Nitrogen Assimilation and its Relevance to Crop Improvement

Nitrogen Metabolism in Plants in the Post-genomic Era

Volumn 42, Issue , 2010, Pages 1-40

  Lea, Peter J ^a   Miflin, Ben J ^b  

^a LANCASTER UNIVERSITY   (United Kingdom)

^b ROTHAMSTED RESEARCH   (United Kingdom)

Author keywords

Ammonia assimilation; Asparaginase; Asparagine synthetase; Gene regulation; Glutamate dehydrogenase; Glutamate synthase; Glutamine synthetase; Overexpression

Indexed keywords

EID: 84948965974     PISSN: None     EISSN: None     Source Type: Book    
DOI: 10.1002/9781444328608.ch1     Document Type: Chapter

References (205)

1. Alonso, J.M., Stepanova, A.N., Leisse, T.J., et al. (2003) Genome-wide insertional mutagenesis of Arabidopsis thaliana. Science 301, 653-657.
2. Ameziane, R., Bernhard, K. & Lightfoot, D. (2000) Expression of the bacterial gdhA gene encoding aNADPH glutamate dehydrogenase in tobacco affects plant growth and development. Plant and Soil 221, 47-57.
3. Andrews, M., Lea, P.J., Raven, J.A., et al. (2004) Can genetic manipulation of plant nitrogen assimilation enzymes result in increased crop yield and greater N-use efficiency? Annals of Applied Biology 145, 25-40.
4. Atkins, C.A., Pate, J.S. & Sharkey, P.J. (1975) Asparagine metabolism - key to the nitrogen nutrition of developing legume seeds. Plant Physiology 56, 807-812.
5. Barsch, A., Carvalho, H.G., Cullimore, J.V., et al. (2006) GS-MS based metabolite profiling implies three interdependent ways of ammonium assimilation in Medicago truncatula root nodules. Journal of Biotechnology 127, 79-83.
6. Bascomb, N.F., Prunkard, D.E. & Schmidt, R.R. (1987) Different rates of synthesis and degradation of two chloroplastic ammonium-inducible NADP-specific glutamate dehydrogenase isoenzymes during induction and deinduction in Chlorella sorokiniana cells. Plant Physiology 83, 85-91.
7. Becker, T.W., Carrayol, E. & Hirel, B. (2000) Glutamine synthetase and glutamate dehydrogenase isoforms inmaize leaves: localization, relative proportion and their role in ammonium assimilation or nitrogen transport. Planta 211, 800-806.
8. Bellucci, M., Ederli, L., DeMarchis, F., et al. (2004) Transformation of Lotus corniculatus plants with Escherichia coli asparagine synthetase A: effect on nitrogen assimilation and plant development. Plant Cell Tissue and Organ Culture 78, 139-150.
9. Bernard, S.M., Møller, A.L.B., Dionisio, G., et al. (2008) Gene expression, cellular localisation and function of glutamine synthetase isozymes in wheat (Triticum aestivum). Plant Molecular Biology 67, 89-105.
10. Blackwell, R.D., Murray, A.J.S., Lea, P.J., et al. (1988) The value of mutants unable to carry out photorespiration. Photosynthesis Research 16, 155-176.
11. Blanco, L., Reddy, P.M., Silvente, S., et al. (2008) Molecular cloning, characterization and regulation of twodifferentNADH-glutamate synthase cDNAs in bean nodules. Plant Cell and Environment 31, 454-472.
12. Boisson, M., Mondon, K., Torney, V., et al. (2005) Partial sequences of nitrogen metabolism genes in hexaploid wheat. Theoretical and Applied Genetics 110, 932-940.
13. Borek, D., Podkowinski, J., Kisiel, A., et al. (1999) Isolation and characterization of cDNA encoding L-asparaginase from Lupinus luteus (Accession, AF112444). Plant Physiology 119, 1568.
14. Borek, D., Michalska, K., Brzezinski, K., et al. (2004) Expression, purification and catalytic activity of Lupinus luteus asparagine β-amidohydrolase and its Escherichia coli homolog. European Journal of Biochemistry 271, 3215-3226.
15. Botella, J.R., Verbelen, J.P. & Valpuesta, V. (1988) Immunocytolocalisation of glutamine synthetase in green leaves and cotyledons of Lycopersicon esculentum. Plant Physiology 88, 943-946.
16. Bowsher, C.G., Lacey, A.E., Hanke, G.T., et al. (2007) The effect of Glc6P uptake and its subsequent oxidation within pea root plastids on nitrite reduction and glutamate synthesis. Journal of Experimental Botany 58, 1109-1118.
17. Brears, T., Liu, C., Knight, T.J., et al. (1993) Ectopic overexpression of asparagine synthetase in transgenic tobacco. Plant Physiology 103, 1285-1290.
18. Brugière, N., Dubois, F., Masclaux, C., et al. (2000) Immunolocalization of glutamine synthetase in senescing tobacco (Nicotiana tabacum L.) leaves suggest that ammonia assimilation is progressively shifted to the mesophyll cytosol. Planta 211, 519-527.
19. Bruneau, L., Chapman, R. & Marsolais, F. (2006) Co-occurrence of both L-asparaginase subtypes in Arabidopsis: At3g16150 encodes a K+-dependent L-asparaginase. Planta 224, 668-679.
20. Cañas, R.A., de la Torre, F., Cánovas, F.M., et al. (2006) High levels of asparagine synthetase in hypocotyls of pine seedlings suggest a role of the enzyme in reallocation of seed-stored nitrogen. Planta 224, 83-95.
21. Cañas, R.A., de la Torre, F., Cánovas, F.M., et al. (2007) Coordination of PsAS1 and PsASPG expression controls timing of re-allocated N utilization in hypocotyls of pine seedlings. Planta 225, 1205-1219.
22. Cánovas, F.M., Avila, C., Cantón, F.R., et al. (2007) Ammonium assimilation and amino acid metabolism in conifers. Journal of Experimental Botany 58, 2307-2318.
