M. .. Materials, 168 6.2.1 Experimental design and treatment

. .. Results, 3.2 The sources for grain filling

, The grain sink

. .. Source-sink-relationship, , p.185

. .. Tolerance, 3.5.2 Tolerance and healthy crop traits, p.190

. .. Discussion,

.. .. Conclusion,

.. .. References,

.. .. Appendix, 6.A.1 Healthy trait correlation with tolerance estimates, p.202

.. .. Highlights,

. .. , The range of source/sink balance generated, p.206

. .. Scale, 208 7.3.1 Limits of the single slope-based estimation of tolerance, The quantification of tolerance and the

. The and . .. Stb-tolerance-traits, 212 7.4.1 Comparison of the experimental results

. .. Environment-e-ect,

.. .. Tolerance,

.. .. Perspectives,

. .. References, 223 References R. Ben Slimane. E ets de la septoriose foliaire sur la sénescence et les ux d'azote pendant le remplissage des grains chez le blé tendre, AgroParisTech, 2010.

I. J. Bingham and C. F. Topp, Potential contribution of selected canopy traits to the tolerance of foliar disease by spring barley, Plant Pathology, vol.58, issue.6, pp.1010-1020, 2009.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

A. Boonman, N. P. Anten, T. A. Dueck, W. J. Jordi, A. Van-der-werf et al., Functional significance of shade-induced leaf senescence in dense canopies: an experimental test using transgenic tobacco, The American Naturalist, vol.168, issue.5, pp.597-607, 2006.

B. Borghi, M. Corbellini, C. Minoia, M. Palumbo, N. D. Fonzo et al., E ects of Mediterranean climate on wheat bread-making quality, European journal of agronomy, vol.6, issue.3-4, pp.145-154, 1997.

A. K. Borrell and G. L. Hammer, Nitrogen dynamics and the physiological basis of stay-green in sorghum, Crop science, vol.40, issue.5, pp.1295-1307, 2000.

P. A. Brading, E. C. Verstappen, G. H. Kema, and J. K. Brown, A gene-for-gene relationship between wheat and Mycosphaerella graminicola, the Septoria tritici blotch pathogen, Phytopathology, vol.92, issue.4, pp.439-445, 2002.

J. K. Brown, L. Chartrain, P. Lasserre-zuber, and C. Saintenac, Genetics of resistance to Zymoseptoria tritici and applications to wheat breeding, Fungal Genetics and Biology, vol.79, pp.33-41, 2015.

R. J. Bryson, N. D. Paveley, W. S. Clark, R. Sylvester-bradley, and R. K. Scott, Use of in-field measurements of green leaf area and incident radiation to estimate the e ects of yellow rust epidemics on the yield of winter wheat, European Journal of Agronomy, vol.7, pp.53-62, 1997.

J. Burke and B. Dunne, Septoria tritici in winter wheat -to spray or not to spray? Irish farmer, pp.14-18, 2006.

J. Cartelle, A. Pedró, R. Savin, and G. A. Slafer, Grain weight responses to postanthesis spikelet-trimming in an old and a modern wheat under mediterranean conditions, European Journal of Agronomy, vol.25, issue.4, pp.365-371, 2006.

A. C. Castro and M. R. Simón, E ect of tolerance to Septoria tritici blotch on grain yield, yield components and grain quality in Argentinean wheat cultivars, Crop Protection, vol.90, pp.66-76, 2016.

H. F. Causin, R. N. Jauregui, and A. J. Barneix, The e ect of light spectral quality on leaf senescence and oxidative stress in wheat, Plant Science, vol.171, issue.1, pp.24-33, 2006.

L. Chartrain, P. A. Brading, J. C. Makepeace, and J. K. Brown, Sources of resistance to Septoria tritici blotch and implications for wheat breeding, Plant Pathology, vol.53, issue.4, pp.454-460, 2004.

J. Chen, Y. Liang, X. Hu, X. Wang, F. Tan et al., Physiological characterization of 'stay green'wheat cultivars during the grain filling stage under field growing conditions, Acta physiologiae plantarum, vol.32, issue.5, pp.875-882, 2010.

P. Cheval, A. Siah, M. Bomble, A. D. Popper, P. Reignault et al., Evolution of QoI resistance of the wheat pathogen Zymoseptoria tritici in Northern France, Crop protection, vol.92, pp.131-133, 2017.

H. J. Cools and B. A. Fraaije, Are azole fungicides losing ground against Septoria wheat disease? resistance mechanisms in Mycosphaerella graminicola, Pest Management Science, vol.64, issue.7, pp.681-684, 2008.

H. J. Cools and B. A. Fraaije, Update on mechanisms of azole resistance in Mycosphaerella graminicola and implications for future control. Pest management science, vol.69, pp.150-155, 2013.

F. Cormier, J. Foulkes, B. Hirel, D. Gouache, Y. Moënne-loccoz et al., Breeding for increased nitrogen-use e ciency: a review for wheat T. aestivum L.), Plant Breeding, vol.135, pp.255-278, 2016.

C. Cowger, M. Ho-er, and C. Mundt, Specific adaptation by Mycosphaerella graminicola to a resistant wheat cultivar, Plant Pathology, vol.49, issue.4, pp.445-451, 2000.

M. Cox, C. Qualset, and D. Rains, Genetic-variation for nitrogen assimilation and translocation in wheat. 2. Nitrogen assimilation in relation to grain-yield and protein, Crop science, vol.25, issue.3, pp.435-440, 1985.

S. J. Crafts-brandner, F. E. Below, V. A. Wittenbach, J. E. Harper, and R. H. Hageman, Di erential senescence of maize hybrids following ear removal II. Selected leaf, Plant Physiology, vol.74, issue.2, pp.368-373, 1984.

S. J. Crafts-brandner, R. Hölzer, and U. Feller, Influence of nitrogen deficiency on senescence and the amounts of RNA and proteins in wheat leaves, Physiologia Plantarum, vol.102, issue.2, pp.192-200, 1998.

T. Curtis and N. Halford, Food security: the challenge of increasing wheat yield and the importance of not compromising food safety, Annals of applied biology, vol.164, issue.3, pp.354-372, 2014.

J. C. Dawson, D. R. Huggins, and S. S. Jones, Characterizing nitrogen use e ciency in natural and agricultural ecosystems to improve the performance of cereal crops in lowinput and organic agricultural systems, Field Crops Research, vol.107, issue.2, pp.89-101, 2008.

S. Demotes-mainard, G. Doussinault, J. M. Meynard-;-demotes-mainard, G. Doussinault, and J. M. Meynard, Abnormalities in the male developmental programme of winter wheat induced by climatic stess at meiosis, Agronomie, vol.15, pp.357-365, 1995.

A. P. Derkx, S. Orford, S. Gri-ths, M. J. Foulkes, and M. J. Hawkesford, Identification of di erentially senescing mutants of wheat and impacts on yield, biomass and nitrogen partitioning, Journal of integrative plant biology, vol.54, issue.8, pp.555-566, 2012.

L. Ding, K. Wang, G. Jiang, D. Biswas, H. Xu et al., E ects of nitrogen deficiency on photosynthetic traits of maize hybrids released in di erent years, Annals of Botany, vol.96, issue.5, pp.925-930, 2005.

H. Dooley, M. Shaw, J. Spink, and S. Kildea, E ect of azole fungicide mixtures, alternations and dose on azole sensitivity in the wheat pathogen Zymoseptoria tritici, Plant Pathology, vol.65, issue.1, pp.124-136, 2016.

H. Dooley, M. W. Shaw, J. Mehenni-ciz, J. Spink, and S. Kildea, Detection of Zymoseptoria tritici SDHI-insensitive field isolates carrying the SdhC-H152R and SdhD-R47W substitutions. Pest management science, vol.72, pp.2203-2207, 2016.

K. E. Duncan and R. J. Howard, Cytological analysis of wheat infection by the leaf blotch pathogen Mycosphaerella graminicola, Mycological Research, vol.104, issue.9, pp.1074-1082, 2000.

L. Echarte, S. Rothstein, and M. Tollenaar, The response of leaf photosynthesis and dry matter accumulation to nitrogen supply in an older and a newer maize hybrid, Crop Science, vol.48, issue.2, pp.656-665, 2008.

D. B. Egli and W. P. Bruening, Source-sink relationships, seed sucrose levels and seed growth rates in soybean, Annals of botany, vol.88, issue.2, pp.235-242, 2001.

R. T. Ellis, Tolerance to the maize rust Puccinia polysora underw, Nature, vol.174, p.1021, 1954.

L. Eriksen, M. W. Shaw, and H. Østergård, A model of the e ect of pseudothecia on genetic recombination and epidemic development in populations of Mycosphaerella graminicola, Phytopathology, vol.91, issue.3, pp.240-248, 2001.

