硝酸还原酶活性对小白菜硝酸盐积累及相关代谢调节的研究
摘要
目前,蔬菜中硝酸盐的积累引起了许多国家的重视,环境卫生学大量试验证明,硝酸盐可以通过细菌的作用在动物体内外还原成亚硝酸盐,亚硝酸盐不仅会使人体中毒缺氧导致高铁血红蛋白症,还会导致人体消化系统的癌变等。由于人体摄取的硝酸盐有81.2%来源于蔬菜,小白菜(Brassica campestris L.ssp Chinesis Makino)是极易富集硝酸盐的蔬菜,又是南方地区占蔬菜上市量达33%的大众化蔬菜,因此研究硝酸盐在小白菜中的积累具有重要意义。影响硝酸盐积累的内外因素很多,其中,硝酸还原酶活性(nitrate reductase activitM NRA)是硝酸盐还原的主要限速因子。
小白菜品种间的硝酸盐含量和硝酸还原酶活性差异显著,本研究主要采用“上海青、苏州青、兴化青梗菜、引02、青丰一号、兔子腿矮脚黄”的高代自交系为试验材料,利用水溶液培养法,围绕NRA对小白菜硝酸盐积累及生理机制影响进行了研究,
结果表明:
1.氮素形态对小白菜体内硝酸盐积累和NRA的影响
在全硝态氮供应的N1处理组的硝酸盐含量显著高于硝/铵等比处理的N2组和全铵态氮处理的N3组;N1组的NRA略高于N2组,并无显著性差异,但N1组和N2组的NRA均极显著高于N3组,全铵态氮的N3处理组硝酸盐含量和NRA都最低。N2处理组的品质和产量最高,N1略低于N2处理组,而N3的品质和产量最低。在全硝态氮供应下,根系对NO_3~-的吸收速率最高,而随着铵态氮比例的增加,其吸收速率随之降低。
2.根部NRA对小白菜硝酸盐和可溶性糖代谢的影响
根系NRA受抑的情况下根系对NO_3~-吸收速率显著下降,而地上部的硝酸盐含量并无显著差异。同时,根系NRA受抑条件下,其根系可溶性糖浓度增加,而地上部可溶性糖浓度则相对减少。
3.小白菜光合性能与NRA关系
1)光照强度是制约NRA和硝酸盐含量的主要因素,尤其对NRA的高低影响极显著。2)NRA的降低会对光合速率产生一定的反作用。而叶绿素含量、气孔导度、胞间CO_2浓度和蒸腾速率在NRA抑制前后无显著变化,不受NRA的直接影响。3)在一定的硝态氮浓度范围内(=8mmol·L~(1)),NRA的升高对净光合速率的升高有促进作用。以上结论表明,光合性能与NRA存在着一定的相互促进和相互制约的关系。
4.钼锰不同施用方式对小白菜NRA和硝酸盐含量的影响
钼显著提高了小白菜的NRA,使NO_3~-显著降低,锰的施用,对NRA的影响不明显,但可以使硝酸盐含量降低,钼和锰的施用均可提高可溶性糖含量和Vc含量,与钼相比,锰对可溶性糖含量和Vc含量的提高更为显著。钼锰配施对提高NRA和降低硝酸盐含量最为明显,同时配施时对可溶性糖含量和Vc含量的提高也最为明显。叶面喷施与营养液根部结合施用对各生理指标的影响效果要明显高于叶面施用而略高于营养液施用。
Concentrations of nitrate in vegetables have been the focus of attention in several countries. A lot of reaserch on environment hygienics show that nitrate may reduce to nitrite which can lead to some diease such as cancer in animals. The nitrate absorbed by human is 81.2% derived from vegetable. Furthermore, nitrate was very easy accumulated in pakchoi which is a very popular vegetable which occupied 33% of the whole vegetable market in China, and NRA (Nitrate Reductase Activity) was a main limit factor in nitrate reduced.
In this study soluble culture was used for physiological mechanism of nitrate accumulation and NRA in pakchoi. There was significant difference of nitrate content and NRA between different genotypes of vegetable crops. Six genotypes were used in this study: Shanghaiqing, Suzhouqing, Xinghua qinggengcai, Yin02,QingfengNo.l,Tuzitui aijiaohuang. The results went as followed:
1 Effects of nitrogen forms on nitrate content and nitrate reductase of pak-choi
The content of nitrate in the N1 system which was offered pure NO_3~- is obviously higher than the system of N2 (NO_3~-/NH_4~+=1/1) and N3(NO_3~-/NH_4~+=0/1).NRA in N1 has a little higher than N2, but NRA in N1 and N2 is much obviously higher than N3.The nitrate content and NRA is the lowest in N3 which is treat with pure ammonium. Qualities and yield of N2 are the highest, N1 is a little lower than N2, N3 is the lowest neither in qualities or yield. The absorb rate of NO_3~- in root is the fastest when pure nitrate was offered, but the absorb rate was decreasing with the ammonium increasing.
2 Effect of NRA in root on the absorb and metabolism in nitrate of pakchoi
The absorb rate of NO_3~- reduced by inhibiting NRA in root, but there is no significant dramatically difference of the nitrate content. Furthermore, if we inhibit NRA in root,the concentration of soluble sugar increased and the concentration reduced in shoot.
3 Studies on the relationship of NRA and photosynthetic characters of pakchoi
The main characters of NRA and nitrate content controlling factor are light intensity especially on NRA. In addition, NRA reduced can feed back the photosynthetic, but chlorophyll content, Cond and Ci did not change obviously after NRA reduced. What's more, the grown of NRA can promote the photosynthetic when the nitrate content is lower than 8mmol·L~(-1).To sum up,the conclusion shows that photosynthetic characters and NRA have some relationship both in promotion and restriction.
4 Effects of Molybdenum and Manganese on NRA and nitrate content
Mo was significantly improved NRA and reduced nitrate in two genotype of pakchoi. However, Mn could not improve NRA significantly, but it can also reduce nitrate content. Both of Mo and Mn could improve the content of soluble sugar and Vc but Mn improved them more significantly compare with Mo. Mo cooperated with Mn had improved NRA and reduced nitrate content the most significant, in addition, it also improved the content of soluble sugar and Vc the most significant. It is the most effective if Mo and Mn fertilized both in leaves and root, and fertilized in root is better than in leaves.
小白菜品种间的硝酸盐含量和硝酸还原酶活性差异显著,本研究主要采用“上海青、苏州青、兴化青梗菜、引02、青丰一号、兔子腿矮脚黄”的高代自交系为试验材料,利用水溶液培养法,围绕NRA对小白菜硝酸盐积累及生理机制影响进行了研究,
结果表明:
1.氮素形态对小白菜体内硝酸盐积累和NRA的影响
在全硝态氮供应的N1处理组的硝酸盐含量显著高于硝/铵等比处理的N2组和全铵态氮处理的N3组;N1组的NRA略高于N2组,并无显著性差异,但N1组和N2组的NRA均极显著高于N3组,全铵态氮的N3处理组硝酸盐含量和NRA都最低。N2处理组的品质和产量最高,N1略低于N2处理组,而N3的品质和产量最低。在全硝态氮供应下,根系对NO_3~-的吸收速率最高,而随着铵态氮比例的增加,其吸收速率随之降低。
2.根部NRA对小白菜硝酸盐和可溶性糖代谢的影响
根系NRA受抑的情况下根系对NO_3~-吸收速率显著下降,而地上部的硝酸盐含量并无显著差异。同时,根系NRA受抑条件下,其根系可溶性糖浓度增加,而地上部可溶性糖浓度则相对减少。
3.小白菜光合性能与NRA关系
1)光照强度是制约NRA和硝酸盐含量的主要因素,尤其对NRA的高低影响极显著。2)NRA的降低会对光合速率产生一定的反作用。而叶绿素含量、气孔导度、胞间CO_2浓度和蒸腾速率在NRA抑制前后无显著变化,不受NRA的直接影响。3)在一定的硝态氮浓度范围内(=8mmol·L~(1)),NRA的升高对净光合速率的升高有促进作用。以上结论表明,光合性能与NRA存在着一定的相互促进和相互制约的关系。
4.钼锰不同施用方式对小白菜NRA和硝酸盐含量的影响
钼显著提高了小白菜的NRA,使NO_3~-显著降低,锰的施用,对NRA的影响不明显,但可以使硝酸盐含量降低,钼和锰的施用均可提高可溶性糖含量和Vc含量,与钼相比,锰对可溶性糖含量和Vc含量的提高更为显著。钼锰配施对提高NRA和降低硝酸盐含量最为明显,同时配施时对可溶性糖含量和Vc含量的提高也最为明显。叶面喷施与营养液根部结合施用对各生理指标的影响效果要明显高于叶面施用而略高于营养液施用。
Concentrations of nitrate in vegetables have been the focus of attention in several countries. A lot of reaserch on environment hygienics show that nitrate may reduce to nitrite which can lead to some diease such as cancer in animals. The nitrate absorbed by human is 81.2% derived from vegetable. Furthermore, nitrate was very easy accumulated in pakchoi which is a very popular vegetable which occupied 33% of the whole vegetable market in China, and NRA (Nitrate Reductase Activity) was a main limit factor in nitrate reduced.
In this study soluble culture was used for physiological mechanism of nitrate accumulation and NRA in pakchoi. There was significant difference of nitrate content and NRA between different genotypes of vegetable crops. Six genotypes were used in this study: Shanghaiqing, Suzhouqing, Xinghua qinggengcai, Yin02,QingfengNo.l,Tuzitui aijiaohuang. The results went as followed:
1 Effects of nitrogen forms on nitrate content and nitrate reductase of pak-choi
The content of nitrate in the N1 system which was offered pure NO_3~- is obviously higher than the system of N2 (NO_3~-/NH_4~+=1/1) and N3(NO_3~-/NH_4~+=0/1).NRA in N1 has a little higher than N2, but NRA in N1 and N2 is much obviously higher than N3.The nitrate content and NRA is the lowest in N3 which is treat with pure ammonium. Qualities and yield of N2 are the highest, N1 is a little lower than N2, N3 is the lowest neither in qualities or yield. The absorb rate of NO_3~- in root is the fastest when pure nitrate was offered, but the absorb rate was decreasing with the ammonium increasing.
2 Effect of NRA in root on the absorb and metabolism in nitrate of pakchoi
The absorb rate of NO_3~- reduced by inhibiting NRA in root, but there is no significant dramatically difference of the nitrate content. Furthermore, if we inhibit NRA in root,the concentration of soluble sugar increased and the concentration reduced in shoot.
3 Studies on the relationship of NRA and photosynthetic characters of pakchoi
The main characters of NRA and nitrate content controlling factor are light intensity especially on NRA. In addition, NRA reduced can feed back the photosynthetic, but chlorophyll content, Cond and Ci did not change obviously after NRA reduced. What's more, the grown of NRA can promote the photosynthetic when the nitrate content is lower than 8mmol·L~(-1).To sum up,the conclusion shows that photosynthetic characters and NRA have some relationship both in promotion and restriction.
4 Effects of Molybdenum and Manganese on NRA and nitrate content
Mo was significantly improved NRA and reduced nitrate in two genotype of pakchoi. However, Mn could not improve NRA significantly, but it can also reduce nitrate content. Both of Mo and Mn could improve the content of soluble sugar and Vc but Mn improved them more significantly compare with Mo. Mo cooperated with Mn had improved NRA and reduced nitrate content the most significant, in addition, it also improved the content of soluble sugar and Vc the most significant. It is the most effective if Mo and Mn fertilized both in leaves and root, and fertilized in root is better than in leaves.
引文
[1] Fytianos K,Zarogiannis P. Nitrate and nitrite accumulation in fresh vegetables from Greece.Bull Environ Contam Toxicol, 1999.62:187-192.
[2] 陈君石，闻芝梅主译．食物、营养与癌症预防．上海：上海医科大学出版社，1999．
[3] Fine, D H, N-nitrate compounds: Occurrence and biological effects (IARC Sci. Publ.No.41),Iyon,France, IARC,1982,379.
[4] 任敬红．国外医学护理分册[M]．2002，21(4)：187-188．
[5] Margaretha B Z, Nitrate accumulation in vegetables and its relationship to quality. Association of Applied Biologists, 1989,115:553-561.
[6] White J W J. Relative significance of daietarg source of nitrate and nitrite.J Agric Food Chem.l975,23(5):886-891.
[7] 周燮，沈康．植物营养生理进展．1985．《南京农业大学科技译丛》，第93页．
[8] Sodium Nitrite Summary of Evaluations Performed by the Joint FAO/WHO Expert Committee on food Additives (JECFA) Report: TRS 859-JECFA44/29,31.995.
[9] 黄勇强，王利群，董英．蔬菜硝酸盐含量测定与食用安全性探讨．现代科学仪器，2003，(1)：56-58．
[10] 冀宏杰．过量施用氮肥对北京市蔬菜硝酸盐含量影响的综合评估．[硕士学位论文]．保定：河北农业大学，1999．
[11] Miller A J, Smith S J. Nitrate transport and compartmentation in cereal root cells[J]. J. Exp.Bot..1996, 47: 843-854.
[12] Crawford N M, Glass ADM.Molecular and physiological aspects of nitrate uptake in plant (Reviews) [J].Trends in Plant Science,1998,3 (10):389-395.
[13] 阮云泽，唐树梅．蔬菜累积硝酸盐的机理及其调控措施研究进展[J]．华南热带农业大学学报，2003，9(4)：14-18．
[14] 高祖明．几种叶菜的硝酸盐和亚硝酸盐积累及其与有关酶活性的关系[J]．植物生理通迅， 1990，(3)：21-24．
[15] Olday,EC.et al.A Physiological basis for different patterns of nitrate accumulation in two spinach cultivars. J. Amer.Soc.Hort.Sci.1976,101(3):217-219.
[16] 庄舜尧，孙秀廷．氮肥对蔬菜硝酸盐积累的影响[J]．土壤学进展，1995，23(3)：29-35．
[17] Marchner H. M. Nutrition of Higher Plants. Harcourt Brace Jovanovich Martin,1986,200-210.
[18] 沈明珠．蔬菜硝酸盐累积的研究．园艺学报，1982，9(4)：41-48．
[19] Cantliffe DI.Nitrate accumulation in spinach grown at different temperatures[J].J.Am.Soc.Hort.Sci.,1972,97(5):674-676.
[20] Reinink K,Eenink A H.Genotypical differences in nitrate accumulation in vegetables and its relationship to quality[J].Annals of Applied Biology,1989,115(3):553-561.
[21] Olday,EC.et al.,1978,J.Amer.Soc.Hort.Sci.99(2):135-138.
[22] 尹元萍．科学施肥控制蔬菜硝酸盐[J]．西南园艺，2002，30(1)：25-26．
[23] 王明霞，王三根，吴文彬等．硝态氮对不同基因型小白菜光合速率及品质的影响[J]．西南农业大学学报，2005，27(3)：386-389．
[24] 陈振德．不同收获时期对蔬菜硝酸盐含量的影响[J]．中国蔬菜，1989，(3)：8-10．
[25] 王朝辉，李生秀，田霄鸿．不同氮肥用量对蔬菜硝态氮累积的影响[J]．植物营养与肥料学报，1998，4(1)：22-28．
[26] 中国农业百科全书[M]．农业化学卷，36-37．
[27] 刘永菊，曹一平，夏江等．不同NPK配比对大白菜产量及硝酸盐累积的影响[J]．土壤肥料，1999，(4)：26-29．
[28] 马娟娟，胡全才，王申贵．钾、锌肥与氮肥配施对油菜体内硝酸盐累积的影响[J]．山西农业科学，2000，28(2)：49-52．
[29] 倪吾钟，何念祖，林荣新．钾对大白菜的营养作用及其生理机制研究[J].;植物营养与肥料学报，1997，3(2)：117-121．
[30] 孙权，高艳明，李建设．氮、磷、钾配合施用对油菜硝酸盐含量的影响[J]．中国生态农业学报，2003，11(3)：84-86．
[31] 林志刚，赵义华，薛耀英．叶菜类蔬菜的硝酸盐积累规律及其控制方法的研究[J]．土壤通报，1993(24)：253-255.
[32] 周根娣等．环境因素对蔬菜累积硝酸盐影响的研究进展[J]．土壤通报，2004，(03)：123-126．
[33] Cantliffe DI. Nitrate accumulation in spinach grown at different temperatures [J].J.Am.Soc.Hort.Sci.,1972,97(5):674-676.
[34] 黄建国，袁玲．重庆市蔬菜硝酸盐、亚硝酸盐含量及其与环境的关系[J]．生态学报，1996，16(4)：383-388．
[35] 周艺敏．氮素化肥对蔬菜硝酸盐积累的影响[J]．华北农学报，1986，4(1)：110-115．
[36] Van der Boon J. Steehuizen J W. Steingraever EG Growth and nitrate concentration of lettuce as affected by total nitrogen and chloride concentration NH_4~+-N/NO_3~桸 ratio and temperature of the recirculating nitrate solution. J Hort Sci. 1990.63(3):309.
[37] 朱祝军，蒋有条．不同形态氮素对不结球白菜生长和硝酸盐积累的影响[J]．植物生理学通迅，1994，30(3)：198-201．
[38] 高祖明，张耀栋，张道勇等．NPK对叶菜类硝酸盐积累和硝酸还原酶、过氧化物酶的影响[J]．园艺学报，1989，16(4)：293-297．
[39] 常丽新．钾肥在小白菜和萝卜上的施用效果[J]．中国蔬菜，2002，(1)：16-17．
[40] 胡勤海，叶兆杰，马生良．杭州市蔬菜硝酸盐污染现状及防治对策[J]．环境污染与防治，1991，13(4)：5-9．
[41] 黎星辉，黄启为，彭建伟．施肥对蔬菜硝酸盐含量的影响，菜园土壤肥力与蔬菜合理施肥[M]．南京：河海大学出版社，1997，202-206．
[42] 王正银，刘金萍，涂从等．几种化学物质用量对叶类蔬菜硝酸盐和营养品质的影响[J]．土壤农化通报，1998，13(2)：16-20．
[43] 黄启为，彭建伟．化肥对蔬菜硝酸盐含量的影响[J]．湖南农业大学学报(自然科学版)，2002，28(5)：387-390．
[44] 李冰，王昌全，冯长春等．喷施不同钙肥对莴笋产量及品质的影响[J]。中国农学通报，2004，20(2)：129-131．
[45] 余光辉，张杨珠，万大娟．几种硝化抑制剂对土壤和小白菜硝酸盐含量及产量的影响[J]．应用生态学报，2006，17(2)：247-250．
[46] 殷允相，杨俊，林孔仪等．银川地区蔬菜硝酸盐类含量、污染评价及防治途径研究[J]．宁夏农林科技，1993，(1)：40-43．
[47] 熊艳，尹增松等．蔬菜中硝酸盐污染现状及其防治措施[J]．云南农业大学学报，2003，18(3)：31-35．
[48] 汪李平，向长萍，王运华．白菜不同基因型硝酸盐含量差异的研究[J].园艺学报，2004，31(1)：43-46．
[49] Rein ink K Genetics of nitrate content of lettuce I. Analysis of generation means[J]. Euphytica,1991, 54: 83-92.
[50] 林家宝，陈火英，林观捷．蔬菜硝酸含量遗传的初步研究[J]．上海农学院学报，1995(2)：98-102．
[1].EvansH J,Nason A.,1952, Arch.Biochem.BioPhys.39:234-235.
[2]Evans H J,NasonA.,1953, Plant Physiol. 28:233-254.
[3] Tang P S,WuH.Y.,1957, Nature, 179:1355-1356.
[4]Kessler E,1964, Ann. Rev. Plant Physiol. 15:57-72.
[5]BeeversL,HagemanR. H., 1969, Ann. Rev. Plant Physiol. 20:495-522.
[6] Simon B C,Christopher E C. Characterization of the nitrite reductase gene Nii and the nitrate-assimilation gene cluster of Stagonospora (Septoria) nodorum[J].Curr Genet,1999,36:282-289.
[7] Samia D,Isabelle Q,Marie-Therese L, et al. Expression of a deregulated tobacco nitrate reductase gene in potato increases biomass production and decreases nitrate concentration in all organs. [J].Planta, 2004, 219:884-893.
[8] Cheneby D,Hallet S, Mondon M,et al.Genetic characterization of the nitrate reducing community base on G nucleotide sequence analysis [J].Microbial Ecology, 2003, 46(1):113-121.
[9]Antipov AN,Sorokin D Y. New enzyme belonging to family of molybdenum-free nitrate reductase[J].Biochemical journal, 2003, 369(1):185-189.
[10]Cheng C L,Dewdney J,Kleinhofs A,er al.A new locus (Nit2) nitrogene regulatory Protein: importance of the number, spacing and orientation of GATA core elements and their flanking sequences upon NIT2 binding [J].Biochemistry, 1988, 33:576-582.
[11] Crawford N M,Campbell W H, Davis R. Nitrate reductase from squash: cDNA clining and nitrate regulation [J].Proc. Natl.Acad. Sci. USA, 1986, 83:8073-8076.
[12]Wilkinson J Q,Crawford N M. identification of the Arabisposis CHL3 gene as the nitrate reductase structural gene NIA2[J]. Plant Cell,1991,3:461-471.
[13] Choi H, Kieinhofs A, An G Nucleotide sequence of rice nitrate reductase genes[J]. Plant Mol.Biol,1989,13:731-733.
[14] Daniel-Vedele F, Dorbe M F, Caboche M, et al. Cloning and analysis of the nitrate erduetase gene form tomato: A comparison of nitrate reductase protein sequences in higher Plants[J].Plants.Genet,1989, 220:88-94.
[15] 王利群，王勇，董英，等．硝酸盐对硝酸还原酶活性的诱导及硝酸还原酶基因的克隆[J]．生物工程学报，2003，19(5)：632-635．
[16] Cherel I,Gonneau M, Meyer C, et al. Biochemical and immunological characterization of nitrate reductase-deficient nia mutants of Nicotiana Plumbaginifolia [J].Plant Physiol.1990,92:659-665.
[17] Kubo Y,Ogura N,Nakagawa H. Limites Protelysis of the nitrate reductase from spinach leave[J].Biol Chem.,1998, 263:19684-19689.
[18] Masatoshi S,Hiroakil,Shinya N,et al. Cloning of a nitrate reductase in activateor (NRI) cDNA form spinacia oleracea L. and expression of mRNA and protein of NRI in cultured spinach cells[J]. Planta, 2003, 216:961-968.
[19] Hoff T, Stummann B M, Henningsssen K W, et al. Cloning and expression of a gene encoding a root specific nitrate reductase in bean[J]. Physiol. Plant,1991,82:197-204.
[20] Jensen P E, Moller M Q, Stummann B M, et al. Identification and characterization of a nitrate reductase gene form bean containing four introns[J]. Physiol. Plant, 1994, 92:613-623.
[21] Miyazkai J, Juricek M, Angelis K, et al.Characterization and sequence of a novel nitrate reductase form barley[J]. Mol.Gen.Genet.,1991, 228:329-334.
[22] Schnorr K M, Juricek M, Huang C, et al Analysis of barley nitrate reductase cDNA and genomic coones[J]. Mol. Genet., 1991, 227:411-416.
[23] 孙淑斌，罗金葵，徐国华，等．小白菜硝酸还原酶基因的克隆与初步鉴定[J] 植物营养与肥料学报2006，12(4)：592-596．
[24] Perng-Kuang Chang, Kenneth C. Ehrlich, John E.Linz, Deepak Bhatnagar, et al.Characterization of the Aspergillus Parasiticus niaD and niiA gene cluster[J]. Curr Genet, 1996, 30:68-75.
[25] Nicholas D. J. D., 1961,Ann.Bev.Plant Physiol.12:63-90.
[26]Guerrero, M.G,Vega, J. M., Losada,M.,The assimilatory nitrate-reduction and its regulation.Ann. Rev. Plant Physiol.,1980,32:169-402.
[27] Hewitt E. J., Ann.Rev. Plant Physiol. 1975, 26:73-158.
[28]Somer D.A.,et al.Plant Physiol,1983, 72:949-952.
[29]Funkhouser E,A.,et al,1983, Plant Physiol. 65:939-943.
[30]Funkhouser E,A.,Ramadoss C.S.,1980, Plant Physiol.65:944-948.
[31] Loppes R, Radnox M. Two short regions of the Promoter are essential of activation and repression of the nitrate reductase gene in Chlamydomonas reinhardtii [J]. Molecular Genetic and Genomics,2002, 268(1): 42-48.
[32] Melzer M J, Edwards R A. Nitrogen control in Bacteria[J]. Microbiol. Rev,1995,59:604-622.
[33] Callaci J J, Smarrelli J Jr. Regulation of the inducible nitrate reductase isoform from soybeans[J].Biochim.Biophys.Acta, 1991, 1088:127-130.
[34] Crawford N M. Nitrate: Nutrient and signal for Plant growth[J]. Plant Cell, 1995, 7: 859-868.
[35] Srivastava H S. Regulation of nitrate reductase activity in higher Plants[J]. Phytochemistry, 1980,19:725-733.
[36] Frost W B. et al. Plant Physiol, 1978, 61:323-331.
[37] Remmol/Ller J L,Campbell. Plant Physiol. 1986, 80:442.
[38] Calza R.Mol Gen Genet.,1987, 209:552.
[39] Campbell W H. Nitrate reductase and its role in nitrate assimilation in Plants[J]. Physiol Plant,1988, 74:214-219.
[40] Wallaee W,O aks A. In Dalling M J(ed), Plant Protaolytic Enzymes(Vol.2). CRC Press, Boca Raton, FL.1986, p.81.
[41]SorgerGJ.et al.Biochem.J, 1974, 140:395.
[42] Gerendas J, Zhu Z, Bendixen R, Ratcliffe R G. Sattelmacher B. Physiological and biochemical Processes related to ammol/Lonium toxicity in higher plants. Zeitschrift fur Pflanzenernahrung and Bodenkunde.1997, 160:239-251.
[43] Raab T K,Terry N. Nitrogen source regulation of growth and Photosynthesis in Beta vulgaris L.Plant Physiology. 1994, 105: 1159-1166.
[44].Magalhaes JR, Huber DW. Ammol/Lonium assimilation in different plants species as affected by nitrogen from and pH control in solution culture. Fertilizer Research, 1989, 21:1-6.
