulation and growth in rice seedlings under high temperature and CO2 concentration conditions[J].Plant&Cell Physiology,2014,55:320-332.
[42]Lebansky B R,Mc Knight T D,Griffing L R.Purification and characterization of a secreted purple phosphatase from soybean suspension cultures[J].Plant Physiology,1992,99:391-395.
[43]Wu W,Lin Y,Liu P,et al.Association of extracellular d NTP utilization with a GmPAP1-like protein identified in cell wall proteomic analysis of soybean roots[J].Journal of Experimental Botany,2018,69:603-617.
[44]Lu L,Qiu W,Gao W,et al.OsPAP10c,a novel secreted acid phosphatase in rice,plays an important role in the utilization of external organic phosphorus[J].Plant,Cell&Environment,2016,39:2247-2259.
[45]Mehra P,Pandey B K,Giri J.Improvement in phosphate acquisition and utilization by a secretory purple acid phosphatase(OsPAP21b)in rice[J].Plant Biotechnology Journal,2017,15:1054-1067.
[46]Nishikoori M,Washio K,Hase A,et al.Cloning and characterization of c DNA of the GPI-Anchored purple acid phosphatase and its root tissue distribution in Spirodela Oligorrhiza[J].Physiologia Plantarum,2001,113:241-248.
[47]Liang C,Sun L,Yao Z,et al.Comparative analysis of Pv PAP gene family and their functions in response to phosphorus deficiency in common bean[J].PLo S One,2012,7:65-65.
[48]Liu P D,Xue Y B,Chen Z J,et al.Characterization of purple acid phosphatases involved in extracellular dNTP utilization in Stylosanthes[J].Journal of Experimental Botany,2016,67:4141-4154.
[49]Hurley B A,Tran H T,Marty N J,et al.The dual-targeted purple acid phosphatase isozyme At PAP26 is essential for efficient acclimation of Arabidopsis to nutritional phosphate deprivation[J].Plant Physiology,2010,153:1112-1122.
[50]Kataya A R A,Schei E,Lillo C.Towards understanding peroxisomal phosphoregulation in Arabidopsis thaliana[J].Planta,2016,243:699-717.
[51]Sun F,Suen P K,Zhang Y,et al.A dual-targeted purple acid phosphatase in Arabidopsis thaliana moderates carbon metabolism and its overexpression leads to faster plant growth and higher seed yield[J].New Phytologist,2012,194:206-219.
[52]Sun Q,Li J,Cheng W,et al.At PAP2,a unique member of the PAP family,functions in the plasma membrane[J].Genes.2018,9(5).
[53]Robinson W D,Park J,Tran H T,et al.The secreted purple acid phosphatase isozymes At PAP12 and At PAP26 play a pivotal role in extracellular phosphate-scavenging by Arabidopsis thaliana[J].Journal of Experimental Botany,2012,63:6531-6542.
[54]Tian J,Wang C,Zhang Q,et al.Overexpression of Os PAP10a,a root-associated acid phosphatase,increased extracellular organic phosphorus utilization in rice[J].Journal of Integrative Plant Biology,2012,54:631-639.
[55]Xiao K,Harrison M J,Wang Z Y.Transgenic expression of a novel M.truncatula phytase gene results in improved acquisition of organic phosphorus by Arabidopsis[J].Planta,2005,222:27-36.
[56]Kong Y,Li X,Ma J,et al.GmPAP4,a novel purple acid phosphatase gene isolated from soybean(Glycine max),enhanced extracellular phytate utilization in Arabidopsis thaliana[J].Plant Cell Reports,2014,33:655-667.
[57]Kong Y,Li X,Wang B,et al.The soybean purple acid phosphatase Gm PAP14 predominantly enhances external phytate utilization in plants[J].Frontiers in Plant Science,2018,9:292.
[58]Robinson W D,Carson I,Ying S,et al.Eliminating the purple acid phosphatase At PAP26 in Arabidopsis thaliana delays leaf senescence and impairs phosphorus remobilization[J].New Phytologist,2012,196:1024-1029.
[59]Kaida R,Satoh Y,Bulone V,et al.Activation ofβ-glucan synthases by wall-bound purple acid phosphatase in tobacco cells[J].Plant Physiology,2009,150:1822-1830.
[60]Ravichandran S,Stone S L,Benkel B,et al.Purple acid phosphatase 5 is required for maintaining basal resistance against pseudomonas syringae in Arabidopsis[J].BMC Plant Biology,2013,13:1-12.
[61]Li W F,Shao G,Lam H M.Ectopic expression of GmPAP3alleviates oxidative damage caused by salinity and osmotic stresses[J].New Phytologist,2008,178:80-91.
[62]Kuang R,Chan K,Yeung E,et al.Molecular and biochemical characterization of At PAP15,a purple acid phosphatase with phytase activity,in Arabidopsis[J].Plant Physiology,2009,151:199-209.
[63]Zhang Y,Yu L,Yung K F,et al.Over-expression of AtPAP2 in Camelina sativa leads to faster plant growth and higher seed yield[J].Biotechnology for Biofuels,2012,5:19.
[64]Zhang Y,Sun F,Fettke J,et al.Heterologous expression of AtPAP2 in transgenic potato influences carbon metabolism and tuber development[J].FEBS Letters,2014,588:3726-3731.
[65]Sun L,Song L,Zhang Y,et al.Arabidopsis PHL2 and PHR1act redundantly as the key components of the central regulatory system controlling transcriptional responses to phosphate starvation[J].Plant Physiology,2016,170:499-514.
[66]Del Pozo J C,Allona I,Rubio V,et al.A type 5 acid phosphatase gene from Arabidopsis thaliana is induced by phosphate starvation and by some other types of phosphate mobilising/oxidative stress conditions[J].The Plant Journal,1999,19:579-589.
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |