Profile – Bertrand Hirel


Bertrand Hirel

Job Title:

Director of Research CNRS


Phone Number:

+33 1 30 83 30 89



1977-1978          BSc. Université of Nancy I (France)

1978-1980          PhD thesis. Université de Nancy I (France)

1980-1984          Doctor of Science Thesis, University of Paris-Sud, Orsay (France).


1981 – 1984          Attaché de Recherche (Scientific Officer) Centre National de la Recherche Scientifique (CNRS), Université of Paris-Sud, Orsay, France.

1984 – 1990          Chargé de Recherche CNRS 1ère Classe (Principal Scientific Officer), Laboratoire du Métabolisme, INRA Versailles, France.

1990                      Directeur de Recherches CNRS 2ème classe (Senior Scientific Officer). Laboratoire du Métabolisme, INRA Versailles, France.

1996 – 1998          Associate Director. Unité de Nutrition Azotée des Plantes. INRA, Versailles, France.

2001- 2002          University of Paris XI, Orsay (France). Associate Professor

1998 – 2010          Directeur de Recherches CNRS 2ème classe (Senior Scientific Officer). Group leader.

2010 – present    Directeur de Recherches CNRS 1ère Classe (Chief Senior Scientific Officer). Group Leader.


Our overall goal is to better understand and improve nitrogen (N) use efficiency (NUE: Nitrogen Use Efficiency) in maize and wheat. Indeed, these two plants, which are with rice the base for food and feed in the world requires significant inputs of N fertilizers (nitrates in particular) for optimum performance. The improvement of NUE is a major issue, considering both the growing demand for agricultural products of the world population, the rising cost of fossil fuels needed to produce N fertilizers and water pollution caused by the application in excess of nitrates to fertilize cultivated soils. Therefore, it is necessary to breed new varieties of maize and wheat that require less N fertilizer (20 to 30% less), but that are able to maintain both high yield and good grain quality (content proteins, in particular).

My research has been to advance knowledge on the regulation of N assimilation in plants with a constant focus on innovation and agricultural applications taking into account the current constraints related to the environment and the development of a sustainable agriculture.


Research Areas:

  • Nitrogen use efficiency

Current N-related Projects:

  • Improving maize nitrogen use efficiency


Peer review articles:

1 – Guiz C., Hirel B., Schedlofsky G., Gadal P., 1979 – Occurrence and influence of light on the relative proportions of two glutamine synthetase in rice leaves. Plant Sci. Let. 15 : 272-277.

2 – Hirel B., Gadal P. 1980- Sur la localisation intracellulaire de deux formes isofonctionnelles de la glutamine synthétase chez le Riz. C.R. Acad. Sci., Paris 291 D : 441-444.

3 – Hirel B., Gadal P., 1980 – Glutamine synthetase in rice. A comparative study of the enzymes from roots and leaves. Plant Physiol. 66 : 619-623.

4 – Hirel B., Gadal P., 1981 – Glutamine synthetase isoforms in pea leaves. Z. Pflanzenphysiol. 102 : 315-319.

5 – Hirel B., Gadal P., 1982  – Glutamine synthetase isoforms in leaves of a C4 plant : Sorghum vulgare  L. Physiol. Plant. 54 : 69-74.

6 – Hirel B., Perrot-Rechenmann C., Suzuki A., Vidal J., Gadal P., 1982 – Glutamine synthetase in spinach leaves. Immunological studies and immuno-cytochemical localization. Plant Physiol. 69 : 983-987.

7 – Hirel B., Perrot-Rechenmann C., Maudinas B., Gadal P., 1982b – Glutamine synthetase in alder (Alnus glutinosa) root nodules. Purification, properties and cytoimmunochemical localization. Physiol. Plant. 55 : 197-200.

8 – Hirel B., Vidal J., Gadal P., 1982 – Evidence for a cytosolic dependent light induction of chloroplastic glutamine synthetase in etiolated rice leaves. Planta 155 : 17-23.

9 – Martin F., Suzuki A., Hirel B., 1982 – A new high performance liquid chromatography assay for glutamine synthetase and glutamate synthase in plant tissue. Anal. Biochem. 125 : 24-29.

10 – McNally S.F., Hirel B., Gadal P., Mann A.F., Stewart G.R., 1983 – Glutamine synthetase in higher plants. Evidence for a specific isoform content related to their possible physiological role and their compartmentation within the leaf. Plant Physiol. 72 : 22-25.

11 – McNally S.F., Orejambo T.O., Hirel B., Stewart G.R., 1983 – Glutamine synthetase isoenzymes of Striga hermontica  and other angiosperm root parasites. J. Exp. Bot. 94 : 47-56.

12 – Martin F., Hirel B., Gadal P., 1983 – Sur l’activité enzymatique ornithine carbamoyl transferase des actinorhizes d’Alnus glutinosa  Gaetn. C.R. Acad. Sci., Paris, série III 295 : 557-559.

13 – Martin F., Hirel B., Gadal P., 1983 – Purification and properties of ornithine carbamyol transferase from Alnus glutinosa  root nodules. Z. Pflanzenphysiol. 5 : 413-422.

14 – McNally S.F., Hirel B., Stewart G.R., 1983 – Nitrogen metabolism of halophytes. IV. The occurrence of multiple forms of glutamine synthetase in leaf tissue. New Phytol. 34 : 610-619.

15 – McNally S.F., Hirel B., 1983 – Glutamine synthetase isoforms in higher plants. Physiol. Vég. 21 : 761-774.

16 – Hirel B., Lavergne D., McNally S.F., Gadal P., 1983 – Some regulatory properties of glutamine synthetase isoforms in a C4 plant Pennisetum americanum  L. (Leeke). Plant Sci. Lett. 111 : 413-422.