23. Canton, F.R., Suarez, M.F., Jose-Estanyol, M., et al. (1999) Expression analysis of a cytosolic glutamine synthetase gene in cotyledons of Scots pine seedlings: developmental, light regulation and spatial distribution of specific transcripts. Plant Molecular Biology 40, 623-634.
24. Carvalho, H. & Cullimore, J. (2003) Regulation of glutamine synthetase isoenzymes and genes in the model legume Medicago truncatula. In: Pandalai, S.G. (ed.) Recent Research Developments in Plant Molecular Biology, Vol. 1, Pt 1. Research Signpost, Trivandrum, India, pp. 57-175.
25. Carvalho, H.G., Lopes-Cardoso, I.A., Lima, L.M., et al. (2003) Nodule-specific modulation of glutamine synthetase in transgenic Medicago truncatula leads to inverse alterations in asparagine synthetase expression. Plant Physiology 133, 243-252.
26. Chang, K.S. & Farnden, K.J. (1981) Purification and properties of asparaginase from Lupinus arboreus and Lupinus angustifolius. Archives of Biochemistry and Biophysics 208, 49-58.
27. Chen, F-L. & Cullimore, J.V. (1988) Two isoenzymes of NADH-dependent glutamate synthase in root nodules of Phaseolus vulgaris L: purification, properties and activity changes during nodule development. Plant Physiology 88, 1411-1417.
28. Chichkova, S., Arellano, J., Vance, C.P., et al. (2001) Transgenic tobacco plants that overexpress alfalfa NADH-glutamate synthase have higher carbon and nitrogen content. Journal of Experimental Botany 52, 2079-2084.
29. Cho, C-W., Lee, H-J., Chung, E., et al. (2007) Molecular characterization of the soybean L-asparaginase gene induced by low temperature stress. Molecules and Cells 23, 280-286.
30. Cock, J.M., Kim, K.D., Miller P.W., et al. (1991) A nuclear gene with many introns encoding ammonium-inducible ammonium inducible chloroplastic NADP-specific glutamate dehydrgenase(s) in Chlorella sorokiniana. Plant Molecular Biology 17, 1023-1044.
31. Collard, B.C.Y. & Mackill, D.J. (2008) Marker-assisted selection: an approach for precision plant breeding in the twenty-first century. Philosophical Transactions of the Royal Society B: Biological Sciences 363, 557-572.
32. Coschigano, K.T., Melo-Oliveira, R., Lim, J., et al. (1998) Arabidopsis gls mutants and distinct Fd-GOGAT genes: implications for photorespiration and primary nitrogen assimilation. Plant Cell 10, 741-752.
33. Cottevieille, M., Larquet, E., Jonic, S., et al. (2008) The subnanometer resolution structure of the glutamate synthase 1.2MDahexamer by cryoelectron microscopy and its oligomerization behaviour in solution - functional implications. Journal of Biological Chemistry 283, 8237-8249.
34. Cullimore, J.V., Gebhardt, C., Saarelainen, R., et al. (1984) Glutamine synthetase of Phaseolus vulgaris organ-specific expression of a multigene family. Journal of Molecular and Applied Genetics 2, 589-600.
35. Davies, K.M. & King, G.A. (1993) Isolation and characterization of a cDNA clone for a harvest induced asparagine synthetase from Asparagus officinalis L. Plant Physiology 102, 1337-1340.
36. Dickson, J.M., Vincze, E., Grant, M.R., et al. (1992) Molecular cloning of the gene encoding developing seed L-asparaginase from Lupinus angustifolius. Plant Molecular Biology 20, 333-336.
37. Donn, G., Tischer, E., Smith, J.A., et al. (1984) Herbicide-resistant alfalfa cells an example of gene amplification in plants. Journal of Molecular and Applied Genetics 2, 621-636.
38. Dubois, F., Tercé-Laforgue, T., Gonzalez-Moro, M.B., et al. (2003) Glutamate dehydrogenase in plants: is there a new story for an old enzyme? Plant Physiology and Biochemistry 41, 565-576.
39. Eckes, P., Schmitt, P., Daub, W., et al. (1989) Overproduction of alfalfa glutaminesynthetase in transgenic tobacco plants. Molecular and General Genetics 217, 263-268.
40. Edwards, J.W., Walker, E.L. & Coruzzi, G.M. (1990) Cell-specific expression in transgenic plants reveals nonoverlapping roles for chloroplast and cytosolic glutaminesynthetase. Proceedings of the National Academy of Sciences of the United States of America 87, 3459-3463.
41. Fei, H.M., Chaillou, S., Hirel, B., et al. (2003) Overexpression of a soybean cytosolic glutamine synthetase gene linked to organ-specific promoters in pea plants grown in different concentrations of nitrate. Planta 216, 467-474.
42. Fei, H., Chaillou, S., Hirel, B., et al. (2006) Effects of the overexpression of a soybean cytosolic glutamine synthetase gene (GS1-5) linked to organ-specific promoters on growth and nitrogen accumulation of pea plants supplied with ammonium. Plant Physiology and Biochemistry 44, 543-550.
43. Feraud, M., Masclaux-Daubresse, C., Ferrario-Méry, S., et al. (2005) Expression of a ferredoxin-dependent glutamate synthase gene in mesophyll and vascular cells and functions of the enzyme in ammonium assimilation in Nicotiana tabacum (L.). Planta 222, 667-677.
44. Ficarelli, A., Tassi, F. & Restivo, F.M. (1999) Isolation and characterization of twocDNA clones encoding for glutamate dehydrogenase in Nicotiana plumbaginifolia. Plant and Cell Physiology 40, 339-342.
45. Fontaine, J-X., Saladino, F., Agrimonti, C., et al. (2006) Control of the synthesis and subcellular targeting of the twoGDHgene products in leaves and stems of Nicotiana plumbaginifolia and Arabidopsis thaliana. Plant and Cell Physiology 47, 410-418.