[45].Mengel K,Viro M. The significance of plant energy status for the uptake and incorporation of NH_4~+ - nitrogen by young rice plants. Soil Sci. Plant Nutr. 1978, 24: 407-416.
[46] 封克，汪晓丽，陈平等．水稻苗期不同时段NO3~-吸收特点及其受NH_4~+的影响[J]．中国农业科
?学，2003，36(3)：307-312．
[47] 封克，汤炎，张素玲．钱离子对不同基因型水稻吸收硝酸根离子的影响[J]．植物生理学通讯，2001，37(3)：192-194．
[48] Minotti P L, Williams D C, Jackson W A. The influence of ammol/Lonium on nitrate reduction in wheat seedlings. Planta, 1969, 86:267-271.
[49] Kronzuker H J, Glass AD M,Siddiqi M Y. Inhibition of nitrate uptake by ammol/Lonium in barley.Analysis of component fluxes.Plant Physiology,1999,120:283-291.
[50] Lycklama J. The absorption of ammol/Lonium and nitrate by Perennial ryegrass. Acta. Bot. Neer.1963, 12:361-462.
[51] Rao K P, Rains D W. Nitrate absorption by barley:Ⅱ.Influence of nitrate reductase activity. Plant Physiology, 1976, 57:59-62.
[52] Samuelson M E, Ohlen L, Lind M, Larsson C M. Nitrate regulation of nitrate uptake and nitrate reductase expression in barley grown at different nitrate: ammol/Lonium ratios at constant relative nitrogen addition rate. Physiologia Plantarum, 1995, 94:254-260.
[53] Youngdahl L J, Pacheco R, Street J J, Vlek P L G The kinetics of ammol/Lonium and nitrate uptake by young rice plants. Plant And Soil,1982,69:225-232.
[54] 廖祥儒，朱新产．氨积累对植物生长发育的效应[J]．农业环境保护，1996，15(6)：281-287．
[55] 何新华，Ann Oak S，李明启．C3和C4植物的氮素利用效率[J]．云南植物研究，1994，16(1)：93-94．
[56] 高廷东．光和不同形态氮互作对小麦幼苗碳氮同化的影响．山东农业大学硕士学位论文2003．
[57] Lillo C,Ruoff P. An unusually rapid light-induced nitrate reductase mRNA pulse and circadian osicillation. Naturuis-senschaften, 1989, 76:526-532.
[58] 林振武．硝酸还原酶的研究动态[J]．植物生理学通讯，1987，(6)：8-13．
[59]Deng M. D., Moureaux T., Cherel I. et al. Effects of nitrogen metabolites on the regulation and circadian expression of tobacco NR. Plant Physiol Biochem.,1991, 29:239-247.
[60] 余让才，李明启，范燕平．高等植物硝酸还原酶的光调控[J]．植物生理学通讯，1997，33(1)：61-65．
[61] Deng M, Moureaux T., Caboche M, et al. A molybdate analog in activating nitrate reductase,deregulates the expression of the nitrate reductase structure gene[J]. Plant Physiol.1989, 91:304-309.
[62] Kaiser W M, Spill D, Brendle-Behnisch E. Adenine nucleotide are apparently involves in the light-dark modulation of spinach-leaf nitrate reductase. Planta,1992,186:236-240.
[63] Kaiser W M, Huber S C. Posttranslational regulation of nitrate reductase in higher Plants [J]. Plant Physiol., 1994, 106:817-821.
[64] Douglas P, Morrice N, Mackintosh C. Identification of a regulatory Phosphorylation site in the hinge region of nitrate reductase from spinach leaves [J]. EFBS Lett. 1995,377:113-117.
[65]Bachmann M, Hunter H L,Liao P C. et al.The inhibitor protein of Phosphorylated nitrate reductase from spinach leaves is al4-3-3 protein[J]. EFBS Lett, 1996. 387:127-131.
[66]Chantarotwong W., Huffaker, RC,Miller,B.L., Granstedt,R.C.,In vivo nitrate reduction in relation to nitrate uptake,nitrate content, and in vivo nitrate reductase activity in intact barley seedlings[J]. Plant Physiol., 1976, 57:519-522.
[67] Mackown C T, Jackson W A, Volk R G Partitioning of Previously accumulated nitrate to translocation reduction and efflux in corn roots[J]. Planta 1983, 157:8-14.
[68] Udayakumar M, Devendra R, Reddy V S,Sastry K S K.Nitrate availability under low irradiance and its effect on the nitrate reductase activity. New Phytol, 1981, 88:290-297.
[69] Pearson J,Steer B T. Planta, 1977. 137:107.
[70] Taluoizte A, Champigny M L. Plant Cell Environ, 1988, 11:149.
[71] Talouize A, Guiraud G, Moyse A, Marol C M, Chmapigny M L,Effect of previous nitrate deprivation on ~(15)N-nitrate absorption and assimilation by wheat seedlings[J]. Plant Physiol, 1984,116:113-122.
[72].Passama L.et al.Plant Soil, 1987, 102:145.
[73] Reed A J,Hageman RH. Plant Physiol, 1980, 66:1184.
[74] 许长蔼，倪晋山．小麦叶内硝酸还原的代谢库[J]．植物生理学报，1990，16(3)：277-283．
[75] Shaner D L, Boyer J S. Nitrate reductase activity in maize leaves regulation by nitrate flux. Plant Physiol, 1976, 58:499-504.
[76] Martinoia E, Heck U, Wiemker A.Vacuoles as storage compartments for nitrate in barley leaves.Nature, 1981,289:292-293.
[77] 许长蔼，倪晋山．NO_3~-上运和液泡内NO_3~-外流对小麦叶内硝酸还原酶活性的调节[J]．植物生理学报，1990，16(4)：340-346．
[78] Yamazaki M. Plant Cell Physiol, 1986, 27:443.
[79] Amy N K,Carrett R H. Purification and characterization of the nitrate reductase from the diatom Thalassiosira pseudonana[J]. Plant Physiol, 1974, 54: 629-637.
[1] Britto D T,Kronzucker H J.NH_4~+ toxicity in higher plants:A critical review [J].J.Plant Physiol.,2002,159:567-584.
[2] 王利群，董英，黄达明等．蔬菜硝酸盐的积累及其生理机制研究进展[J]．江苏农业科学,2006，(6)：78-811。
[3] 林观捷，林家宝，陈火英．影响蔬菜硝酸盐含量积累因素的探讨[J]．上海农学院学报，1995，13(1)：47-52．
[4] Smiciklas KD,Below F E. Role of nitrogen form in determining yield of field-grown maize [J].Crop Science , 1992 , 32 : 1220-1225.
[5] 张春兰，高祖明，张耀栋等．氮素形态和NO_3~-N:与NH_4~+-N配比对菠菜生长和品质的影响[J]．南京农业大学学报，1990，13(3)：70-74．
[6] 陈巍，罗金葵，姜慧梅等．不同形态氮素比例对不同小白菜品种生物量和硝酸盐含量的影响[J]．土壤学报，2004，41(3)：420-425．
[7] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社．
[8] 封克，汪晓丽，陈平等．水稻苗期不同时段NO_3~-吸收特点及其受NH~+的影响[J]．中国农业科学，2003，36(3)：307-312．
[9] Guerrero, M G, Vega, J M., Losada,M.,The assimilatory nitrate-reduction and its regulation.Ann.Rev. Plant Physiol.,1980,32:369-402.
[10] Cokojiob O A．氮肥施用的生态观点[J]．土壤学进展，1992(1)：19-21．
[11] Granstedt R C.Huffaker R C. Identification of the leaf vacuole as a major nitrate storage pool. Plant Physiol.l982,70:410-413.
[12] Raab T K,Terry N.Nitrogen source regulation of growth and Photosynthesis in Beta vulgaris L.Plant Physiology. 1994,105:1159-1166.
[13] Gerendas J, Zhu Z. BendixenR.RatcliffeRG. Sattelmacher B. Physiological and biochemical Processes related to ammol/Lonium toxicity in higher plants. Zeitschrift fur Pflanzenernahrung and Bodenkunde.1997,160:239-251.
[14] 孙传范，戴廷波，曹卫星．不同施氮水平下增铵营养对小麦生长和氮素利用的影响[J]．植物营养与肥料学报，2003，9(1)：33-38。
[15] Kronzuker H J,Glass A D M,Siddiqi M Y. Inhibition of nitrate uptake by ammol/Lonium in barley.Analysis of component fluxes. Plant Physiology, 1999, 120:283-291.
[16]Samuelson M E, Ohlen L, Lind M, Larsson C M. Nitrate regulation of nitrate uptake and nitrate reductase expression in barley grown at different nitrate: ammol/Lonium ratios at constant relative nitrogen addition rate. Physiologia Plantarum,1995,94:254-260.]