J. R. Evans, Photosynthesis and nitrogen relationships in leaves of C3 plants, Oecologia, vol.78, pp.9-19, 1989.

R. E. Evenson and D. Gollin, Assessing the impact of the green revolution, Science, vol.300, pp.758-762, 1960.

Z. Eyal, I. Maize, and W. I. Center, The Septoria diseases of wheat : concepts and methods of disease management / International Maize and Wheat Improvement Center, 1987.

Z. , Integrated control of Septoria diseases of wheat, References FAO. FAO Statistical pocketbook -World food and Agriculture, chapter Crops, vol.65, pp.28-29, 1981.

U. Feller and A. Fischer, Nitrogen-metabolism in senescing leaves, Critical reviews in plant sciences, vol.13, issue.3, pp.241-273, 1994.

R. A. Fischer, Understanding the physiological basis of yield potential in wheat, The Journal of Agricultural Science, vol.145, issue.2, pp.99-113, 2007.

R. A. Fischer and Y. M. Stockman, Kernel number per spike in wheat (Triticum aestivum l.): responses to preanthesis shading, Australian Journal of Plant Physiology, vol.7, issue.2, pp.169-180, 1980.

H. Fones and S. Gurr, The impact of Septoria tritici blotch disease on wheat: An EU perspective, Septoria tritici blotch disease of wheat: Tools and techniques to study the pathogen Zymoseptoria tritici, vol.79, pp.3-7, 2015.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

M. J. Foulkes, M. J. Hawkesford, P. B. Barraclough, M. J. Holdsworth, S. Kerr et al., Identifying traits to improve the nitrogen economy of wheat: Recent advances and future prospects, Field crops research, vol.114, issue.3, pp.329-342, 2009.

B. Fraaije, H. Cools, J. Fountaine, D. Lovell, J. Motteram et al., Role of ascospores in further spread of QoI-resistant cytochrome b alleles (G143A) in field populations of Mycosphaerella graminicola, Phytopathology, vol.95, issue.8, pp.933-941, 2005.

B. A. Fraaije, J. A. Lucas, W. S. Clark, and F. J. Burnett, QoI resistance development in populations of cereal pathogens in the UK, Proceedings of the BCPC International Congress, pp.689-694, 2003.

B. A. Fraaije, C. Bayon, S. Atkins, H. J. Cools, J. A. Lucas et al., Risk assessment studies on succinate dehydrogenase inhibitors, the new weapons in the battle to control Septoria leaf blotch in wheat, Molecular Plant Pathology, vol.13, issue.3, pp.263-275, 2012.

V. Fröhlich and U. Feller, E ect of phloem interruption on senescence and protein remobilization in the flag leaf of field-grown wheat, Biochemie und Physiologie der P anzen, vol.187, issue.2, pp.80118-80124, 1991.

J. Fu and B. Lee, Changes in photosynthetic characteristics during grain filling of a functional stay-green rice SNU-SG1 and its F1 hybrids, Journal of crop science and biotechnology, vol.11, pp.75-82, 2008.

O. Gaju, V. Allard, P. Martre, J. W. Snape, E. Heumez et al., Identification of traits to improve the nitrogen-use e ciency of wheat genotypes, Field Crops Research, vol.123, issue.2, pp.139-152, 2011.

O. Gaju, V. Allard, P. Martre, J. L. Gouis, D. Moreau et al., Nitrogen partitioning and remobilization in relation to leaf senescence, grain yield and grain nitrogen concentration in wheat cultivars, Field Crops Research, vol.155, pp.213-223, 2014.

Y. Gan, B. D. Gossen, L. Li, G. Ford, and S. Banniza, Cultivar type, plant population, and Ascochyta blight in chickpea, Agronomy journal, vol.99, issue.6, pp.1463-1470, 2007.

H. C. Godfray, The challenge of feeding 9-10 billion people equitably and sustainably, The Journal of Agricultural Science, vol.152, issue.S1, pp.2-8, 2014.

M. J. Gooding, P. J. Gregory, K. E. Ford, and S. Pepler, Fungicide and cultivar a ect post-anthesis patterns of nitrogen uptake, remobilization and utilization e ciency in wheat, Journal of Agricultural Science, vol.143, issue.6, pp.503-518, 2005.

D. Gouache, M. Bancal, B. De-solan, and P. Gate, Tolérance du blé tendre aux stress biotiques et abiotiques. Innovations Agronomiques, vol.35, pp.75-87, 2014.