46. Forde, B.G. & Lea, P.J. (2007) Glutamate in plants: metabolism, regulation and signalling. Journal of Experimental Biology 58, 2339-2358.
47. Finnemann, J. & Schjoerring, J.K. (2000) Post-translational modification of cytosolic glutamine synthetase by reversible phosphorylation and 14-3-3 protein interaction. Plant Journal 24, 171-181.
48. Forde, B.G. & Cullimore, J.V. (1989) The molecular biology of glutamine synthetase in higher plants. In: Miflin, B.J. (ed.) Oxford Surveys of Plant Molecular and Cell Biology, Vol. 6. Oxford University Press: Oxford, England, UK, pp. 247-299.
49. Forde, B.G., Day, H.M., Turton, J.F., et al. (1989) Two glutamine-synthetase genes from Phaseolus vulgaris L display contrasting developmental and spatial patterns of expression in transgenic Lotus corniculatus plants. Plant Cell 1, 391-401.
50. Fuentes, S.I., Allen, D.J., Ortiz-Lopez, A., et al. (2001) Over-expression of cytosolic glutamine synthetase increases photosynthesis and growth at low nitrogen concentrations. Journal of Experimental Botany 52, 1071-1081.
51. Gallais, A. & Hirel, B. (2004) An approach to the genetics of nitrogen use efficiency in maize. Journal of Experimental Botany 55, 295-306.
52. Gallardo, F., Fu, J.M., Canton, F.R., et al. (1999) Expression of a conifer glutamine synthetase gene in transgenic poplar. Planta 210, 19-26.
53. Gálvez-Valdivieso, G., Osuna, D., Maldonado, J.M., et al. (2005) Purification of a functional asparagine synthetase (PVAS2) from common bean (Phaseolus vulgaris), a protein predominantly found in root tissues. Plant Science 168, 89-95.
54. Garcia-Gutierrez, A., Dubois, F., Canton, F.R., et al. (1998) Two different modes of early development and nitrogen assimilation in gymnosperm seedlings. Plant Journal 13, 187-199.
55. Gebhardt, C., Oliver, J.E., Forde, B.G., et al. (1986) Primary structure and differential expression of glutamine-synthetase genes in nodules, roots and leaves of Phaseolus vulgaris. Embo Journal 5, 1429-1435.
56. Glevarec, G., Bouton, S., Jaspard, E., et al. (2004) Respective roles of the glutamine synthetase/glutamate synthase cycle and glutamate dehydrogenase in ammonium and amino acid metabolism during germination and post-germinative growth in the model legume Medicago truncatula. Planta 219, 286-297.
57. Gomes, M.A. & Sodek, L. (1984) Allantoinase and asparaginase activities in maturing fruits of nodulated and non-nodulated soybeans. Physiologia Plantarum 62, 105-109.
58. Good, A.G., Shrawat, A.K. & Muench, D.G. (2004) Can less yield more? Is reducing nutrient input into the environment compatible withmaintaining crop production? Trends in Plant Science 9, 597-605.
59. Good, A.G., Johnson, S.J., De Pauw, M., et al. (2007) Engineering nitrogen use efficiency with alanine aminotransferase. Canadian Journal of Botany 85, 252-262.
60. Goto, S., Akagawa, T., Kojima, S., et al. (1998) Organization and structure of NADHdependent glutamate synthase gene from rice plants. Biochimica et Biophysica Acta 1387, 298-308.
61. Grant, M. & Bevan, M.W. (1994) Asparaginase gene expression is regulated in a complex spatial and temporal pattern in nitrogen-sink tissues. Plant Journal 5, 695-704.
62. Gregerson, R.G., Miller, S.S., Twary, S.N., et al. (1993) Molecular characterization of NADH-dependent Glu synthase from alfalfa nodules. Plant Cell 5, 215-226.
63. Habash, D.Z., Massiah, A.J., Rong, H.L., et al. (2001) The role of cytosolic glutamine synthetase in wheat. Annals of Applied Biology 138, 83-89.
64. Habash, D.Z., Bernard, S., Schondelmaier, J., et al. (2007) The genetics of nitrogen use in hexaploid wheat: N utilisation, development and yield. Theoretical and Applied Genetics 114, 403-419.
65. Halford, N.G., Muttucumaru N., Curtis, T.Y., et al. (2007) Genetic and agronomic approaches to decreasing acrylamide precursors in crop plants. Food Additives and Contaminants 24, 26-36.
66. Hanson, J., Hanssen, M., Wiese, A., et al. (2008) The sucrose regulated transcription factor bZIP11 affects amino acid metabolism by regulating the expression of ASPARAGINE SYNTHETASE 1 and PROLINE DEHYDROGENASE 2. Plant Journal 53, 935-949.
67. Hayakawa, T., Nakamura, T., Hattori, F., et al. (1994) Cellular-localization of NADHdependent glutamate-synthase protein in vascular bundles of unexpanded leaf blades and young grains of rice plants. Planta 193, 455-460.
68. Hejazi, M., Piotukh, K., Mattow, J., et al. (2002) Isoaspartyl dipeptidase activity of plant-type asparaginases. Biochemical Journal 364, 129-136.
69. Hemon, P., Robbins, M.P. & Cullimore, J.V. (1990) Targeting of glutamine synthetase to the mitochondria of transgenic tobacco. Plant Molecular Biology 15, 895-904.
70. Herrera-Rodriguez, M.B., Carrasco-Ballesteros, S., Maldonado, J.M., et al. (2002) Three genes showing distinct regulatory patterns encode the asparagine synthetase of sunflower (Helianthus annuus). New Phytologist 155, 33-45.
71. Herrera-Rodriguez, M.B, Maldonado, J.M. & Pérez-Vicente, R. (2004) Light and metabolic regulation ofHAS1, HAS1.1 and HAS2, three asparagine synthetase genes in Helianthus annuus. Plant Physiology and Biochemistry 42, 511-518.
72. Herrera-Rodriguez, M.B., Maldonado, J.M. & Pérez-Vicente, R. (2006) Role of asparagine and asparagine synthetase genes in sunflower (Helianthus annuus) germination and natural senescence. Journal of Plant Physiology 163, 1061-1070.