17 – Hirel B., McCashin B., Layzell D.B., McNally S.F., Canvin D.T., 1983 – Isoforms of glutamine synthetase having C3, C4 and intermediate photosynthetic pathways. Can. J. Bot. 61 : 2257-2259.

18 – Ferrario S., Hirel B., Gadal P., 1983 – Immunochemical characterization of nitrate reductase from spinach leaves and roots. Biochem. Biophys. Res. Comm. 113 : 733-737.

19 – Hirel B., McNally S.F., Gadal P., Sumar N., Stewart G.R., 1984 – Cytosolic glutamine synthetase in higher plants. A comparative immunological study. Eur. J. Biochem. 138 : 63-66.

20 – Hirel B., Weatherley C., Cretin C., Bergounioux C., Gadal P., 1984 – Multiple subunit composition of chloroplastic glutamine synthetase of Nicotiana tabacum  L. Plant Physiol. 74 : 448-450.

21 – Hirel B., Nato A., Martin F., 1984c – Glutamine synthetase in ribulose-1,5 bisphosphate carboxylase deficient tobacco mutants in cell suspension culture. Plant Cell Reports 3 : 106-108.

22 – Nato F., Hirel B., Nato A., Gadal P., 1984 – Chloroplastic glutamine synthetase from tobacco leaves. A glycosylated protein. FEBS Lett. 175 : 443-446.

23 – Oaks A., Hirel B., 1985. Nitrogen assimilation in roots. Ann. Rev. Plant Physiol. 36 : 345-365.

24 – Hoarau J., Hirel B., Nato A., 1986 – A new artificial electron donor for nitrate reductase of higher plants and other organisms. Plant Physiol. 80 : 946-949.

25 – Hirel B., Bouet C., King B., Layzell D., Jacobs F., Verma D.P.S., 1987 – Glutamine synthetase genes are regulated by ammonia provided externally or by symbiotic nitrogen fixation. EMBO J. 5 : 1167-1171.

26 – Thalouran P., Rey L., Hirel B., Renaudin S., Fer A., 1987 – Activity and immunocytochemical localization of glutamine synthetase in Lathrea clandestina; Protoplasma 141: 95-100.

27 – Brangeon J., Hirel B., Forchioni A., 1989 – Immunogold localization of glutamine synthetase  in soybean leaves, roots and nodules. Protoplasma 151 : 88-97.

28 – Miao G.H., Hirel B., Marsolier M.C., Ridge R.W.,Verma D.P.S., 1991 – Ammonia regulated expression of a soybean gene encoding cytosolic glutamine synthetase in transgenic Lotus corniculatus. The Plant Cell 3 : 11-22.

29 – Hoarau J., Nato A., Lavergne D., Flipo V., Hirel B., 1991. – Nitrate reductase activity during a culture cycle of tobacco cells : the participation of a membrane bound form enzyme. Plant Sci. 79 : 192-204.

30 – Zehnacker C., Becker T.W., Suzuki A., Carrayol E., Caboche M., Hirel B., 1992. – Purification and properties of tobacco ferredoxin-dependent glutamate synthase and isolation of corresponding cDNA clones. Light inducibility and organ-specificiy in the gene transcription and protein expression. Planta 187 : 266-274.

31 – Becker T.W., Caboche M., Carrayol E., Hirel B., 1992. – Nucleotide sequence of a tobacco cDNA encoding plastidic glutamine synthetase and light-inducibility, organ specificity and diurnal rhytmicity in the expression of the corresponding genes of tobacco and tomato. Plant Mol. Biol. 19 : 367-379.

32 – Hirel, B., Marsolier M.C., Hoarau A., Hoarau J., Brangeon J., Schafer R., Verma D.P.S., 1992. – Forcing expression of a soybean root glutamine synthetase gene in tobacco leaves induces a native gene encoding cytosolic enzyme. Plant Mol. Biol. 20 : 207-218.

33 – Marsolier, M.C., Carrayol, E., Hirel, B., 1993. – Multiple functions of promoter sequences involved in ammonia regulation and organ-specific expression of a soybean glutamine synthetase gene in transgenic Lotus corniculatus. The Plant Journal 3 : 405-414.

34 – Becker, T.W., Perrot-Rechenman, C., Suzuki, A., Hirel, B., 1993. – Subcellular and immunocytochemical localization of the enzymes involved in ammonia assimilation in mesophyll and bundle sheath strands of maize leaves. Planta 191 : 129-136.

35 – Becker, T.W., Nef, C., Zehnacker, C., Hirel, B., 1993. – Implication of the phytochrome in light regulation of the tomato gene(s) encoding ferredoxin-dependent glutamate synthase. Plant Physiol. Biochem. 31 : 725-729.

36- Marsolier, M.C., Hirel, B., 1993. – Metabolic and developmental control of soybean cytosolic glutamine synthetase genes expression. Proceedings of the Third international Symposium on Inorganic Nitrogen Assimilation. Physiol. Plant. 89: 613-617.

37- Marsolier, M.C., Debrosses, G., Hirel, B. 1994. – Identification of several soybean glutamine syntetase transcripts highly or specifically expressed in nodules. Expression studies of the corresponding genes in trangenic Lotus corniculatus. Plant Mol. Biol. 27: 1-15.

38- Pérez-Garcia, A., Canovas, F.M., Gallardo, F., Hirel, B., de Vicente, A. 1995. – Differential expression of glutamine synthetase isoforms in detached leaflets infected with Pseudomonas syringae pv.tomato. Mol. Plant Microbe Interac. 8 : 96-103.

39 – Dubois F., Brugière N., Sangwan R.S., Hirel B., 1996. – Localization of tobacco cytosolic glutamine synthetase enzymes and the corresponding transcripts shows organ- and cell-specific patterns of protein synthesis and gene expression. Plant Mol. Biol. 31: 803-817.

40 – Migge A., Meya G., Carrayol E., Hirel B., Becker T.W., 1996. – Regulation of the subunit composition of tomato plastidic glutamine synthetase by light and the nitrogen source. Planta 200: 213-220.

41 – Schnorr K.M., Laloue M., Hirel B., 1996. – Isolation of cDNAs encoding two purine biosynthetic enzymes of soybean and expression of the corresponding transcripts in roots and root nodules. Plant Mol. Biol. 32: 751-757.

42 – Bauer D., Biehler K., Fock H., Carrayol E., Hirel B., Migge A., Becker T., 1997. – A role for cytosolic glutamine synthetase in the remobilization of leaf nitrogen during water stress in tomato. Physiol. Plant. 99: 241-248.

43 – Carrayol E., Tercé-Laforgue T., Desbrosses G., Pruvot-Maschio G., Poirier S., Ratet P., Hirel B., 1997. – Ammonia regulated expression of a soybean gene encoding cytosolic glutamine synthetase is not conserved in two heterologous plant systems. Plant Sci. 125: 75-85.

44 – Hirel B., Phillipson B., Murchie E., Suzuki A., Kunz C., Ferrario S. Limami A., Chaillou S., Deléens E., Brugière N., Chaumont-Bonnet M., Foyer C.H., Morot-Gaudry J.F., 1997. – Manipulating the pathway of ammonia assimilation in transgenic non-legumes and legumes. J. Plant Nutr. Soil Sci. 160: 283-290.

45 – Migge A., Carrayol E., Kunz C., Hirel B., Fock H., Becker T., 1997. – The expression of the tobacco genes encoding plastidic glutamine synthetase or ferredoxin-dependent glutamate synthase does not depend on the rate of nitrate reduction and is unaffected by supression of photorespiration. J. Exp. Bot. 48: 1175-1184.

46 – Vincent R., Fraisier V., Chaillou S., Limami M.A., Deléens E., Phillipson B., Douat C., Boutin J.P., Hirel B., 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.

47 – Migge A., Carrayol E., Hirel B., Lohmann M., Meya G., Becker T.W., 1998. – Regulation of the subunit composition of plastidic glutamine synthetase of the wild-type and of the phytochrome-deficient aurea mutant of tomato by blue/UV-A- or by UV-B-light. Plant Mol. Biol. 37: 689-700.

48 – Migge A., Carrayol E., Hirel B., Lohmann M., Meya G., Becker T.W., 1998. – Two nitrite reductase isoforms are present in tomato cotyledons and are regulated differently by UV-A or UV-B light and during plant development. Planta 207:  229-234.

49 – Migge A., Carrayol E., Hirel B., Lohmann M., Meya G., Becker T.W., 1998. – Influence of UV-A or UV-B light and of nitrogen source on the induction of ferredoxin-dependent glutamate synthase in etiolated tomato cotyledons. Plant Physiol. Biochem. 36: 789-797.

50 – Tercé-Laforgue T., Carrayol E., Cren M., Desbrosses G., Hecht V., Hirel, B., 1999. – A strong constitutive positive element is essential for the ammonium-regulated expression of a soybean gene encoding cytosolic glutamine synthetase. Plant Mol. Biol. 39, 551-564.

51 – Ruiz M.T., Prosser I., Hirel B., Clarckson D.T. (1999) – A cDNA sequence (Accession No. X94299) encoding a cytosolic subunit of glutamine synthetase in the model legume Lotus japonicus. (PGR99-031). Plant Physiol. 119: 1148.

52 – Lancien M., Ferrario-Méry S., Roux Y., Bismuth E., Masclaux C., Hirel B., Gadal P., Hodges M., 1999. – Simultaneous expression of NAD-dependent isocitrate dehydrogenase and other Krebs cycle genes after nitrate resupply to short-term nitrogen starved Nicotiana tabacum. Plant Physiol. 120: 717-725.

53 – Limami A., Phillipson B., Ameziane R., Pernollet N., Jiang Q., Roy R., Deleens E., Chaumont-Bonnet M., Gresshoff P.M., Hirel B., 1999. – Does root glutamine synthetase control plant biomass production in Lotus japonicus L. Planta  209, 495-502.

54 – Brugière N., Dubois F., Limami A., Lelandais M., Roux Y., Sangwan R., Hirel B. 1999. – Glutamine synthetase in the phloem plays a major role in controlling proline production. The Plant Cell 11: 1995-2011.

55 – Cren M., Hirel B. 1999. – Glutamine synthetase in higher plants: regulation of gene and protein expression from the organ to the cell. Plant Cell Physiol. 40: 1187-1193.

56 – Migge A., Carrayol E., Hirel B., Becker T.W. 2000. – Leaf-specific overexpression of plastidic glutamine synthetase stimulates the growth of transgenic tobacco seedlings. Planta  2: 252-260.

57 – Harrison J., Brugière N., Phillipson B., Ferrario-Méry S., Becker T., Limami A., Hirel B. 2000. – Manipulating the pathway of ammonia assimilation through genetic manipulation and breeding. Consequences on plant physiology and development. Plant and Soil 221: 81-93.

58 – Ferrario-Méry S., Suzuki A., Kunz C., Valadier M.H., Roux Y., Hirel B., Foyer C. 2000. – Modulation of amino acid metabolism in transformed tobacco plants deficient in Fd-GOGAT. Plant and Soil 221: 67-79.

59 – Masclaux C., Valadier M.H., Brugière N., Morot-Gaudry J.F., Hirel B. 2000. – Characterization of the sink/source transition in tobacco (Nicotiana tabacum L.) shoots in relation to nitrogen management and leaf senescence. Planta 211: 510-518.