P. L. Gregersen, The Molecular and Physiological Basis of Nutrient Use E ciency in Crops, chapter 5. Senescence and Nutrient Remobilization in Crop Plants, pp.83-102, 2011.

P. L. Gregersen, A. Culetic, L. Boschian, and K. Krupinska, Plant senescence and crop productivity, Plant Molecular Biology, vol.82, issue.6, pp.603-622, 2013.

M. J. Gri-n and N. Fischer, Laboratory studies on benzimidazole resistance in Septoria tritici, EPPO Bulletin, vol.15, pp.505-511, 1985.

Y. Guo and S. Gan, Convergence and divergence in gene expression profiles induced by leaf senescence and 27 senescence-promoting hormonal, pathological and environmental stress treatments, Plant, Cell & Environment, vol.35, issue.3, pp.644-655, 2012.

A. J. Hall and R. A. Richards, Prognosis for genetic improvement of yield potential and water-limited yield of major grain crops, Field Crops Research, vol.143, pp.18-33, 2013.

A. J. Hall, R. Savin, and G. A. Slafer, Is time to flowering in wheat and barley influenced by nitrogen? A critical appraisal of recent published reports, European Journal of Agronomy, vol.54, pp.40-46, 2014.

J. P. Hébrard, Azote et qualité des blés: fractioner pour plus de protéines, Perspectives Agricoles, issue.68, pp.21-29, 1999.

S. J. Inglese and N. D. Paul, Tolerance of Senecio vulgaris to infection and disease caused by native and alien rust fungi, Phytopathology, vol.96, issue.7, pp.718-726, 2006.

C. F. Jenner, T. D. Ugalde, and D. Aspinall, The physiology of starch and protein deposition in the endosperm of wheat, Australian journal of plant physiology, vol.18, issue.3, pp.211-226, 1991.

A. K. Joshi, M. Kumari, V. P. Singh, C. M. Reddy, S. Kumar et al., Stay green trait: variation, inheritance and its association with spot blotch resistance in spring wheat (Triticum aestivum L.), Euphytica, vol.153, issue.1, pp.59-71, 2007.

G. H. Kema, D. Z. Yu, F. H. Rijkenberg, M. W. Shaw, and R. P. Baayen, Histology of the pathogenesis of Mycosphaerella graminicola in wheat, Phytopathology, vol.86, issue.7, pp.777-786, 1996.

J. Keon, J. Antoniw, R. Carzaniga, S. Deller, J. L. Ward et al., Transcriptional adaptation of Mycosphaerella graminicola to programmed cell death (PCD) of its susceptible wheat host. Molecular plant-microbe interactions, vol.20, pp.178-193, 2007.

T. Kichey, B. Hirel, E. Heumez, F. Dubois, and J. L. Gouis, In winter wheat (Triticum aestivum L.), post-anthesis nitrogen uptake and remobilisation to the grain correlates with agronomic traits and nitrogen physiological markers, Field Crops Research, vol.102, issue.1, pp.22-32, 2007.

E. J. Kirby, Analysis of leaf, stem and ear growth in wheat from terminal spikelet stage to anthesis, Field Crops Research, vol.18, issue.2, pp.127-140, 1988.

T. Kramer, B. H. Gildemacher, M. Van-der, J. E. Ster, and . Parlevliet, Tolerance of spring barley cultivars to leaf rust, Puccinia hordei, Euphytica, vol.29, issue.2, pp.209-216, 1980.

M. Kremer and G. M. Ho-mann, E ects of leaf infections of Drechslera tritici-repentis on the carbohydrate and nitrogen metabolism of wheat plants, Journal of Plant Diseases and Protection, vol.100, issue.3, pp.259-277, 1993.

J. Levitt, Stress and strain terminology, pp.1-8, 1972.

D. J. Lovell, S. R. Parker, T. Hunter, D. J. Royle, and R. R. Coker, Influence of crop growth and structure on the risk of epidemics by Mycosphaerella graminicola (Septoria tritici) in winter wheat, Plant Pathology, vol.46, issue.1, pp.126-138, 1997.

J. P. Lynch, E. Glynn, S. Kildea, and J. Spink, Yield and optimum fungicide dose rates for winter wheat (Triticum aestivum L.) varieties with contrasting ratings for resistance to Septoria tritici blotch, Field Crops Research, vol.204, pp.89-100, 2017.