73. Herrera-Rodriguez, M.B., Pérez-Vicente, R. & Maldonado, J.M. (2007) Expression of asparagine synthetase genes in sunflower (Helianthus annuus) under various environmental stresses. Plant Physiology and Biochemistry 45, 33-38.
74. Hirel, B., Bertin, P., Quillere, I., et al. (2001) Towards a better understanding of the genetic and physiological basis for nitrogen use efficiency in maize. Plant Physiology 125, 1258-1270.
75. Hirel, B., Martin, A., Tercé-Laforgue, T., et al. (2005) Physiology of maize I: a comprehensive and integrated view of nitrogen metabolism in a C4 plant. Physiologia Plantarum 124, 167-177.
76. Hirel, B., Chardon, F. & Durand, J. (2007) The contribution of molecular physiology to the improvement of nitrogen use efficiency in crops. Journal of Crop Science and Biotechnology 10, 123-132.
77. Hoshida, H., Tanaka, Y., Hibino, T., et al. (2000) Enhanced tolerance to salt stress in transgenic rice that overexpresses chloroplast glutamine synthetase. PlantMolecular Biology 43, 103-111.
78. 4+ production in leaves of wild-type and glutamine synthetase-2 antisense oilseed rape. Plant Physiology 130, 989-998.
79. Ishiyama, K., Inoue, E., Tabuchi, M., et al. (2004a) Biochemical background and compartmentalized functions of cytosolic glutamine synthetase for active ammonium assimilation in rice roots. Plant and Cell Physiology 45, 1640-1647.
80. Ishiyama, K., Inoue, E., Watanabe-Takahashi, A., et al. (2004b) Kinetic properties and ammonium-dependent regulation of cytosolic isoenzymes of glutamine synthetase in Arabidopsis. Journal of Biological Chemistry 279, 16598-16605.
81. Jaspard, E. (2006) A computational analysis of the three isoforms of glutamate dehydrogenase reveals structural features of the isoform EC. 1.4.1.4 supporting a key role in ammonium assimilation by plants. Biology Direct 1, 38
82. Jing, Z.P., Gallardo, F., Pascual, M.B., et al. (2004) Improved growth in a field trial of transgenic hybrid poplar overexpressing glutamine synthetase. New Phytologist 164, 137-145.
83. Joy, K.W. (1988) Ammonia, glutamine and asparagine: a carbon-nitrogen interface. Canadian Journal of Botany 66, 2103-2109.
84. Kendall, A., Wallsgrove, R.M., Hall, N.P., et al. (1986) Carbon and nitrogen-metabolism in barley (Hordeum vulgare L) mutants lacking ferredoxin-dependent glutamate synthase. Planta 168, 316-323.
85. Keys, A.J. (2006) The re-assimilation of ammonia produced by photorespiration and the nitrogen economy of C-3 higher plants. Photosynthesis Research 87, 165-175.
86. Keys, A.J., Bird, I.F., Cornelius, M J., et al. (1978) Photorespiratory nitrogen cycle. Nature 275, 741-743.
87. Kichey, T., Le Gouis, J., Sangwan, B., et al. (2005) Changes in the cellular and subcellular localization of glutamine synthetase and glutamate dehydrogenase during flag leaf senescence in wheat (Triticum aestivum L.). Plant and Cell Physiology 46, 964-974.
88. Kirby, E.G., Gallardo, F., Man, H., et al. (2006) The overexpression of glutamine synthetase in transgenic poplar: a review. Silvae Genetica 55, 278-284.
89. Kozaki, A. & Takeba, G. (1996) Photorespiration protects C3 plants from photooxidation. Nature 384, 557-560.
90. Krajewski, W.W., Collins, R., Holmberg-Schiavone, L., et al. (2008) Crystal structure of mammalian glutamine synthetases illustrate substrate-induced conformational changes and provide opportunities for drug and herbicide design. Journal ofMolecular Biology 375, 217-228.
91. Lam, H.M., Peng, S.S. & Coruzzi, G.M. (1994) Metabolic regulation of the gene encoding glutamine-dependent asparagine synthetase in Arabidopsis thaliana. Plant Physiology 106, 1347-1357.
92. Lam, H.M., Coschigano, K., Schultz, C., et al. (1995) Use of Arabidopsis mutants and genes to study amide amino acid biosynthesis. Plant Cell 7, 887-898.
93. Lam, H.M., Hsieh, M.H. & Coruzzi, G. (1998) Reciprocal regulation of distinct asparagine synthetase genes by light and metabolites in Arabidopsis thaliana. Plant Journal 16, 345-353.
94. Lam, H.M., Wong, P., Chan, H.K., et al. (2003) Overexpression of the asn1 gene enhances nitrogen status in seeds of Arabidopsis. Plant Physiology 132, 926-935.
95. Lancien, M., Martin, M., Hsieh, M.H., et al. (2002) Arabidopsis glt1-T mutant defines a role of NADH-GOGAT in the non-photorespiratory ammonium assimilatory pathway. Plant Journa1 29, 347-358.
96. Lea, P.J. & Azevedo, R.A. (2007) Nitrogen use efficiency. 2. Amino acid metabolism. Annals of Applied Biology 151, 269-275.
97. Lea, P.J. & Miflin, B.J. (1974) Alternative route for nitrogen assimilation in higher plants. Nature 251, 614-616.
98. Lea, P.J. & Miflin, B.J. (2003) Glutamate synthase and the synthesis of glutamate in plants. Plant Physiology and Biochemistry 41, 555-564.
99. Lea, P.J. & Thurman, D.A. (1972) Intracellular location and properties of plant Lglutamate dehydrogenases. Journal of Experimental Botany 23, 440-449.
100. Lea, P.J., Fowden, L. & Miflin, B.J. (1978) The purification and properties of asparaginase from Lupinus species. Phytochemistry 17, 217-222.