[17] 董晓英，李式军，沈仁芳．白菜不同品种对硝酸盐吸收积累差异原因初探[J] 园艺学报，2003，30(4)：470-472．
[1] Fytianos K,Zarogiannis P. Nitrate and nitrite accumulation in fresh vegetables from Greece.Bull Environ Contain Toxicol, 1999,62:187-192.
[2] 陈君石、闻芝梅主译．食物、营养与癌症预防．上海：上海医科大学出版社，1999．
[3] White J W J. Relative significance of daietarg source of nitrate and nitrite.[J]. Agric Food Chem.l975,23(5):886-891.
[4] Evans H J, Nason A. Pyridine nucleotide nitrate reductase from extracts of higher plants[J]. Plant Physiol,1953, 28:233-254.
[5] 郭世荣 无土栽培学[M]．北京：中国农业出版社，2004．125-129．
[6] Deng M,Moureaux T, Caboche M. Tungstate.A molybdate analog inactivating nitrate reductase,deregulates the expression of the nitrate reductase structural gene.[J]. Plant Physiol,1989,91:304-309.
[7] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．123-128．
[8] Hageman R H, D P Hucklesby.Methods in Enzymology[J].1971,23 (A): 491-503.
[9] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．195-197．
[10] 林立欣，李寅安，王美菊．萝卜杂种优势与配合力的研究[J]．中国农业科学，1986年(2)：54-60
[11] 司江英，汪晓丽，陈平等．硝酸还原酶抑制剂和NH_4~+对不同基因型水稻苗期NO_3~-吸收的影响[J]．扬州大学学报(农业与生命科学版)．2004，25(1)：59-62．
[12] Robert H. Desirée Gómez F. Tobacco plants that lack expression of functional nitrate reductase in roots shows changes in growth rates and metabolite accumulation.[J].Journal of Experimental Botany,2001,52,No.359:1251-1258.
[1] 林振武．硝酸还原酶的研究动态[J]．植物生理学通讯，1987，(6)：8-13．
[3] 刘丽，甘志军，王宪泽．植物氮代谢硝酸还原酶水平调控机制的研究进展[J]．西北植物学报，2004，24(7)：1355-1361．
[2] Magalhase J R. The Responses of Ammonium assimilation enzymes to nitrogen form in different Plant[J].Journal of Plant Nutrition,1991,14(2):175-185.
[4] Deng M, Moureaux T, Caboche M. Tungstate, a molybdate analog inactivating nitrate reductase,deregulates the expression of the nitrate reductase structural gene[J]. Plant Physiol, 1989,91:304-309.
[5] 李合生主编．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．123-128．
[6] Mohotti A J,Lawlor D W. Diurnal variation of Photo synthesis and Photo-inhibitionin tea:effects of irradianee and nitrogen supply during growth in the field[J].Journal of ExperimentalBotany,2002,53(367):313-322.
[7] Huber J L,Huber S C,Campelli W H. Reversible light/dark modulation of spinach leaf nitrate reductase activity involved protein phosphorylation[J].Arch Biochem Biophys,1992, 296(1):58-65.
[8] Champigny M L. Integration of photosynthetic carbon and nitrogen metabolism in higher plants[J].Photosynthesis Research,1995, 46:117-127.
[9] Zavala J A, Ravetta A D.Allocation of Photoassimilates to biomass,resin and carbohydrates in Grindelia chiloensis as affected by light intensity[J].Field crops research,2001,69:143-149.
[10] 关义新，林葆，凌碧莹．光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J]．作物学报，2000，26(6)：806-812．
[11] 朱为民．蔬菜硝酸盐积累的基因型差异与营养调控及分子生物学的研究[D]．南京：南京农业大学，1997．
[1] Kaiser B N. The role of molybdenum in agricultural plant production. Annals of Botany , 2005 , 96(5):745-754.
[2] 庞静，胡承孝，王运华等．钼对黄棕壤上冬小麦碳代谢的影响．华中农业大学学报，2001，20(1)：33-35．
[3] Fang Z,Zhang Y G,Zhang F S,et al.Mechanisms of difference in Mo efficiency between wheat and oil seed rape. Pedosphere[J] 2000,10(3) :213-220.
[4] 喻敏，王运华．高等植物中的钼．植物学通报，1999，16(5)：547-554．
[5] 杜新民．锰对小白菜产量和品质的影响[J]．2007，27(5)：48-51．
[6] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．248-249．
[7] Hageman R H,Hucklesby D P.Methods in Enzymology[J].1971,23(A): 491-503.
[8] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．195-197．
[9] 王立克，洪法水．钼浸种对苜蓿硝酸还原酶活性及营养成分的影响[J]．中国草地．1995，15(3)：37-38．
[10] 杜应琼，廖新荣，黄志尧等．硼、钼对花生氮代谢的影响[J]．作物学报，2001，27(5)：612-616．
[11] 付其如，何纪荣．钼对葡萄叶片硝酸还原酶活性及生长发育的影响．四川师范学院学报，1996,17(1)：56-60．
[12] 张录强．钼(Mo)的生理及其生物学效应[J]．生物学通报，1999,34(11)：24-26．
[13] Chitralekha C, Ninnala N. Sugarcane yield and physiology in response to molybdenum supply.Sugar Cane, 1999,(3):19-20.
[14] 宋志荣．施锰对马铃薯产量和品质的影响[J]．中国农学通报，2005，21(3)：222-223．
[15] 杨保川，王昌全，项虹艳等．叶面喷施钼锰对莴笋生长发育和硝酸盐含量的影响[J]．农业环境科学学报，2006，25(增刊)：96-99．
[16] 杜慧玲，王建锁，姚永平．钾锌锰配施对甜椒品质的影响[J]．中国农学通报，2000，16(5)：32-33．
[17] 刘鹏，徐根娣，周子仙等．Mo和Mn对七子花几种生理效应的影响[J]．林业科学研究，2002，15(2)：156-162．
[18] 关义新，林葆，凌碧莹．光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J]．作物学报，2000，26(6)：806-812．
[2] 陈君石，闻芝梅主译．食物、营养与癌症预防．上海：上海医科大学出版社，1999．
[3] Fine, D H, N-nitrate compounds: Occurrence and biological effects (IARC Sci. Publ.No.41),Iyon,France, IARC,1982,379.
[4] 任敬红．国外医学护理分册[M]．2002，21(4)：187-188．
[5] Margaretha B Z, Nitrate accumulation in vegetables and its relationship to quality. Association of Applied Biologists, 1989,115:553-561.
[6] White J W J. Relative significance of daietarg source of nitrate and nitrite.J Agric Food Chem.l975,23(5):886-891.
[7] 周燮，沈康．植物营养生理进展．1985．《南京农业大学科技译丛》，第93页．
[8] Sodium Nitrite Summary of Evaluations Performed by the Joint FAO/WHO Expert Committee on food Additives (JECFA) Report: TRS 859-JECFA44/29,31.995.
[9] 黄勇强，王利群，董英．蔬菜硝酸盐含量测定与食用安全性探讨．现代科学仪器，2003，(1)：56-58．
[10] 冀宏杰．过量施用氮肥对北京市蔬菜硝酸盐含量影响的综合评估．[硕士学位论文]．保定：河北农业大学，1999．
[11] Miller A J, Smith S J. Nitrate transport and compartmentation in cereal root cells[J]. J. Exp.Bot..1996, 47: 843-854.
[12] Crawford N M, Glass ADM.Molecular and physiological aspects of nitrate uptake in plant (Reviews) [J].Trends in Plant Science,1998,3 (10):389-395.
[13] 阮云泽，唐树梅．蔬菜累积硝酸盐的机理及其调控措施研究进展[J]．华南热带农业大学学报，2003，9(4)：14-18．
[14] 高祖明．几种叶菜的硝酸盐和亚硝酸盐积累及其与有关酶活性的关系[J]．植物生理通迅， 1990，(3)：21-24．
[15] Olday,EC.et al.A Physiological basis for different patterns of nitrate accumulation in two spinach cultivars. J. Amer.Soc.Hort.Sci.1976,101(3):217-219.
[16] 庄舜尧，孙秀廷．氮肥对蔬菜硝酸盐积累的影响[J]．土壤学进展，1995，23(3)：29-35．
[17] Marchner H. M. Nutrition of Higher Plants. Harcourt Brace Jovanovich Martin,1986,200-210.
[18] 沈明珠．蔬菜硝酸盐累积的研究．园艺学报，1982，9(4)：41-48．
[19] Cantliffe DI.Nitrate accumulation in spinach grown at different temperatures[J].J.Am.Soc.Hort.Sci.,1972,97(5):674-676.
[20] Reinink K,Eenink A H.Genotypical differences in nitrate accumulation in vegetables and its relationship to quality[J].Annals of Applied Biology,1989,115(3):553-561.
[21] Olday,EC.et al.,1978,J.Amer.Soc.Hort.Sci.99(2):135-138.