Y. References, C. T. Ma, D. A. Mackown, and . Van-sanford, Di erential e ects of partial spikelet removal and defoliation on kernel growth and assimilate partitioning among wheat cultivars, Field Crops Research, vol.47, issue.96, pp.16-16, 1996.

A. Martin, X. Belastegui-macadam, I. Quilleré, M. Floriot, M. Valadier et al., Nitrogen management and senescence in two maize hybrids di ering in the persistence of leaf greenness: agronomic, physiological and molecular aspects, New Phytologist, vol.167, issue.2, pp.483-492, 2005.

C. Masclaux-daubresse, M. Reisdorf-cren, and M. Orsel, Leaf nitrogen remobilisation for plant development and grain filling, Plant Biology, vol.10, pp.23-36, 2008.

C. Masclaux-daubresse, F. Daniel-vedele, J. Dechorgnat, F. Chardon, L. Gaufichon et al., Nitrogen uptake, assimilation and remobilization in plants: challenges for sustainable and productive agriculture, Annals of Botany, vol.105, issue.7, pp.1141-1157, 2010.
URL : https://hal.archives-ouvertes.fr/hal-01203920

C. Mccartney, A. Brûlé-babel, and L. Lamari, Inheritance of race-specific resistance to Mycosphaerella graminicola in wheat, Phytopathology, vol.92, issue.2, pp.138-144, 2002.

G. M. Murray, R. H. Martin, and B. R. Cullis, Relationship of the severity of Septoria tritici blotch of wheat to sowing time, rainfall at heading and average susceptibility of wheat cultivars in the area, Australian journal of agricultural research, vol.41, issue.2, pp.307-315, 1990.

B. Ney, M. Bancal, P. Bancal, I. J. Bingham, J. M. Foulkes et al., Crop architecture and crop tolerance to fungal diseases and insect herbivory. Mechanisms to limit crop losses, European Journal of Plant Pathology, vol.135, issue.3, pp.561-580, 2013.
URL : https://hal.archives-ouvertes.fr/hal-01000864

L. D. Noodén, J. W. Hillsberg, and M. J. Schneider, Induction of leaf senescence in Arabidopsis thaliana by long days through a light-dosage e ect, Physiologia Plantarum, vol.96, issue.3, pp.491-495, 1996.

K. Ono, I. Terashima, and A. Watanabe, Interaction between nitrogen deficit of a plant and nitrogen content in the old leaves, Plant and Cell Physiology, vol.37, issue.8, pp.1083-1089, 1996.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

M. Parry, M. Reynolds, C. Salvucci, P. Raines, X. Andralojc et al., Raising yield potential of wheat. II. Increasing photosynthetic capacity and e ciency, Journal of Experimental Botany, vol.62, pp.453-467, 2001.

A. Pask, Optimising nitrogen storage in wheat canopies for genetic reduction in fertiliser nitrogen inputs, 2009.

A. J. Pask, R. Sylvester-bradley, P. D. Jamieson, and M. J. Foulkes, Quantifying how winter wheat crops accumulate and use nitrogen reserves during growth, Field crops research, vol.126, pp.104-118, 2012.

J. B. Passioura, Soil conditions and plant growth, Plant, Cell & Environment, vol.25, issue.2, pp.311-318, 2002.

I. Rajcan and M. Tollenaar, Source: sink ratio and leaf senescence in maize: II. Nitrogen metabolism during grain filling, Field Crops Research, vol.60, issue.3, pp.255-265, 1999.

S. Recous, P. Loiseau, J. M. Machet, and B. Mary, Transformations et devenir de l'azote de l'engrais sous cultures annuelles et sous prairies, Les colloques de l'INRA, vol.83, pp.105-120, 1997.

M. P. Reynolds, A. Pellegrineschi, and B. Skovmand, Sink-limitation to yield and biomass: a summary of some investigations in spring wheat, Annals of applied biology, vol.146, issue.1, pp.39-49, 2005.

S. Rieben, O. Kalinina, B. Schmid, and S. L. Zeller, Gene flow in genetically modified wheat, Plos One, vol.6, issue.12, p.2011

C. Robert, C. Fournier, B. Andrieu, and B. Ney, Coupling a 3D virtual wheat (Triticum aestivum) plant model with a Septoria tritici epidemic model (Septo3D): a new approach to investigate plant-pathogen interactions linked to canopy architecture, Functional Plant Biology, vol.35, pp.997-1013, 2008.
URL : https://hal.archives-ouvertes.fr/hal-01192058

A. A. Rosielle and W. J. Boyd, Genetics of host-pathogen interactions to the Septoria species of wheat, 1985.

M. C. Rousseaux, A. J. Hall, and R. A. Sánchez, Far-red enrichment and photosynthetically active radiation level influence leaf senescence in field-grown sunflower, Physiologia Plantarum, vol.96, issue.2, pp.217-224, 1996.