101. Lea, P.J., Sodek, L., Parry, M.A.J., et al. (2007). Asparagine in plants. Annals of Applied Biology 150, 1-26.
102. Lea, U.S., Leydecker, M-T., Quilleré, I., et al. (2006) Posttranslational regulation of nitrate reductase strongly affects the levels of free amino acids and nitrate, whereas transcriptional regulation has onlyminor influence. Plant Physiology 140, 1085-1094.
103. Leegood, R.C., Lea, P.J., Adcock, M.D., et al. (1995) The regulation and control of photorespiration. Journal of Experimental Botany 46, 1397-1414.
104. Lehmann, T. & Ratajczak, L. (2008) The pivotal role of glutamate dehydrogenase (GDH) in the mobilization of C and N from the storage material to asparagine in germinating seeds of yellow lupine. Journal of Plant Physiology 165, 149-158.
105. Li, M.G., Villemur, R., Hussey, P.J., et al. (1993) Differential expression of 6-glutaminesynthetase genes in Zea mays. Plant Molecular Biology 23, 401-407.
106. Li, R-J., Hua W. & Lu, Y-T. (2006) Arabidopsis cytosolic glutamine synthetase AtGLN1;1 is a potential substrate of AtCRK3 involved in leaf senescence. Biochemical and Biophysical Research Communications 342, 119-126.
107. Lightfoot, D.A., Green, N.K. & Cullimore, J.V. (1988) The chloroplast-located glutamine-synthetase of Phaseolus vulgaris L - nucleotide-sequence, expression in different organs and uptake into isolated-chloroplasts. Plant Molecular Biology 11, 191-202.
108. Lightfoot, D.A., Mungur, R., Ameziane, R., et al. (2007) Improved drought tolerance of transgenic Zea mays plants that express the glutamate dehydrogenase gene (gdhA) of E. coli. Euphytica 156, 103-116.
109. Lima, L., Seabra, A., Melo, P., et al. (2006) Phosphorylation and subsequent interaction with 14-3-3 proteins regulate plastid glutamine synthetase in Medigago truncatula. Planta 223, 558-567.
110. Lough T.J., Chang K.S., Carne A., et al. (1992a) L-asparaginase from developing seeds of Lupinus arboreus. Phytochemistry 31, 1519-1527.
111. Lough T.J., Reddington B.D., Grant M.T., et al. (1992b) The isolation and characterization of cDNA clone encoding L-asparaginase from developing seeds of lupin (Lupinus arboreus). Plant Molecular Biology 19, 391-399.
112. Loulakakis, K.A. & Roubelakis-Angelakis, KA. (1991) Plant NAD(H) glutamate dehydrogenase consists of two subunit polypeptides and their participation in the 7 isoenzymes occurred in an ordered ratio. Plant Physiology 97, 104-111.
113. Loulakakis, K.A. & Roubelakis-Angelakis, K.A. (1996) The seven NAD(H)glutamate dehydrogenase isoenzymes exhibit similar anabolic and catabolic activities. Physiologia Plantarum 96, 29-35.
114. 4+ assimilation in Zea mays. Plant Physiology 94, 647-656.
115. Martin, A., Lee, J., Kichey, T., et al. (2006) Two cytosolic glutamine synthetase isoforms of maize are specifically involved in the control of grain production. Plant Cell 8, 3252-3274.
116. Masclaux-Daubresse, C., Valadier, M.H., Carrayol, E., et al. (2002) Diurnal changes in the expression of glutamate dehydrogenase and nitrate reductase are involved in the C/N balance of tobacco source leaves. Plant Cell and Environment 25, 1451-1462.
117. Masclaux-Daubresse, C., Carrayol, E. & Valadier, M.H. (2005) The two nitrogen mobilisation- and senescence-associated GS1 and GDH genes are controlled by C and N metabolites. Planta 221, 580-588.
118. Masclaux-Daubresse, C., Reisdorf-Cren, M., Pageau, K., et al. (2006) Glutamine synthetase-glutamate synthase pathway and glutamate dehydrogenase play distinct roles in the sink source nitrogen cycle in tobacco. Plant Physiology 140, 444-456.
119. Melo-Oliveira, R., Oliveira, I.C. & Coruzzi, G.M. (1996) Arabidopsis mutant analysis and gene regulation define a non-redundant role for glutamate dehydrogenase in nitrogen assimilation. Proceedings of the National Academy of Sciences of the United States of America 93, 4718-4723.
120. Michalska, K., Borek, D., Hernandez-Santoyo, A., et al. (2006a) Crystal packing of plant type L-asaparaginase from Escherichia coli. Acta Crystallographica D64, 309-320.
121. Michalska, K., Bujacz, G. & Jaskolski, M. (2006b) Crystal structure of plant asparaginase. Journal of Molecular Biology 360, 105-116.
122. Michalska, K., Hernandez-Santoyo, A. & Jaskolski, M. (2008) The Mechanism of autocatalytic activation of plant-type L-asparaginases. Journal of Biological Chemistry 283, 13388-13397.
123. Mickelson, S., See, D., Meyer, F.D., Garner, J.P., et al. (2003) Mapping of QTL associated with nitrogen storage and remobilization in barley (Hordeum vulgare L.) leaves. Journal of Experimental Botany 54, 801-812.
124. Miflin, B.J. & Habash, D.Z. (2002) The role of glutamine synthetase and glutamate dehydrogenase in nitrogen assimilation and possibilities for improvement in the nitrogen utilization of crops. Journal of Experimental Botany 53, 979-987.
125. Miflin, B.J. & Lea, P.J. (1976) Pathway of nitrogen assimilation in plants. Phytochemistry 15, 873-885.
126. Migge, A., Carrayol, E., Hirel, B., et al. (2000) Leaf-specific overexpression of plastidic glutamine synthetase stimulates the growth of transgenic tobacco seedlings. Planta 210, 252-260.
127. Miyashita, Y. & Good, A.G. (2008) NAD(H)-dependent glutamate dehydrogenase is essential for the survival of Arabidopsis thaliana during dark-induced carbon starvation. Journal of Experimental Botany 59, 667-680.