[22] 尹元萍．科学施肥控制蔬菜硝酸盐[J]．西南园艺，2002，30(1)：25-26．
[23] 王明霞，王三根，吴文彬等．硝态氮对不同基因型小白菜光合速率及品质的影响[J]．西南农业大学学报，2005，27(3)：386-389．
[24] 陈振德．不同收获时期对蔬菜硝酸盐含量的影响[J]．中国蔬菜，1989，(3)：8-10．
[25] 王朝辉，李生秀，田霄鸿．不同氮肥用量对蔬菜硝态氮累积的影响[J]．植物营养与肥料学报，1998，4(1)：22-28．
[26] 中国农业百科全书[M]．农业化学卷，36-37．
[27] 刘永菊，曹一平，夏江等．不同NPK配比对大白菜产量及硝酸盐累积的影响[J]．土壤肥料，1999，(4)：26-29．
[28] 马娟娟，胡全才，王申贵．钾、锌肥与氮肥配施对油菜体内硝酸盐累积的影响[J]．山西农业科学，2000，28(2)：49-52．
[29] 倪吾钟，何念祖，林荣新．钾对大白菜的营养作用及其生理机制研究[J].;植物营养与肥料学报，1997，3(2)：117-121．
[30] 孙权，高艳明，李建设．氮、磷、钾配合施用对油菜硝酸盐含量的影响[J]．中国生态农业学报，2003，11(3)：84-86．
[31] 林志刚，赵义华，薛耀英．叶菜类蔬菜的硝酸盐积累规律及其控制方法的研究[J]．土壤通报，1993(24)：253-255.
[32] 周根娣等．环境因素对蔬菜累积硝酸盐影响的研究进展[J]．土壤通报，2004，(03)：123-126．
[33] Cantliffe DI. Nitrate accumulation in spinach grown at different temperatures [J].J.Am.Soc.Hort.Sci.,1972,97(5):674-676.
[34] 黄建国，袁玲．重庆市蔬菜硝酸盐、亚硝酸盐含量及其与环境的关系[J]．生态学报，1996，16(4)：383-388．
[35] 周艺敏．氮素化肥对蔬菜硝酸盐积累的影响[J]．华北农学报，1986，4(1)：110-115．
[36] Van der Boon J. Steehuizen J W. Steingraever EG Growth and nitrate concentration of lettuce as affected by total nitrogen and chloride concentration NH_4~+-N/NO_3~桸 ratio and temperature of the recirculating nitrate solution. J Hort Sci. 1990.63(3):309.
[37] 朱祝军，蒋有条．不同形态氮素对不结球白菜生长和硝酸盐积累的影响[J]．植物生理学通迅，1994，30(3)：198-201．
[38] 高祖明，张耀栋，张道勇等．NPK对叶菜类硝酸盐积累和硝酸还原酶、过氧化物酶的影响[J]．园艺学报，1989，16(4)：293-297．
[39] 常丽新．钾肥在小白菜和萝卜上的施用效果[J]．中国蔬菜，2002，(1)：16-17．
[40] 胡勤海，叶兆杰，马生良．杭州市蔬菜硝酸盐污染现状及防治对策[J]．环境污染与防治，1991，13(4)：5-9．
[41] 黎星辉，黄启为，彭建伟．施肥对蔬菜硝酸盐含量的影响，菜园土壤肥力与蔬菜合理施肥[M]．南京：河海大学出版社，1997，202-206．
[42] 王正银，刘金萍，涂从等．几种化学物质用量对叶类蔬菜硝酸盐和营养品质的影响[J]．土壤农化通报，1998，13(2)：16-20．
[43] 黄启为，彭建伟．化肥对蔬菜硝酸盐含量的影响[J]．湖南农业大学学报(自然科学版)，2002，28(5)：387-390．
[44] 李冰，王昌全，冯长春等．喷施不同钙肥对莴笋产量及品质的影响[J]。中国农学通报，2004，20(2)：129-131．
[45] 余光辉，张杨珠，万大娟．几种硝化抑制剂对土壤和小白菜硝酸盐含量及产量的影响[J]．应用生态学报，2006，17(2)：247-250．
[46] 殷允相，杨俊，林孔仪等．银川地区蔬菜硝酸盐类含量、污染评价及防治途径研究[J]．宁夏农林科技，1993，(1)：40-43．
[47] 熊艳，尹增松等．蔬菜中硝酸盐污染现状及其防治措施[J]．云南农业大学学报，2003，18(3)：31-35．
[48] 汪李平，向长萍，王运华．白菜不同基因型硝酸盐含量差异的研究[J].园艺学报，2004，31(1)：43-46．
[49] Rein ink K Genetics of nitrate content of lettuce I. Analysis of generation means[J]. Euphytica,1991, 54: 83-92.
[50] 林家宝，陈火英，林观捷．蔬菜硝酸含量遗传的初步研究[J]．上海农学院学报，1995(2)：98-102．
[1].EvansH J,Nason A.,1952, Arch.Biochem.BioPhys.39:234-235.
[2]Evans H J,NasonA.,1953, Plant Physiol. 28:233-254.
[3] Tang P S,WuH.Y.,1957, Nature, 179:1355-1356.
[4]Kessler E,1964, Ann. Rev. Plant Physiol. 15:57-72.
[5]BeeversL,HagemanR. H., 1969, Ann. Rev. Plant Physiol. 20:495-522.
[6] Simon B C,Christopher E C. Characterization of the nitrite reductase gene Nii and the nitrate-assimilation gene cluster of Stagonospora (Septoria) nodorum[J].Curr Genet,1999,36:282-289.
[7] Samia D,Isabelle Q,Marie-Therese L, et al. Expression of a deregulated tobacco nitrate reductase gene in potato increases biomass production and decreases nitrate concentration in all organs. [J].Planta, 2004, 219:884-893.
[8] Cheneby D,Hallet S, Mondon M,et al.Genetic characterization of the nitrate reducing community base on G nucleotide sequence analysis [J].Microbial Ecology, 2003, 46(1):113-121.
[9]Antipov AN,Sorokin D Y. New enzyme belonging to family of molybdenum-free nitrate reductase[J].Biochemical journal, 2003, 369(1):185-189.
[10]Cheng C L,Dewdney J,Kleinhofs A,er al.A new locus (Nit2) nitrogene regulatory Protein: importance of the number, spacing and orientation of GATA core elements and their flanking sequences upon NIT2 binding [J].Biochemistry, 1988, 33:576-582.
[11] Crawford N M,Campbell W H, Davis R. Nitrate reductase from squash: cDNA clining and nitrate regulation [J].Proc. Natl.Acad. Sci. USA, 1986, 83:8073-8076.
[12]Wilkinson J Q,Crawford N M. identification of the Arabisposis CHL3 gene as the nitrate reductase structural gene NIA2[J]. Plant Cell,1991,3:461-471.
[13] Choi H, Kieinhofs A, An G Nucleotide sequence of rice nitrate reductase genes[J]. Plant Mol.Biol,1989,13:731-733.
[14] Daniel-Vedele F, Dorbe M F, Caboche M, et al. Cloning and analysis of the nitrate erduetase gene form tomato: A comparison of nitrate reductase protein sequences in higher Plants[J].Plants.Genet,1989, 220:88-94.
[15] 王利群，王勇，董英，等．硝酸盐对硝酸还原酶活性的诱导及硝酸还原酶基因的克隆[J]．生物工程学报，2003，19(5)：632-635．
[16] Cherel I,Gonneau M, Meyer C, et al. Biochemical and immunological characterization of nitrate reductase-deficient nia mutants of Nicotiana Plumbaginifolia [J].Plant Physiol.1990,92:659-665.
[17] Kubo Y,Ogura N,Nakagawa H. Limites Protelysis of the nitrate reductase from spinach leave[J].Biol Chem.,1998, 263:19684-19689.
[18] Masatoshi S,Hiroakil,Shinya N,et al. Cloning of a nitrate reductase in activateor (NRI) cDNA form spinacia oleracea L. and expression of mRNA and protein of NRI in cultured spinach cells[J]. Planta, 2003, 216:961-968.
[19] Hoff T, Stummann B M, Henningsssen K W, et al. Cloning and expression of a gene encoding a root specific nitrate reductase in bean[J]. Physiol. Plant,1991,82:197-204.
[20] Jensen P E, Moller M Q, Stummann B M, et al. Identification and characterization of a nitrate reductase gene form bean containing four introns[J]. Physiol. Plant, 1994, 92:613-623.
[21] Miyazkai J, Juricek M, Angelis K, et al.Characterization and sequence of a novel nitrate reductase form barley[J]. Mol.Gen.Genet.,1991, 228:329-334.
[22] Schnorr K M, Juricek M, Huang C, et al Analysis of barley nitrate reductase cDNA and genomic coones[J]. Mol. Genet., 1991, 227:411-416.