S. A. Ruuska, G. J. Rebetzke, A. F. Van-herwaarden, R. A. Richards, N. A. Fettell et al., Genotypic variation in water-soluble carbohydrate accumulation in wheat, Functional Plant Biology, vol.33, issue.9, pp.799-809, 2006.

S. C. Salmon and H. H. Laude, Twenty years of testing varieties and strain of winter wheat at the Kansas agricultural experiment station. Technical bulletin 80, Kansas state college of agriculture and applied science, 1932.

R. Sanchez-bragado, G. Molero, M. P. Reynolds, and J. L. Araus, Relative contribution of shoot and ear photosynthesis to grain filling in wheat under good agronomical conditions assessed by di erential organ ?13c, Journal of experimental botany, vol.65, issue.18, pp.5401-5413, 2014.

R. Sanchez-bragado, M. D. Serret, and J. L. Araus, The nitrogen contribution of di erent plant parts to wheat grains: Exploring genotype, water, and nitrogen e ects, Frontiers in Plant Science, vol.7, 1986.

M. Sanchez-garcia, C. Royo, N. Aparicio, J. Martin-sanchez, and F. Alvaro, Genetic improvement of bread wheat yield and associated traits in Spain during the 20th century, The Journal of Agricultural Science, vol.151, issue.1, pp.105-118, 2013.

A. Sánchez-vallet, M. C. Mcdonald, P. S. Solomon, and B. A. Mcdonald, Is Zymoseptoria tritici a hemibiotroph?, Septoria tritici blotch disease of wheat: Tools and techniques to study the pathogen Zymoseptoria tritici, vol.79, pp.29-32, 2015.

S. Savary, N. P. Castilla, F. A. Elazegui, C. G. Mc-laren, M. A. Ynalvez et al., Direct and indirect e ects of nitrogen supply and disease source structure on rice sheath blight spread, Phytopathology, vol.85, issue.9, pp.959-965, 1995.

J. Schafer, Annual review of phytopathology, vol.9, pp.235-252, 1971.

A. L. Scharen and J. M. Krupkinsk, E ect of Septoria nodorum infection on CO2 absorption and yield of wheat, Phytopathology, vol.59, issue.9, p.1298, 1969.

R. A. Serrago, I. Alzueta, R. Savin, and G. A. Slafer, Understanding grain yield responses to source-sink ratios during grain filling in wheat and barley under contrasting environments, Field Crops Research, vol.150, pp.42-51, 2013.

M. W. Shaw and D. J. Royle, Factors determining the severity of epidemics of Mycosphaerella graminicola (Septoria tritici) on winter wheat in the UK, Plant Pathology, vol.42, issue.6, pp.882-899, 1993.

V. J. Shearman, R. Sylvester-bradley, R. K. Scott, and M. J. Foulkes, Physiological processes associated with wheat yield progress in the UK, Crop Science, vol.45, issue.1, pp.175-185, 2005.

P. R. Shewry and . Wheat, Journal of experimental botany, vol.60, issue.6, pp.1537-1553, 2009.

P. R. Shewry, R. A. Mitchell, P. Tosi, Y. Wan, C. Underwood et al., An integrated study of grain development of wheat (cv. Hereward), Journal of Cereal Science, vol.56, issue.1, pp.21-30, 2012.

M. R. Simón, A. E. Perelló, C. A. Cordo, S. Larrán, P. E. Van-der-putten et al., Association between Septoria tritici blotch, plant height, and heading date in wheat, Agronomy Journal, vol.97, issue.4, pp.1072-1081, 2005.

T. R. Sinclair and P. D. Jamieson, Grain number, wheat yield, and bottling beer: an analysis, Field Crops Research, vol.98, issue.1, pp.60-67, 2006.

G. A. Slafer and F. H. Andrade, Changes in physiological attributes of the dry matter economy of bread wheat (Triticum aestivum) through genetic improvement of grain yield potential at di erent regions of the world, Euphytica, vol.58, issue.1, pp.37-49, 1991.

G. A. Slafer and R. Savin, Postanthesis green area duration in a semidwarf and a standardheight wheat cultivar as a ected by sink strength, Australian Journal of Agricultural Research, 1994.