128. Møller, M.G., Taylor, C., Rasmussen S.K., et al. (2003) Molecular cloning and characterisation of two genes encoding asparagine synthetase in barley (Hordeum vulgare L.). Biochimica et Biophysica Acta 1628, 123-132.
129. Moorhead, G., Douglas, P., Cotelle, V., et al. (1999) Phosphorylation-dependent interactions between enzymes of plant metabolism and 14-3-3 proteins. Plant Journal 18, 1-12.
130. Morey, K.J., Ortega, J.L. & Sengupta-Gopalan, C. (2002) Cytosolic glutamine synthetase in soybean is encoded by a multigene family, and the members are regulated in an organ-specific and developmental manner. Plant Physiology 128, 182-193.
131. Mottram, D.S., Wedzicha, B.L., Dodson, A.T. (2002) Acrylamide is formed in the Maillard reaction. Nature 419, 448-449.
132. Mouilleron, S. & Golinelli-Pimpaneau, B. (2007) Conformational changes in ammoniachanneling glutamine amidotransferases. Current Opinion in Structural Biology 17, 653-664.
133. Muhitch, M.J. (2003) Distribution of the glutamine synthetase isozymeGS(p1) inmaize (Zea mays). Journal of Plant Physiology 160, 601-605.
134. Murray D.R. & Kennedy, I.R. (1980) Changes in the activity of enzymes of nitrogen metabolism in seed coats and cotyledons during embryo development in pea seeds. Plant Physiology 66, 782-785.
135. Myashita, Y. & Good, A.G. (2008) NAD(H)-glutamate dehydrogenase is essential for the survival of Arabidopsis thaliana during dark induced carbon starvation. Journal of Experimental Botany 59, 667-680.
136. Oaks, A. & Ross, D.W. (1984) Asparagine synthetase in Zea mays. Canadian Journal of Botany 62, 68-73.
137. Obara, M., Kajiura, M., Fukuta, Y., et al. (2001) Mapping of QTLs associated with cytosolic glutamine synthetase andNADH-glutamate synthase in rice (Oryza sativa L.). Journal of Experimental Botany 52, 1209-1217.
138. Obara, M., Sato, T., Sasaki, S., et al. (2004) Identification and characterization of a QTL on chromosome 2 for cytosolic glutamine synthetase content and panicle number in rice. Theoretical and Applied Genetics 110, 1-11.
139. Oliveira, I.C., Brears, T., Knight, T.J., et al. (2002) Overexpression of cytosolic glutamine synthetase. Relation to nitrogen, light, and photorespiration. Plant Physiology 129, 1170-1180.
140. Ortega, J.L., Temple, S.J. & Sengupta-Gopalan, C. (2001) Constitutive overexpression of cytosolic glutamine synthetase (GS(1)) gene in transgenic alfalfa demonstrates that GS(1) may be regulated at the level of RNA stability and protein turnover. Plant Physiology 126, 109-121.
141. Osuna, D., Gálvez-Valdivieso, G., Piedras, P., et al. (2001) Cloning, characterization andmRNA expression analysis of PVAS1, a type I asparagine synthetase gene from Phaseolus vulgaris. Planta 213, 402-410.
142. Paczek, V., Dubois, F., Sangwan, R., et al. (2002) Cellular and subcellular localization of glutamine synthetase and glutamate dehydrogenase in grapes, gives new insights on the regulation of C and N metabolism. Planta 216, 245-254.
143. Pageau, K., Reisdorf-Cren, M., Morot-Gaudry, J-F., et al. (2006) The two senescencerelated markers, GS1 (cytosolic glutamine synthetase) and GDH (glutamate dehydrogenase), involved in nitrogen mobilization, are differentially regulated during pathogen attack and by stress hormones and reactive oxygen species in Nicotiana tabacum L. leaves. Journal of Experimental Botany 57, 547-557.
144. Peat, L.J. & Tobin, A.K. (1996) The effect of nitrogen nutrition on the cellular localisation of glutamine synthetase isoforms in barley roots. Plant Physiology 111, 1109-1117.
145. Peterman, T.K. & Goodman, H.M. (1991) The glutamine synthetase gene family of Arabidopsis thaliana: light regulation and differential expression in leaves, roots and seeds. Molecular and General Genetics 230, 145-154.
146. Pryor, A. (1990) A maize glutamic dehydrogenase null mutant is cold temperature sensitive. Maydica 35, 367-372.
147. Purnell, M.P. & Botella, J.R. (2007) Tobacco isoenzyme 1 of NAD(H)-dependent glutamate dehydrogenase catabolizes glutamate in vivo. Plant Physiology 143, 530-539.
148. Purnell, M.P., Skopelitis, D.S., Roubelakis-Angelakis, K.A., et al. (2005) Modulation of higher-plant NAD(H)-dependent glutamate dehydrogenase activity in transgenic tobacco via alteration of beta subunit levels. Planta 222, 167-180.
149. Rastogi, R., Chourey, P.S. & Muhitch, M.J. (1998) The maize glutamine synthetase GS(1-2) gene is preferentially expressed in kernel pedicels and is developmentallyregulated. Plant and Cell Physiology 39, 443-446.
150. Restivo, F.M. (2004) Molecular cloning of glutamate dehydrogenase genes of Nicotiana plumbaginifolia: structure analysis and regulation of their expression by physiological conditions. Plant Science 166, 971-982.
151. Rhodes, D., Rich, P.J. & Brunk, D.G. (1989) Amino acid metabolism of Lemna minor L.: IV. 15N-Labeling kinetics of the amide and amino groups of glutamine and asparagine. Plant Physiology 89, 1161-1171.
152. Ribarits, A., Mamun, A.N.K., Li, S.P., et al. (2007) Combination of reversible male sterility and doubled haploid production by targeted inactivation of cytoplasmic glutamine synthetase in developing anthers and pollen. Plant Biotechnology Journal 5, 483-494.