60 – Brugière N., Dubois F., Masclaux C., Sangwan R., Hirel B. 2000. – Immunolocalization of glutamine synthetase in senescing tobacco (Nicotiana tabacum L.) leaves suggests that ammonia assimilation is progressively shifted to the mesophyll cytosol. Planta  211: 519-527.

61 – Becker T.W., Carrayol E., Hirel B. 2000. – Glutamine synthetase and glutamate dehydrogenase isoforms in maize leaves: Localization, relative proportion and their role in ammonium assimilation or nitrogen transport. Planta 211: 800-806.

62 – Ferrario-Méry S., Masclaux C., Suzuki A., Valadier M.H., Hirel B., Foyer C.H. 2001. – Glutamine and a-ketoglutarate are metabolite signals involved in nitrate reductase gene transcription in untransformed and transformed tobacco plants deficient in ferredoxin-glutamine-a-ketoglutarate aminotransferase. Planta 213: 265-271.

63 – Masclaux C., Quilleré I., Gallais A., Hirel B. 2001. – The challenge of remobilisation in plant nitrogen economy. A survey of physio-agronomic and molecular approaches. Annals of Apl. Biol. 138: 69-81.

64 – Hirel B., Bertin P., Quillere I., Bourdoncle W., Attagnant C., Dellay C., Gouy A., Cadiou S., Retailliau C., Falque M., Gallais A., 2001. – Towards a better understanding of the genetic and physiological basis for nitrogen use efficiency in maize. Plant Physiol. 125: 1258-1270.

65 – Clément A., Prévôt J.C., Pujade-Renaud V., Hirel B., Jacob J.L. 2001. – Glutamine synthetase of Hevea brasiliensis latex: characterisation and physiological regulation. J. Rubb. Res. 4: 56-68.

66 – Ferrario-Mery S., Hodges M., Hirel B., Foyer C.H. 2001. – Photorespiration-dependent increases in phophoenolpyruvatecarboxylase, isocitrate dehydrogenase and glutamate dehydrogenase in transformed tobacco plants deficient in ferredoxin-dependent glutamine-a-ketoglutarate amino transferase. Planta 214: 877-886.

67 – Paczek V., Dubois F., Sangwan R., Morot-Gaudry J.F., Roubelakis-Angelakis K.A., Hirel B.  2002. – Cellular and subcellular localisation of glutamine synthetase and glutamate dehydrogenase in grapes give new insights on the regulation of C and N metabolism. Planta 216: 245-254.

68 – Ferrario-Méry S., Valadier M.H., Godefroy N., Miallier D., Hirel, B., Foyer C.H., Suzuki A. 2002. Diurnal changes in ammonia assimilation in transformed tobacco plants expressing ferredoxin-dependent glutamate synthase mRNA in the antisense orientation. Plant Sci. 163: 59-67.

69 – Masclaux-Daubresse C., Valadier M.H., Carrayol E., Reisdorf-Cren M., Hirel B. 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 Env. 25: 1451-1462.

70 – Reisdorf-Cren M., Carrayol E., Tercé-Laforgue  T., Hirel  B. 2002. – A novel HMG A like-protein binds differentially to the AT-rich regions located in the far distal and proximal parts of a soybean glutamine synthetase gene (GS15) promoter. Plant Cell Physiol 43: 1006-1016.

71 – Limami A.M., Rouillon C., Glevarec G., Gallais A., Hirel B. 2002. – Genetic and physiological analysis of germination efficiency in maize in relation to N metabolism reveals the importance of cytosolic glutamine synthetase. Plant Physiol. 130: 1860-1870.

72 – Fei H., Vessey K., Chaillou S., Hirel B., Mahon J. 2002. –  Overexpression of a soybean cytosolic glutamine synthetase gene linked with organ-specific promoters in pea plants grown in different concentration of nitrate. Planta 216: 467-474.

73 – Harrison J., Pou de Crecenzo M.A., Sené O., Hirel B. 2003. –  Does lowering glutamine synthetase activity in nodules modify nitrogen metabolism and growth of Lotus japonicus L.? Plant Physiol. 133: 253-262.

74 – Dubois F., Tercé-Laforgue T., Gonzalez-Moro M.B., Estavillo M.B., Sangwan R.,  Gallais A., Hirel B. 2003 Glutamate dehydrogenase in plants: is there a new story for an old enzyme? Plant Physiol Biochem. 41: 565-576.

75 – Hirel B., Limami A. 2003. – Prospects for improving nitrogen use efficiency: insights given by 15N-labelling experiments. Phytochem Rev.  2: 133-144.

76 – Gallais A., Hirel B. 2004. – An approach of the genetics of nitrogen use efficiency in maize. J Exp. Bot. 396: 295-306.

77 – Harrison J., Hirel B., Limami A. 2004. – Variation in nitrate uptake and assimilation between two ecotypes of Lotus japonicus L. and their recombinant inbred lines. Physiol. Plant. 120: 124-131.

78 – Tercé-Laforgue T., Mäck G., Hirel B. 2004. – New insights towards the function of glutamate dehydrogenase revealed during source-sink transition of tobacco (Nicotiana tabacum L.) plants grown under different nitrogen regimes. Physiol. Plant. 120: 220-228.

79 – Tercé-Laforgue T., Dubois F., Ferrario-Mery S., Pou de Crecenzo M.A., Sangwan R., Hirel B. 2004. – Glutamate dehydrogenase of tobacco (Nicotiana tabacum L.) is mainly induced in the cytosol of phloem companion cells when ammonia is provided either externally or released during photorespiration. Plant Physiol. 136: 4308-4317.

80 – Bahrman N., Gouy A., Devienne-Barret F., Hirel B., Vedele F., LeGouis J. 2004. – Differential changes in root protein patterns of two wheat varieties under high and low nitrogen nutrition levels. Plant Sci. 168: 81-87.