[23] 孙淑斌，罗金葵，徐国华，等．小白菜硝酸还原酶基因的克隆与初步鉴定[J] 植物营养与肥料学报2006，12(4)：592-596．
[24] Perng-Kuang Chang, Kenneth C. Ehrlich, John E.Linz, Deepak Bhatnagar, et al.Characterization of the Aspergillus Parasiticus niaD and niiA gene cluster[J]. Curr Genet, 1996, 30:68-75.
[25] Nicholas D. J. D., 1961,Ann.Bev.Plant Physiol.12:63-90.
[26]Guerrero, M.G,Vega, J. M., Losada,M.,The assimilatory nitrate-reduction and its regulation.Ann. Rev. Plant Physiol.,1980,32:169-402.
[27] Hewitt E. J., Ann.Rev. Plant Physiol. 1975, 26:73-158.
[28]Somer D.A.,et al.Plant Physiol,1983, 72:949-952.
[29]Funkhouser E,A.,et al,1983, Plant Physiol. 65:939-943.
[30]Funkhouser E,A.,Ramadoss C.S.,1980, Plant Physiol.65:944-948.
[31] Loppes R, Radnox M. Two short regions of the Promoter are essential of activation and repression of the nitrate reductase gene in Chlamydomonas reinhardtii [J]. Molecular Genetic and Genomics,2002, 268(1): 42-48.
[32] Melzer M J, Edwards R A. Nitrogen control in Bacteria[J]. Microbiol. Rev,1995,59:604-622.
[33] Callaci J J, Smarrelli J Jr. Regulation of the inducible nitrate reductase isoform from soybeans[J].Biochim.Biophys.Acta, 1991, 1088:127-130.
[34] Crawford N M. Nitrate: Nutrient and signal for Plant growth[J]. Plant Cell, 1995, 7: 859-868.
[35] Srivastava H S. Regulation of nitrate reductase activity in higher Plants[J]. Phytochemistry, 1980,19:725-733.
[36] Frost W B. et al. Plant Physiol, 1978, 61:323-331.
[37] Remmol/Ller J L,Campbell. Plant Physiol. 1986, 80:442.
[38] Calza R.Mol Gen Genet.,1987, 209:552.
[39] Campbell W H. Nitrate reductase and its role in nitrate assimilation in Plants[J]. Physiol Plant,1988, 74:214-219.
[40] Wallaee W,O aks A. In Dalling M J(ed), Plant Protaolytic Enzymes(Vol.2). CRC Press, Boca Raton, FL.1986, p.81.
[41]SorgerGJ.et al.Biochem.J, 1974, 140:395.
[42] Gerendas J, Zhu Z, Bendixen R, Ratcliffe R G. Sattelmacher B. Physiological and biochemical Processes related to ammol/Lonium toxicity in higher plants. Zeitschrift fur Pflanzenernahrung and Bodenkunde.1997, 160:239-251.
[43] Raab T K,Terry N. Nitrogen source regulation of growth and Photosynthesis in Beta vulgaris L.Plant Physiology. 1994, 105: 1159-1166.
[44].Magalhaes JR, Huber DW. Ammol/Lonium assimilation in different plants species as affected by nitrogen from and pH control in solution culture. Fertilizer Research, 1989, 21:1-6.
[45].Mengel K,Viro M. The significance of plant energy status for the uptake and incorporation of NH_4~+ - nitrogen by young rice plants. Soil Sci. Plant Nutr. 1978, 24: 407-416.
[46] 封克，汪晓丽，陈平等．水稻苗期不同时段NO3~-吸收特点及其受NH_4~+的影响[J]．中国农业科
?学，2003，36(3)：307-312．
[47] 封克，汤炎，张素玲．钱离子对不同基因型水稻吸收硝酸根离子的影响[J]．植物生理学通讯，2001，37(3)：192-194．
[48] Minotti P L, Williams D C, Jackson W A. The influence of ammol/Lonium on nitrate reduction in wheat seedlings. Planta, 1969, 86:267-271.
[49] Kronzuker H J, Glass AD M,Siddiqi M Y. Inhibition of nitrate uptake by ammol/Lonium in barley.Analysis of component fluxes.Plant Physiology,1999,120:283-291.
[50] Lycklama J. The absorption of ammol/Lonium and nitrate by Perennial ryegrass. Acta. Bot. Neer.1963, 12:361-462.
[51] Rao K P, Rains D W. Nitrate absorption by barley:Ⅱ.Influence of nitrate reductase activity. Plant Physiology, 1976, 57:59-62.
[52] Samuelson M E, Ohlen L, Lind M, Larsson C M. Nitrate regulation of nitrate uptake and nitrate reductase expression in barley grown at different nitrate: ammol/Lonium ratios at constant relative nitrogen addition rate. Physiologia Plantarum, 1995, 94:254-260.
[53] Youngdahl L J, Pacheco R, Street J J, Vlek P L G The kinetics of ammol/Lonium and nitrate uptake by young rice plants. Plant And Soil,1982,69:225-232.
[54] 廖祥儒，朱新产．氨积累对植物生长发育的效应[J]．农业环境保护，1996，15(6)：281-287．
[55] 何新华，Ann Oak S，李明启．C3和C4植物的氮素利用效率[J]．云南植物研究，1994，16(1)：93-94．
[56] 高廷东．光和不同形态氮互作对小麦幼苗碳氮同化的影响．山东农业大学硕士学位论文2003．
[57] Lillo C,Ruoff P. An unusually rapid light-induced nitrate reductase mRNA pulse and circadian osicillation. Naturuis-senschaften, 1989, 76:526-532.
[58] 林振武．硝酸还原酶的研究动态[J]．植物生理学通讯，1987，(6)：8-13．
[59]Deng M. D., Moureaux T., Cherel I. et al. Effects of nitrogen metabolites on the regulation and circadian expression of tobacco NR. Plant Physiol Biochem.,1991, 29:239-247.
[60] 余让才，李明启，范燕平．高等植物硝酸还原酶的光调控[J]．植物生理学通讯，1997，33(1)：61-65．
[61] Deng M, Moureaux T., Caboche M, et al. A molybdate analog in activating nitrate reductase,deregulates the expression of the nitrate reductase structure gene[J]. Plant Physiol.1989, 91:304-309.
[62] Kaiser W M, Spill D, Brendle-Behnisch E. Adenine nucleotide are apparently involves in the light-dark modulation of spinach-leaf nitrate reductase. Planta,1992,186:236-240.
[63] Kaiser W M, Huber S C. Posttranslational regulation of nitrate reductase in higher Plants [J]. Plant Physiol., 1994, 106:817-821.
[64] Douglas P, Morrice N, Mackintosh C. Identification of a regulatory Phosphorylation site in the hinge region of nitrate reductase from spinach leaves [J]. EFBS Lett. 1995,377:113-117.
[65]Bachmann M, Hunter H L,Liao P C. et al.The inhibitor protein of Phosphorylated nitrate reductase from spinach leaves is al4-3-3 protein[J]. EFBS Lett, 1996. 387:127-131.
[66]Chantarotwong W., Huffaker, RC,Miller,B.L., Granstedt,R.C.,In vivo nitrate reduction in relation to nitrate uptake,nitrate content, and in vivo nitrate reductase activity in intact barley seedlings[J]. Plant Physiol., 1976, 57:519-522.
[67] Mackown C T, Jackson W A, Volk R G Partitioning of Previously accumulated nitrate to translocation reduction and efflux in corn roots[J]. Planta 1983, 157:8-14.
[68] Udayakumar M, Devendra R, Reddy V S,Sastry K S K.Nitrate availability under low irradiance and its effect on the nitrate reductase activity. New Phytol, 1981, 88:290-297.
[69] Pearson J,Steer B T. Planta, 1977. 137:107.
[70] Taluoizte A, Champigny M L. Plant Cell Environ, 1988, 11:149.
[71] Talouize A, Guiraud G, Moyse A, Marol C M, Chmapigny M L,Effect of previous nitrate deprivation on ~(15)N-nitrate absorption and assimilation by wheat seedlings[J]. Plant Physiol, 1984,116:113-122.
[72].Passama L.et al.Plant Soil, 1987, 102:145.
[73] Reed A J,Hageman RH. Plant Physiol, 1980, 66:1184.
[74] 许长蔼，倪晋山．小麦叶内硝酸还原的代谢库[J]．植物生理学报，1990，16(3)：277-283．
[75] Shaner D L, Boyer J S. Nitrate reductase activity in maize leaves regulation by nitrate flux. Plant Physiol, 1976, 58:499-504.
[76] Martinoia E, Heck U, Wiemker A.Vacuoles as storage compartments for nitrate in barley leaves.Nature, 1981,289:292-293.
[77] 许长蔼，倪晋山．NO_3~-上运和液泡内NO_3~-外流对小麦叶内硝酸还原酶活性的调节[J]．植物生理学报，1990，16(4)：340-346．
[78] Yamazaki M. Plant Cell Physiol, 1986, 27:443.
[79] Amy N K,Carrett R H. Purification and characterization of the nitrate reductase from the diatom Thalassiosira pseudonana[J]. Plant Physiol, 1974, 54: 629-637.