153. Richards, N.G.J. & Kilberg, M.S. (2006) Asparagine synthetase chemotherapy. Annual Review of Biochemistry 76, 629-654.
154. Riedel, J., Tischner, R. & Máck, R. (2001) The chloroplastic glutamine synthetase (GS-2) of tobacco is phosphorylated and associated with 14-3-3 proteins inside the chloroplast. Planta 213, 396-401.
155. Sakakibara, H., Kawabata, S., Takahashi, H., et al. (1992) Molecular-cloning of the family of glutamine-synthetase genes from maize - expression of genes for glutaminesynthetase and ferredoxin-dependent glutamate synthase in photosynthetic and nonphotosynthetic tissues. Plant and Cell Physiology 33, 49-58.
156. Sakakibara, H., Fujii, K. & Sugiyama, T. (1995) Isolation and characterization of a cDNA that encodes maize glutamate-dehydrogenase. Plant and Cell Physiology 36, 789-797.
157. Sakakibara, H., Shimizu, H., Hase, T., et al. (1996) Molecular identification and characterization of cytosolic isoforms of glutamine synthetase in maize roots. Journal of Biological Chemistry 271, 29561-29568.
158. Sakurai, N., Hayakawa, T., Nakamura, T., et al. (1996) Changes in the cellular localization of cytosolic glutamine synthetase protein in vascular bundles of rice leaves at various stages of development. Planta 200, 306-311.
159. Schmid, M., Davison, T.S., Henz, S.R., et al. (2005)Agene expression map of Arabidopsis thaliana development. Nature Genetics 37, 501-506.
160. Schoenbeck, M.A., Temple, S.J., Trepp, G.B., et al. (2000) Decreased NADH glutamate synthase activity in nodules and flowers of alfalfa (Medicago sativa L.) transformed with an antisense glutamate synthase transgene. Journal of Experimental Botany 51, 29-39.
161. Seiffert, B., Zhou, Z.W., Wallbraun, M., et al. (2004) Expression of a bacterial asparagine synthetase gene in oilseed rape (Brassica napus) and its effect on traits related to nitrogen efficiency. Physiologia Plantarum 121, 656-665.
162. Shi, L., Twary, S.N., Yoshioka, H., et al. (1997) Nitrogen assimilation in alfalfa: isolation and characterization of an asparagine synthetase gene showing enhanced expression in root nodules and dark-adapted leaves. Plant Cell 9, 1339-1356.
163. Shimizu, T., Matsuoka, Y. & Shirasawa, T. (2005) Biological significance of isoaspartate and its repair system. Biological and Pharmaceutical Bulletin 28, 1590.
164. Shimojima, M., Hoffmann-Benning, S., Garavito, R.M., et al. (2005) Ferredoxindependent glutamate synthase moonlights in plant sulfolipid biosynthsesis by forming a complex with SQD1. Archives of Biochemistry and Biophysics 436, 206-214.
165. Sieciechowicz, K.A., Joy, K.W. & Ireland, R.J. (1988) The metabolism of asparagine in plants. Phytochemistry 27, 663-671.
166. Silvente, S., Reddy, P.M., Khandual, S., et al. (2008) Evidence for sugar signalling in the regulation of asparagine synthetase gene expressed in Phaseolus vulgaris roots and nodules. Journal of Experimental Botany 59, 1279-1294.
167. Skopelitis, D.S., Paranychianakis, N.V., Paschalidis K.A., et al. (2006) Abiotic stress generates ROS that signal expression of anionic glutamate dehydrogenases to form glutamate for proline synthesis in tobacco and grapevine. Plant Cell 18, 2767-2781.
168. Skopelitis, D.S., Paranychianakis, N.V., Kouvarakis, A., et al. (2007) The isoenzyme 7 of tobacco NAD(H)-dependent glutamate dehydrogenase exhibits high deaminating and low aminating activities in vivo. Plant Physiology 145, 1726-1734.
169. Sodek, L. & Lea, P.J. (1993) Asparaginase from the testa of developing lupin and pea seeds. Phytochemistry 34, 51-56.
170. Sodek, L., Lea, P.J. & Miflin, B.J. (1980) Distribution and properties of a potassiumdependent asparaginase isolated from developing seeds of Pisum sativum and other plants. Plant Physiology 65, 22-26.
171. Somerville, C.R. (1986) Analysis of photosynthesis with mutants of higher-plants and algae. Annual Review of Plant Physiology and Plant Molecular Biology 37, 467-507.
172. Somerville, C.R. & Ogren, W.L. (1980) Inhibition of photosynthesis in Arabidopsis mutants lacking leaf glutamate synthase activity. Nature 286, 257-259.
173. Stewart, G.R., Shatilov, V.R., Turnbull, M.H., et al. (1995) Evidence that glutamatedehydrogenase plays a role in the oxidative deamination of glutamate in seedlings of Zea mays. Australian Journal of Plant Physiology 22, 805-809.
174. Suzuki, A. & Knaff, D.B. (2005) Glutamate synthase: structural, mechanistic and regulatory properties, and role in the amino acid metabolism. Photosynthesis Research 83, 191-217.
175. Swarup, R., Bennett, M.J. & Cullimore, J.V. (1991) Expression of glutamine-synthetase genes in cotyledons of germinating Phaseolus vulgaris L. Planta 183, 51-56.
176. Syntichaki, K.M., Loulakakis, K.A. & Roubelakisangelakis, K.A. (1996) The aminoacid sequence similarity of plant glutamate dehydrogenase to the extremophilic archaeal enzyme conforms to its stress-related function. Gene 168, 87-92.
177. Ta, T-C., Faris, M.A. & Macdowall, F.D.H. (1986) Pathways of nitrogen metabolism in nodules of alfalfa (Medicago sativa L.). Plant Physiology 80, 1002-1005.
178. Tabuchi, M., Sugiyama, K., Ishiyama, K., et al. (2005) Severe reduction in growth rate and grain filling of rice mutants lacking OsGS1;1, a cytosolic glutamine synthetase1; 1. Plant Journal 42, 641-651.