81 – Boisson M., Mondon K., Torney V., Nicot N., Laine A.L., Bahrman N., Gouy A., Daniel-Vedele F., Hirel B., Sourdille P., Dardevet M., Ravel C., LeGouis J. 2005. – Partial sequences of nitrogen metabolism genes in hexaploid wheat. Theor. Appl. Genet.110: 932-940.

82 – Hirel B., Martin A., Tercé-Laforgue T., Gonzalez-Moro M.B., Estavillo J.M., 2005. – Physiology of maize I: A comprehensive and integrated view of nitrogen metabolism in a C4 plant. Physiol. Plant. 124: 167-177.

83 – Hirel B., Andrieu B., Valadier M.H., Renard S., Quilleré I., Chelle M., Pommel B., Fournier C., Drouet J.L. 2005. – Physiology of maize II: Identification of physiological markers representative of the nitrogen status of maize (Zea mays L.) leaves, during grain filling. Physiol. Plant. 124: 178-188.

84 – Kichey T., Le Gouis J., Hirel B., Dubois F. 2005. – Evolution of the cellular and subcellular localization of glutamine synthetase and glutamate dehydrogenase during flag leaf senescence in wheat (Triticum aestivum L.). Plant Cell Physiol. 46: 964-974.

85 – Martin A., Belastegui-Macadam X.,  Quilleré I., Floriot M., Valadier M.H., Pommel B., Andrieu B., Donnison I., Hirel B. 2005. – Nitrogen management and senescence in two maize hybrids differing in the persistence of leaf greenness. Agronomic, physiological and molecular aspects. New Phytol. 167: 483-492.

86 – Kichey T., Heumez E., Pocholle P., Pageau K., Vanacker H., Dubois F., Le Gouis J., Hirel B. 2006. – Combined agronomic and physiological aspects of nitrogen management in wheat (Triticum aestivum L.). Dynamic and integrated views highlighting the central role of the enzyme glutamine synthetase. New Phytol. 169: 265-278.

87 – Pommel B., Gallais A., Coque M., Quilleré I., Hirel B., Prioul J.L., Andrieu B. 2006. – Carbon and Nitrogen allocation and grain filling in three maize hybrids differing in leaf senescence. Eur. J. of Agronom. 24: 203-211.

88 – Hirel B., Gallais A. 2006 – Rubisco synthesis, turnover and degradation: some new thoughts on an old problem. New Phytol. 169: 445-448.

89 – Coque M., Bertin P., Hirel B., Gallais A. 2006. – Genetic variation and QTLs for 15N natural abundance in a set of maize recombinant inbred lines. Field Crop Res. 97 : 310-321.

90 – Fontaine J.X., Saladino F., Agrimonti C., Bedu M., Tercé-Laforgue T., Tétu T., Hirel B., Restivo F.M. Dubois F. 2006. – Control of the Synthesis and of the Subcellular Targeting of the Two GDH Genes Products in Leaves and Stems of Nicotiana plumbaginifolia and Arabidopsis thaliana. Plant Cell Physiol. 47 : 410-418.

91 – Gallais A., Coque M., Quilleré I., Prioul J.L., Hirel B.  2006. Modelling post-silking N-fluxes in maize using 15N-labelling-field experiments. New Phytol. 172 : 696-607.

92 – Martin A., Lee J., Kichey T.,  Gerentes D., Zivy M., Tatou C., Balliau T., Valot B., Davanture M., Dubois F., Tercé-Laforgue T., Coque M., Gallais A., Gonzalez-Moro M.B., Bethencourt L., Quilleré I., Habash D.Z., Lea P.J., Charcosset A., Perez P., Murigneux A., Sakakibara H., Edwards K.J., Hirel B. 2006. Two cytosolic glutamine synthetase isoforms of maize (Zea mays L.) are specifically involved in the control of grain production. The Plant Cell 18 : 3252-3274.

93 – Fei H., Chaillou S., Hirel B., Polowick P., Mahon J., Vessey K. 2006. Effects of the overexpression of a soybean cytosolic glutamine synthetase gene (GS15) linked to organ-specific promoters on growth and nitrogen accumulation of pea plants supplied with ammonium. Plant Physiol. Biochem 44 : 543-550.

94 – Gallais A., Coque M., Quilléré I., Le Gouis J., Prioul J.L., Hirel. B. 2007. Estimating proportions of N remobilization and of post-silking N uptake allocated to maize kernels by 15N labelling. Crop Sci. 47: 685-691.

95 – Kichey T., Hirel B., Heumez E., Dubois F., Le Gouis J. 2007. In winter wheat (Triticum aestivum L.), post anthesis nitrogen uptake and remobilisation to the grain correlates with agronomic traits and nitrogen physiological markers. Field Crop Res. 102 : 22-32.

96 – Hirel B., Chardon F., Durand J. 2007. The contribution of molecular physiology to the improvement of nitrogen use efficiency in crops. J. Crop Sci. Biotech. 10 : 129-136.

97 – Hirel B., Le Gouis J., Ney B., Gallais A. 2007. The challenge of improving nitrogen use efficiency in crop plants : towards a more central role for genetic variability and quantitative genetics within integrated approaches. J Exp. Bot. 58: 2369-2387.

98 – Bernard S.M., Blom Møller A.L., Dionisio G., Jahn T.P., Kichey T., Dubois F., Baudo M., Lopes M.S., Tercé-Laforgue T., Foyer C.H., Parry M., Forde B.G., Araus J.L., Hirel B., Schjoerring J.K., Habash D.Z. 2008 Gene expression, cellular localisation and function of glutamine synthetase isozymes in wheat (Triticum aestivum L.). Plant Mol. Biol. 67 : 89-105.

99 – Coque M., Martin  A., Veyrieras J.B., Hirel B., Gallais A. 2008.  Genetic variation for N-remobilization and postsilking N-uptake in a set of maize recombinant inbred lines. 3. QTL detection and coincidences. Theor. Appl. Genet. 117: 729-747.