G. A. Slafer and R. Savin, Source-sink relationships and grain mass at di erent positions within the spike in wheat, Field Crops Research, vol.37, issue.94, pp.90080-90089, 1994.

G. A. Slafer, R. Savin, and V. O. Sadras, Coarse and fine regulation of wheat yield components in response to genotype and environment, Field Crops Research, vol.157, issue.0, pp.71-83, 2014.

O. A. Somasco, C. O. Qualset, and D. G. Gilchrist, Single-gene resistance to Septoria tritici blotch in the spring wheat cultivar 'Tadinia', Plant Breeding, vol.115, issue.4, pp.261-267, 1996.

G. Spano, N. D. Fonzo, C. Perrotta, C. Platani, G. Ronga et al., Physiological characterization of 'stay green' mutants in durum wheat, Journal of Experimental Botany, vol.54, issue.386, pp.1415-1420, 2003.

J. Spiertz and N. Devos, Agronomical and physiological-aspects of the role of nitrogen in yield formation of cereals, Plant and soil, vol.75, issue.3, pp.379-391, 1983.

G. Steinberg, Cell biology of Zymoseptoria tritici: Pathogen cell organization and wheat infection, Fungal Genetics and Biology, vol.79, pp.17-23, 2015.

S. M. Tabib-gha-ary, J. D. Faris, T. L. Friesen, R. G. Visser, T. A. Van-der-lee et al., New broad-spectrum resistance to Septoria tritici blotch derived from synthetic hexaploid wheat, Theoretical and Applied Genetics, vol.124, issue.1, pp.125-142, 2012.

E. A. Tambussi, J. Bort, J. J. Guiamet, S. Nogues, and J. Luis-araus, The photosynthetic role of ears in C3 cereals: Metabolism, water use e ciency and contribution to grain yield, Critical reviews in plant sciences, vol.26, issue.1, pp.1-16, 2007.

H. Thomas and L. De-villiers, Gene expression in leaves of Arabidopsis thaliana induced to senesce by nutrient deprivation, Journal of Experimental Botany, vol.47, issue.12, pp.1845-1852, 1996.

H. Thomas and C. J. Howarth, Five ways to stay green, Journal of Experimental Botany, vol.51, issue.S1, pp.329-337, 2000.

D. K. Tompkins, D. B. Fowler, and A. T. Wright, Influence of agronomic practices on canopy microclimate and Septoria development in no-till winter wheat produced in the Parkland region of Saskatchewan, Canadian Journal of Plant Science, vol.73, issue.1, pp.331-344, 1993.

S. F. Torriani, J. P. Melichar, C. Mills, N. Pain, H. Sierotzki et al., Zymoseptoria tritici: A major threat to wheat production, integrated approaches to control, Fungal Genetics and Biology, vol.79, pp.8-12, 2015.

D. A. Vansanford and C. T. Mackown, Cultivar di erences in nitrogen remobilization during grain fill in soft red winter-wheat, Crop science, vol.27, issue.2, pp.295-300, 1987.

P. M. Vitousek, Beyond global warming: ecology and global change, Ecology, vol.75, issue.7, pp.1861-1876, 1994.

L. Weaver and R. Amasino, Senescence is induced in individually darkened Arabidopsis leaves, but inhibited in whole darkened plants, Plant Physiology, vol.127, pp.876-886, 2001.

R. E. Wilson, Inheritance of resistance to Septoria tritici in wheat, Septoria of Cereals, 1985.

A. Wingler, S. Purdy, J. A. Maclean, and N. Pourtau, The role of sugars in integrating environmental signals during the regulation of leaf senescence, Journal of Experimental Botany, vol.57, issue.2, pp.1460-2431, 2006.

J. C. Zadoks, T. T. Chang, and C. F. Konzak, A decimal code for the growth stages of cereals, Weed Research, vol.44, pp.415-421, 1974.

J. Zhan and B. A. Mcdonald, The interaction among evolutionary forces in the pathogenic fungus Mycosphaerella graminicola, Fungal Genetics and Biology, vol.41, issue.6, pp.590-599, 2004.

Y. Zhang, N. Sun, J. Hong, Q. Zhang, C. Wang et al., E ect of source-sink manipulation on photosynthetic characteristics of flag leaf and the remobilization of dry mass and nitrogen in vegetative organs of wheat, Journal of integrative agriculture, vol.13, issue.8, pp.60665-60671, 2014.