[1] Britto D T,Kronzucker H J.NH_4~+ toxicity in higher plants:A critical review [J].J.Plant Physiol.,2002,159:567-584.
[2] 王利群，董英，黄达明等．蔬菜硝酸盐的积累及其生理机制研究进展[J]．江苏农业科学,2006，(6)：78-811。
[3] 林观捷，林家宝，陈火英．影响蔬菜硝酸盐含量积累因素的探讨[J]．上海农学院学报，1995，13(1)：47-52．
[4] Smiciklas KD,Below F E. Role of nitrogen form in determining yield of field-grown maize [J].Crop Science , 1992 , 32 : 1220-1225.
[5] 张春兰，高祖明，张耀栋等．氮素形态和NO_3~-N:与NH_4~+-N配比对菠菜生长和品质的影响[J]．南京农业大学学报，1990，13(3)：70-74．
[6] 陈巍，罗金葵，姜慧梅等．不同形态氮素比例对不同小白菜品种生物量和硝酸盐含量的影响[J]．土壤学报，2004，41(3)：420-425．
[7] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社．
[8] 封克，汪晓丽，陈平等．水稻苗期不同时段NO_3~-吸收特点及其受NH~+的影响[J]．中国农业科学，2003，36(3)：307-312．
[9] Guerrero, M G, Vega, J M., Losada,M.,The assimilatory nitrate-reduction and its regulation.Ann.Rev. Plant Physiol.,1980,32:369-402.
[10] Cokojiob O A．氮肥施用的生态观点[J]．土壤学进展，1992(1)：19-21．
[11] Granstedt R C.Huffaker R C. Identification of the leaf vacuole as a major nitrate storage pool. Plant Physiol.l982,70:410-413.
[12] Raab T K,Terry N.Nitrogen source regulation of growth and Photosynthesis in Beta vulgaris L.Plant Physiology. 1994,105:1159-1166.
[13] Gerendas J, Zhu Z. BendixenR.RatcliffeRG. Sattelmacher B. Physiological and biochemical Processes related to ammol/Lonium toxicity in higher plants. Zeitschrift fur Pflanzenernahrung and Bodenkunde.1997,160:239-251.
[14] 孙传范，戴廷波，曹卫星．不同施氮水平下增铵营养对小麦生长和氮素利用的影响[J]．植物营养与肥料学报，2003，9(1)：33-38。
[15] Kronzuker H J,Glass A D M,Siddiqi M Y. Inhibition of nitrate uptake by ammol/Lonium in barley.Analysis of component fluxes. Plant Physiology, 1999, 120:283-291.
[16]Samuelson M E, Ohlen L, Lind M, Larsson C M. Nitrate regulation of nitrate uptake and nitrate reductase expression in barley grown at different nitrate: ammol/Lonium ratios at constant relative nitrogen addition rate. Physiologia Plantarum,1995,94:254-260.]
[17] 董晓英，李式军，沈仁芳．白菜不同品种对硝酸盐吸收积累差异原因初探[J] 园艺学报，2003，30(4)：470-472．
[1] Fytianos K,Zarogiannis P. Nitrate and nitrite accumulation in fresh vegetables from Greece.Bull Environ Contain Toxicol, 1999,62:187-192.
[2] 陈君石、闻芝梅主译．食物、营养与癌症预防．上海：上海医科大学出版社，1999．
[3] White J W J. Relative significance of daietarg source of nitrate and nitrite.[J]. Agric Food Chem.l975,23(5):886-891.
[4] Evans H J, Nason A. Pyridine nucleotide nitrate reductase from extracts of higher plants[J]. Plant Physiol,1953, 28:233-254.
[5] 郭世荣 无土栽培学[M]．北京：中国农业出版社，2004．125-129．
[6] Deng M,Moureaux T, Caboche M. Tungstate.A molybdate analog inactivating nitrate reductase,deregulates the expression of the nitrate reductase structural gene.[J]. Plant Physiol,1989,91:304-309.
[7] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．123-128．
[8] Hageman R H, D P Hucklesby.Methods in Enzymology[J].1971,23 (A): 491-503.
[9] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．195-197．
[10] 林立欣，李寅安，王美菊．萝卜杂种优势与配合力的研究[J]．中国农业科学，1986年(2)：54-60
[11] 司江英，汪晓丽，陈平等．硝酸还原酶抑制剂和NH_4~+对不同基因型水稻苗期NO_3~-吸收的影响[J]．扬州大学学报(农业与生命科学版)．2004，25(1)：59-62．
[12] Robert H. Desirée Gómez F. Tobacco plants that lack expression of functional nitrate reductase in roots shows changes in growth rates and metabolite accumulation.[J].Journal of Experimental Botany,2001,52,No.359:1251-1258.
[1] 林振武．硝酸还原酶的研究动态[J]．植物生理学通讯，1987，(6)：8-13．
[3] 刘丽，甘志军，王宪泽．植物氮代谢硝酸还原酶水平调控机制的研究进展[J]．西北植物学报，2004，24(7)：1355-1361．
[2] Magalhase J R. The Responses of Ammonium assimilation enzymes to nitrogen form in different Plant[J].Journal of Plant Nutrition,1991,14(2):175-185.
[4] Deng M, Moureaux T, Caboche M. Tungstate, a molybdate analog inactivating nitrate reductase,deregulates the expression of the nitrate reductase structural gene[J]. Plant Physiol, 1989,91:304-309.
[5] 李合生主编．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．123-128．
[6] Mohotti A J,Lawlor D W. Diurnal variation of Photo synthesis and Photo-inhibitionin tea:effects of irradianee and nitrogen supply during growth in the field[J].Journal of ExperimentalBotany,2002,53(367):313-322.
[7] Huber J L,Huber S C,Campelli W H. Reversible light/dark modulation of spinach leaf nitrate reductase activity involved protein phosphorylation[J].Arch Biochem Biophys,1992, 296(1):58-65.
[8] Champigny M L. Integration of photosynthetic carbon and nitrogen metabolism in higher plants[J].Photosynthesis Research,1995, 46:117-127.
[9] Zavala J A, Ravetta A D.Allocation of Photoassimilates to biomass,resin and carbohydrates in Grindelia chiloensis as affected by light intensity[J].Field crops research,2001,69:143-149.
[10] 关义新，林葆，凌碧莹．光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J]．作物学报，2000，26(6)：806-812．
[11] 朱为民．蔬菜硝酸盐积累的基因型差异与营养调控及分子生物学的研究[D]．南京：南京农业大学，1997．
[1] Kaiser B N. The role of molybdenum in agricultural plant production. Annals of Botany , 2005 , 96(5):745-754.
[2] 庞静，胡承孝，王运华等．钼对黄棕壤上冬小麦碳代谢的影响．华中农业大学学报，2001，20(1)：33-35．
[3] Fang Z,Zhang Y G,Zhang F S,et al.Mechanisms of difference in Mo efficiency between wheat and oil seed rape. Pedosphere[J] 2000,10(3) :213-220.
[4] 喻敏，王运华．高等植物中的钼．植物学通报，1999，16(5)：547-554．
[5] 杜新民．锰对小白菜产量和品质的影响[J]．2007，27(5)：48-51．
[6] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．248-249．
[7] Hageman R H,Hucklesby D P.Methods in Enzymology[J].1971,23(A): 491-503.
[8] 李合生．植物生理生化实验原理和技术[M]．北京：高等教育出版社，2000．195-197．
[9] 王立克，洪法水．钼浸种对苜蓿硝酸还原酶活性及营养成分的影响[J]．中国草地．1995，15(3)：37-38．
[10] 杜应琼，廖新荣，黄志尧等．硼、钼对花生氮代谢的影响[J]．作物学报，2001，27(5)：612-616．
[11] 付其如，何纪荣．钼对葡萄叶片硝酸还原酶活性及生长发育的影响．四川师范学院学报，1996,17(1)：56-60．
[12] 张录强．钼(Mo)的生理及其生物学效应[J]．生物学通报，1999,34(11)：24-26．
[13] Chitralekha C, Ninnala N. Sugarcane yield and physiology in response to molybdenum supply.Sugar Cane, 1999,(3):19-20.
[14] 宋志荣．施锰对马铃薯产量和品质的影响[J]．中国农学通报，2005，21(3)：222-223．
[15] 杨保川，王昌全，项虹艳等．叶面喷施钼锰对莴笋生长发育和硝酸盐含量的影响[J]．农业环境科学学报，2006，25(增刊)：96-99．
[16] 杜慧玲，王建锁，姚永平．钾锌锰配施对甜椒品质的影响[J]．中国农学通报，2000，16(5)：32-33．
[17] 刘鹏，徐根娣，周子仙等．Mo和Mn对七子花几种生理效应的影响[J]．林业科学研究，2002，15(2)：156-162．
[18] 关义新，林葆，凌碧莹．光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J]．作物学报，2000，26(6)：806-812．