100 – Fontaine J.X., Ravel C., Pageau K., Heumez E., Dubois F., Hirel B., Le Gouis J. 2009 A quantitative genetic study for elucidating the contribution of glutamine synthetase, glutamate dehydrogenase and other nitrogen-related physiological traits to the agronomic performance of common wheat. Theor. Appl. Genet. 119: 645-662.

101 – Cañas R.A., Quilleré I., Christ A., Hirel B. 2009. Nitrogen metabolism in the developing ear of maize (Zea mays L.): analysis of two lines contrasting in their mode of nitrogen management. New. Phytol 184: 340-352.

102 – Labboun S., Tercé-Laforgue T., Roscher A., Bedu M., Restivo F.M., Velanis C.N, Skopelitis D.S., Moshou P.N., Roubelakis-Angelakis K.A., Suzuki A., Hirel B. 2009.  Resolving the role of plant glutamate dehydrogenase: I. In vivo real time nuclear magnetic resonance spectroscopy experiments. Plant. Cell. Physiol. 50 : 1761-1773.

103 – Broyard C., Fontaine J.X., Molinie R., Cailleu D., Tercé-Laforgue T., Dubois F., Hirel B., Dubois F., Mesnard F. 2009. Metabolomic profiling of two cytosolic glutmaine synthetase maize mutants (Zea mays L.) using 1H-nuclear magnetic resonance. Phytochem. Anal. 21: 102-109.

104 – Fontaine J.X., Molinie R., Tercé-Laforgue T., Cailleu D., Hirel B., Dubois F., Mesnard F. 2010. Use of 1H-NMR metabolomics to precise the function of the third glutamate dehydrogenase gene in Arabidospis thaliana. C. R. de Chimie 13 : 453–458.

105 – Cañas R.A., Quilleré I., Lea P.J., Hirel B. 2010. Analysis of amino acid metabolism in the ear of maize mutants deficient in two cytosolic glutamine synthetase isoenzymes highlights the importance of asparagine for nitrogen translocation within sink organs. Plant Biotech. J. 8 : 966-978.

106 – Cañas R.A., Amiour N., Quilleré I., Hirel B. 2011. An integrated statistical analysis of the genetic variability of nitrogen metabolism in the ear of three maize inbred lines (Zea mays L). J Exp. Bot. 62 :2309-2318.

107 – Hirel B., Tétu T., Lea P.J., Dubois F. 2011. Improving nitrogen use efficiency in crops for a sustainable agriculture. Sustainability 3 : 1452-1485.

108 – Cañas R.A., Quilleré I., Gallais A., Hirel B. 2012. Can genetic variability for nitrogen metabolism in the developing ear of maize be exploited to improve yield ? New Phytol. 194 : 440-452.

109 – Amiour N., Imbaud S., Clement G., Agier N., Zivy M., Valot B., Balliau T., Armengaud P., Quilleré I. , Cañas R.A., Tercé-Laforgue T., Hirel B. 2012. The use of metabolomics integrated with transcriptomic and proteomic studies for identifying key steps involved in the control of nitrogen metabolism in crops such as maize. J. Exp. Bot. 63 : 5017-5033.

110 – Fontaine J.X., Tercé-Laforgue T., Armengaud P. , Clément G., Renou J.P., Pelletier S., Catterou M., Azzopardi M., Gibon Y., Lea P.J., Hirel B., Dubois F. 2012. Characterization of a NADH-dependent glutamate dehydrogenase mutant of Arabidopsis demonstrates the key role of this enzyme in root carbon and nitrogen metabolism. The Plant Cell 24: 4044-4065.

111 – Fontaine J.X., Tercé-Laforgue T., Bouton S., Pageau K.,  Lea P.J., Dubois F., Hirel B., 2013. Further insights into the isoenzyme composition and activity of glutamate dehydrogenase in Arabidopsis Thaliana. Plant Signaling and Behavior. 8: 3, e233291-5.

112 – Tercé-Laforgue T., Bedu M., Dargel-Graffin C., Dubois F., Gibon Y., Restivo F.M., Hirel B., 2013. Resolving the role of plant glutamate dehydrogenase: II. Physiological Characterization of plants overexpressing individually or simultaneously the two enzyme subunits. Plant. Cell. Physiol. 54 : 1634-1647.

113 – Marchi L., Degola F., Polverini E., Tercé-Laforgue T., Dubois F., Hirel B., Restivo F., 2013. Glutamate dehydrogenase isoenzyme 3 (GDH3) of Arabidopsis thaliana is regulated by a combined effect of nitrogen and cytokinin. Plant Physiol. Biochem. 73 : 368-374.

114 – Simons M., Saha R., Guillard L., Clément G., Armengaud P., Cañas R., Maranas C.D., Lea P.J., Hirel B., 2014. Nitrogen use efficiency in maize (Zea mays L.): from “omics” studies to metabolic modelling. J. Exp. Bot. 65: 5657-5671.

115 – Simons M., Saha R., Amiour A., Kumar A., Guillard L., Clément G ., Miquel M., Li Z., Mouille G., Lea P.J., Hirel B., Maranas C.D. 2014. Assessing the metabolic impact of nitrogen availability using a compartmentalized maize leaf genome-scale model. Plant Physiol. 166 : 1659-1674.

116 – Amiour N., Imbaud S., Clément G., Agier N., Zivy M., Valot B., Balliau T., Quilleré I., Tercé-Laforgue T., Dargel-Graffin C., Hirel B. 2014. An integrated “omics” approach to the characterization of maize (Zea mays L.) mutants deficient in the expression of two genes encoding cytosolic glutamine synthetase. BMC Genomics 15: 1005.