179. Tabuchi, M., Abiko, T. & Yamaya, T. (2007) Assimilation of ammonium-ions and reutilization of nitrogen in rice (Oryza sativa L.). Journal of Experimental Botany 58, 2319-2327.
180. Taira, M., Valtersson, U., Burkhardt, B., et al. (2004) Arabidopsis thaliana GLN2-encoded glutamine synthetase is dual targeted to leaf mitochondria and chloroplasts. Plant Cell 16, 2048-2058.
181. Tanksley, S.D. (1993) Mapping polygenes. Annual Review of Genetics 27, 205-233.
182. Tercé-Laforgue, T., Dubois, F., Ferrario-Méry, S., et al. (2004a). Glutamate dehydrogenase of tobacco is mainly induced in the cytosol of phloem companion cells when ammonia is provided either externally or released during photorespiration. Plant Physiology 136, 4308-4317.
183. Tercé-Laforgue, T., Máck, G., Hirel, B. (2004b) New insights towards the function of glutamate dehydrogenase revealed during source-sink transition of tobacco (Nicotiana tabacum) plants grown under different nitrogen regimes. Physiologia Plantarum 120, 220-228.
184. Thum, K.E., Shasha, D.E., Lejay, L.V., et al. (2003) Light- and carbon-signaling pathways: modeling circuits of interactions. Plant Physiology 132, 440-452.
185. Thurman, D.A., Palin, C. & Laycock, M.V. (1965) Isoenzymatic nature of L-glutamate dehydrogenase of higher plants. Nature 207, 193-194.
186. Tingey, S.V., Tsai, F.Y., Edwards, J.W., et al. (1988) Chloroplast and cytosolic glutaminesynthetase are encoded by homologous nuclear genes which are differentially expressed in vivo. Journal of Biological Chemistry 263, 9651-9657.
187. Tobin, A.K. & Yamaya, T. (2001) Cellular compartmentation of ammonium assimilation in rice and barley. Journal of Experimental Botany 52, 591-604.
188. Tonin, G.S. & Sodek, L. (1990) Asparaginase, allantoinase and glutamine synthetase in soybean cotyledons grown in vitro. Phytochemistry 29, 2829-2831.
189. Trepp, G.B., Van De Mortel, M., Yoshioka, H., et al. (1999) NADH-glutamate synthase in alfalfa root nodules. Genetic regulation and cellular expression. Plant Physiology 119, 817-828.
190. Tsai, F-Y. & Coruzzi, G.M. (1990) Dark-induced and organ-specific expression of two asparagine synthetase genes in Pisum sativum. EMBO Journal 9, 323-332.
191. Tsai, F-Y. & Coruzzi, G.M. (1991) Light represses transcription of asparagine synthetase genes in photosynthetic and non-photosynthetic organs of plants. Mo1ecular Cell Biology 11, 4966-4972.
192. Turano, F.J., Thakkar, S.S., Fang, T., et al. (1997) Characterisation and expression of NAD(H)-dependent glutamate dehydrogenase genes in Arabidopsis. Plant Physiology 113, 1329-1341.
193. Unno, H., Uchida, T., Sugawara, H., et al. (2006) Atomic structure of plant glutamine synthetase: a key enzyme for plant productivity. Journal of Biological Chemistry 281, 29287-29296.
194. van den Heuvel, R.H.H., Curti, B., Vanoni, M.A., et al. (2004) Glutamate synthase a fascinating pathway from glutamine to glutamate. Cellular and Molecular Life Sciences 61, 669-681.
195. 2 fixation. Plant Journal 8, 345-358.
196. Vanoni, M.A. & Curti, B. (2008) Structure-function studies of glutamate synthases: a class of self-regulated iron-sulfur flavoenzymes essential for nitrogen assimilation. IUBMB Life 60, 287-300.
197. Vanoni, M.A., Dosenna, L., van den Heuvel, R.H.H., et al. (2005) Structure function studies on the complex iron sulfur flavoprotein glutamate synthase: the key enzyme of ammonia assimilation. Photosynthesis Research 83, 219-238.
198. Vincent, R., Fraisier, V., Chaillou, S., et al. (1997) Overexpression of a soybean gene encoding cytosolic glutamine synthetase in shoots of transgenic Lotus corniculatus L plants triggers changes in ammonium assimilation and plant development. Planta 201, 424-433.
199. Wallsgrove, R.M., Lea, P.J. & Miflin, B.J. (1979) Distribution of the enzymes of nitrogen assimilation within the pea leaf cell. Plant Physiology 63, 233-236.
200. Wallsgrove, R.M., Keys, A.J., Bird, I.F., et al. (1980) The location of glutamine-synthetase in leaf-cells and its role in the reassimilation of ammonia released in photorespiration. Journal of Experimental Botany 31, 1005-1017.
201. Wallsgrove, R.M., Turner, J.C., Hall, N.P., et al. (1987) Barley mutants lacking chloroplast glutamine-synthetase - biochemical and genetic-analysis. Plant Physiology 83, 155-158.
202. Winichayakul, S., Moyle R.L., Coupe, S.A., et al. (2004a) Analysis of the asparagus (Asparagus officinalis) asparagine synthetase gene promoter identifies evolutionarily conserved cis-regulatory elements that mediate Suc-repression. Functional Plant Biology 31, 63-72.
203. Winichayakul, S., Moyle, R.L., Ryan, D.J., et al. (2004b) Distinct cis-elements in the Asparagus officinalis asparagine synthetase promoter respond to carbohydrate and senescence signals. Functional Plant Biology 31, 573-582.
204. Yamaya, T., Obara, M., Nakajima, H., et al. (2002) Genetic manipulation and quantitative-trait locimapping for nitrogen recycling in rice. Journal of Experimental Botany 53, 917-925.
205. Yamada, K., Lim, J., Dale, J.M., et al. (2003) Empirical analysis of transcriptional activity in the Arabidopsis genome. Science 302, 842-846.