117 – Tercé-Laforgue T., Clément G., Marchi L., Restivo F.M., Lea P.J., Hirel B. 2015. Resolving the role of plant NAD-glutamate dehydrogenase:III. Overexpressing individually or simultaneously the two enzyme subunits under salt stress induces changes in the leaf metabolic profile and increases plant biomass production. Plant Cell Physiol. 56 : 1918-1929.

118- Cormier F., Foulkes J., Hirel B., Gouache D.,  Moenne-Loccoz Y.,  Le Gouis J. 2016. Breeding for increased nitrogen use efficiency: a review for wheat (T. Aestivum L.). Plant Breeding 135 : 255-278.

119 – Verzeaux J., Roger D., Lacoux J., Nivelle E., Adam C., Habbib H., Hirel B., Dubois F.,  Tetu T. 2016. In winter wheat, no-till increases mycorrhizal colonization thus reducing the need for nitrogen fertilization. Agronomy 6 :38.

120 -Verzeaux J., Alahmad A., Habbib H., Nivelle E.,  Roger D., Lacoux J., Decocq, G., Hirel B., Catterou, M., Spicher, F., Dubois F., Duclercq J., Tetu T. 2016. Cover crops prevent from deleterious effect of nitrogen fertilization on bacterial diversity by maintaining carbon concentration in plowed soil. Geoderma 281: 49-57.

Book chapters:

1 – Hirel B. 1980. Les formes isofonctionnelles de la glutamine synthétase chez le riz (Oryza sativa  L.). Doctorat de 3ème cycle. Université de Nancy I. UER PC.B.

2 – Gadal P., Vidal J., Jacquot J.P., Hirel B., Perrot-Rechenmann C., Suzuki A., Crétin C. 1983. Light regulation of carbon and nitrogen metabolism enzymes: PEP carboxylase, NADP malate dehydrogenase and glutamate synthase. In “photosynthesis and Plant Productivity”, Metzner H. ed., Wissensh. Verlags. mbH Stuttgart, 249-254.

3 – Hirel B., 1984. Les isoenzymes de la glutamine synthétase chez les angiospermes. Thèse de Doctorat d’Etat, Université de Paris-Sud, Orsay.

4 – Nato A., Hoarau J., Brangeon J., Hirel B., Suzuki A. 1985. Regulation of carbon and nitrogen assimilation pathways in tobacco cell suspension cultures in relation with structural and biochemical development of the photosynthetic apparatus.In “International Symposium on Primary and Secondary Metabolism of Plant Cell Culture”. Giessen (FRG 5-8 Septembre, Springer-Verlag, Berlin, Heidelberg (Ed.) pp. 43-57.

5 – Hirel B., Bouet C., Gadal P. 1986. Molecular aspects of differentiation in the nitrogen metabolism of roots and root nodules. Colloque I.N.R.A. “Nutrition azotée des légumineuses”. Versailles, Novembre 1985, pp. 109-121.

6 – Verma D.P.S., Delauney A., Kuhse J., Hirel B., Shafer R., Raju K. 1988.  Metabolites and protein factors controllind nodulin gene expression. In nitrogen fixation : Hundred years after. H.Bothe, F. de Bruijn and E. Newton eds. Studgart/New-York. Gustav Fisher Publishers. pp 599-604.

7 – Verma P.P.S., Delauney A.J., Guida M., Hirel B., Shafer R., Koh S. 1988. Control of expression of nodulin genes. In Molecular Genetics of Plant-Microbe Interactions , R. Palacios and D.P.S. Verma  Eds. (St. Paul, MN, APS Press), pp 315-320.

8 – Hirel B., Miao G.H.,Verma D.P.S. 1992. Metabolic and developmental control of glutamine synthetase genes in legume and non-legume plants. In control of plant gene expression, Verma, D.P.S. ed. CRC Press, Inc. Boca Raton, FL. ICBN 0-8493-8866-X. pp. 443-458.

9 – Meyer, C., Hirel, B., Caboche, M., Morot-Gaudry, J.F. 1993. L’assimilation de l’azote chez les plantes. La Recherche (Septembre). pp. 956-962.

10 – Hirel, B., Marsolier, M.C. 1993. Current knowledge and perspectives for the manipulation of the ammonia assimilation pathway via transgenic plants. E.E.C. workshop “Genetic techniques for adapting crop plant to lowered fertilizers inputs and nutritional stresses” Weston Super Mare, U.K. 15-16 December 1992. Working document ref. F.II.3 -SJ/0011. Research and Technological Development Programme in the field of Competitiveness Agriculture and Management of Agricultural Resourses (1989-1993) CAMAR. pp. 99-113.

11 – Brugière N., Suzuki A., Hirel B. 1997.  Assimilation de l’ammonium. In Assimilation de l’azote chez les plantes : aspects physiologique, biochimique et moléculaire, J.F. Morot-Gaudry ed., INRA Editions, 85-107.

12 – Ferrario-Méry S., Murchie E., Hirel B., Galtier N., Quick., Foyer C.H. 1997. – Manipulation of the pathways of sucrose biosynthesis and nitrogen assimilation in transformed plants to improve photosynthesis and productivity. In A Molecular Approach to Primary Metabolism in Plants, C.H. Foyer and P. Quick, eds, Taylor and Francis, London, 125-153.

13 – Limami A.M., Leach F., Brugière N., Aziz A., Martin-Tanguy R.L., Malmberg R.L., Hirel B., Tepfer D. 1999. Altered polyamine metabolism and stress-induced proline accumulation in genetically manipulated tobacco. In Nutrient managment under salinity and water stress. Dahlia Greidinger International Symposium. Haifa, Israel, 17-29.

14 – Morot-Gaudry J.F, Hirel, B. 1999. Improvement of nitrogen use efficiency in plant for more efficient an environmentally compatible agriculture. In: Symposium of trans-century tobacco agricultural science and technology outlook and sutainable development strategies. China Commerce Publishing House. ISBN 7-5044-3987-8S°§23. pp. 16-23.

15 – Gallais A., Bertin P., Hirel B. 2000. Physiological and genetic bases of variation for nitrogen use efficiency in grain maize. Congrès CEE COST-814, Pordenone (Italie), 10-12 mai 2000.

16 – Harrison J., Brugière N., Phillipson B., Ferrario-Méry S., Becker T., Limami A., Hirel B. 2000. Manipulating the pathway of ammonia assimilation through genetic manipulation and breeding. Consequences on plant physiology and development. In: Nitrogen in a sustainable ecosystem from the cell to the plant. M.A. Martins-Loucao and S.H. Lips eds. Backhuys Publishers. Leiden. ISBN 90-5782-063-3 pp. 89- 101.

17 – Hirel B., Lea P.J. 2001. Ammonia assimilation. In: Plant Nitrogen. INRA Springer Eds. pp 79-99.

18 – Brugière N., Suzuki A, Hirel B. 2001. Ammonium Assimilation In: Nitrogen Assimilation by Plants, Physiological, Biochemical and Molecular Aspects, 2001, Ed JF Morot-Gaudry, Science Publishers, Inc., Enfield (NH), USA, Plymouth, UK,470p, ISBN 1-57808-139-4 pp. 71-94.

19 – Hirel B., Lea P.J. 2002. The biochemistry, Molecular Biology, and genetic manipulation of primary ammonia assimilation . In: Photosynthetic Nitrogen Assimilation and Associated Carbon and Respiratory Metabolism. Foyer C.H. and Noctor G. eds. Adavances in Photosynthesis and Respiration. Kluwer Academic Publishers. Dordrecht, Boston, London. pp. 71-92.

20 – Hirel B., Harrison J., Limami A. 2003. Improvement of Nitrogen Utilization. In: Improvement strategies for Leguminosae Biotechnology. Jaiwal P.K. and Singh R.P. eds. Kluwer Academic Publishers. pp. 201-220.

21 – Hirel B., LeGouis J., Falque M., Quétier F., Murigneux A., Rogwoski P., Charcosset, A. 2004. Le maïs et le blé, céréales  modèles pour la recherche en biologie intégrative et son application à la sélection. In: La Génomique en Biologie Végétale. INRA eds. pp. 453-471.

22 – Hirel B., Lemaire G. 2005.  From agronomy and ecophysiology to molecular genetics for improving nitrogen use efficiency in crops. In: Enhancing the efficiency of nitrogen utilisation in plants. Basra A., Goyal S and Tishner, eds. Journal of Crop Improvement. Food Product Press, the Haworth Press Inc. 15 : 213-257.

23 – Hirel B., Le Gouis J., Bernard M., Perez P., Falque M., Quétier F., Joets J., Montalent P., Rogwoski P., Murigneux A., Charcosset A. 2007. Genomics and Plant Breeding: maize and wheat. In : Functional Plant Genomics, JF Morot-Gaudry, Lea P, Briat JF, eds, Science Publishers, Enfield (NH), Jersey, Plymouth, pp 614-635.

24 – Vessey J.K., Fei H., Chaillou S., Hirel B., Polowick P., Mahon J.D. 2008. Effect of the overexpression of a soybean cytosolic glutamine synthetase gene linked to organ-specific promoters on growth and nitrogen accumulation of pea plants. In : Biological nitrogen fixation : towards aleviation through sustainable agriculture. Current Plant Science and Biotechnology in Agriculture. 42 : 20-209.

25 – Le Gouis J., Fontaine J.X, Laperche A., Heumez E., Devienne-Barret F., Brancourt-Hulmel M., Dubois F., Hirel B. 2008. Genetic analysis of wheat nitrogen use efficiency: coincidence between QTL for agronomical and physiological traits. In : The 11th International Wheat Genetics Symposium proceedings. R Appels, R Eastwood, E Lagudah, P Langridge, M Mackay Lynne, eds, Sydney University Press. ISBN: 978192089914. Pp 1-3.

26 – Loulakakis K.A., Morot-Gaudry J.F., Velanis C.N., Skopelitis D.S.,  Moschou P.N., Hirel B., Roubelakis-Angelakis K.A. 2009. Advancements in nitrogen metabolism in grapevine. In : Grapevine molecular physiology and biotechnology, 2nd Edition. KA Roubelakis-Angelakis ed. Springer, Netherlands, pp 161-205.

27 – Hirel B., Gallais A. 2011. Nitrogen use efficiency – Physiological, molecular and genetic investigations towards crop improvement, In : J.L. Prioul, C. Thévenot and T. Molnar (eds). Advances in Maize (Essential Reviews in Experimental Biology) (Vol.3), Society for Experimental Biology, UK, pp. 285-310.

28 – Hirel B., Lea P.J., 2011. The molecular genetics of nitrogen use efficiency in crops. In: M.J. Hawkesford, P.B. Barraclough (eds). The Molecular and Physiological Basis of Nutrient Use Efficiency in Crops. Wiley-Blackwell, Chichester, UK, 139-164.

29 – Hirel B., Gallais A. 2011. Des plantes transgnéniques pour une meilleure utilisation de l’azote. In : A. Ricroch, Y. Dattée, M. Fellous (eds). Biotechnologies végétales : environnement, alimentation, santé. Vuibert-AFBV, 161-169.

30 – Hirel B. 2014. Less is more. International innovation. Issue 167. pp. 79-81.


– Hirel B., Perez P., 2007 – Improvement of the kernel production of maize through the modulation of glutamine synthetase activity. WIPO Patent WO/2008/044150.

– Hirel B., Perez P. 2013 Kernel productivity of plants through the modulation of glutamine synthetase activity. US 20130185829 A1.