O. Ziv and Z. , Assessment of yield component losses caused in plants of spring wheat cultivars by selected isolates of Septoria tritici, Phytopathology, vol.68, issue.5, pp.791-796, 1978.

E. Zuckerman, A. Eshel, and Z. Eyal, Physiological aspects related to tolerance of spring wheat cultivars to Septoria tritici blotch, Phytopathology, vol.87, issue.1, pp.60-65, 1997.

. Ahdb, Ahdb Recommended List -winter wheat 2016/17. tables, 2016.

M. Bancal, R. B. Slimane, and P. Bancal, Zymoseptoria tritici development induces local senescence in wheat leaves, without a ecting their monocarpic senescence under two contrasted nitrogen nutrition, Environmental and Experimental Botany, vol.132, pp.154-162, 2016.

P. Bancal, M. Bancal, F. Collin, and D. Gouache, Identifying traits leading to tolerance of wheat to Septoria tritici blotch, Field Crops Research, vol.180, pp.176-185, 2015.
URL : https://hal.archives-ouvertes.fr/hal-01535231

R. Ben-slimane, P. Bancal, and M. Bancal, Down-regulation by stems and sheaths of grain filling with mobilized nitrogen in wheat, Field Crops Research, vol.140, issue.0, pp.59-68, 2013.
URL : https://hal.archives-ouvertes.fr/hal-01000853

L. Breiman, Random forests, Machine Learning, vol.45, pp.5-32, 2001.

. Hgca and . Hgca, , vol.16, 2015.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

R. Piessens, E. De-doncker-kapenga, C. Uberhuber, and D. Kahaner, Quadpack: a Subroutine Package for Automatic Integration, pp.978-3540125531, 1983.

. R-core-team, R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, 2017.

, RAGT. Blé tendre d'hiver. Leaflet, 2016.

J. C. Zadoks, T. T. Chang, and C. F. Konzak, A decimal code for the growth stages of cereals, .tb01084.x . References AHDB. Wheat disease maagement guide. AHDB, vol.44, pp.415-421, 1974.

P. Bancal, Data: Précocité et surface de feuille étendard, 2017.

D. Bates, M. Mächler, B. Bolker, and S. Walker, Fitting linear mixed-e ects models using lme4, Journal of Statistical Software, vol.67, issue.1, pp.1-48, 2015.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

L. Borrás, G. A. Slafer, and M. E. Otegui, Seed dry weight response to source sink manipulations in wheat, maize and soybean: a quantitative reappraisal, Field Crops Research, vol.86, issue.23, pp.131-146, 2004.

L. Breiman, Manual On Setting Up, Using, And Understanding Random Forests

L. Breiman, Random forests, Machine Learning, vol.45, pp.5-32, 2001.

H. F. Causin, R. N. Jauregui, and A. J. Barneix, The e ect of light spectral quality on leaf senescence and oxidative stress in wheat, Plant Science, vol.171, issue.1, pp.24-33, 2006.

P. Cheval, A. Siah, M. Bomble, A. D. Popper, P. Reignault et al., Evolution of QoI resistance of the wheat pathogen Zymoseptoria tritici in Northern France, Crop protection, vol.92, pp.131-133, 2017.

F. Collin, P. Bancal, J. Foulkes, and M. O. Bancal, A statistical analysis of gxe contribution to leaf senescence during grain filling in wheat, ESA 14 -Growing landscapes -Cultivating innovative agricultural systems, pp.13-14, 2016.

F. Collin, D. Gouache, M. Bancal, and P. Bancal, Tolerance of wheat to Septoria tritici blotch; genetic vs environmental variations of key traits, 9th International Symposium on Septoria Diseases of Cereals, p.76, 2016.

J. Conway, D. Eddelbuettel, T. Nishiyama, S. K. Prayaga, and N. Ti-n, RPostgreSQL: R interface to the PostgreSQL database system, 2016.

H. J. Cools and B. A. Fraaije, Update on mechanisms of azole resistance in Mycosphaerella graminicola and implications for future control. Pest management science, vol.69, pp.150-155, 2013.

M. Crawley, Statistics: an introduction using R, chapter Mixed-e ects models, pp.627-660, 2007.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

D. Gouache, Toward a new indicator of cultivar resistance to Septoria tritici blotch accounting for cultivar phenology. unpublished, 2009.

D. Gouache, A. Bensadoun, F. Brun, C. Pagé, D. Makowski et al., Modelling climate change impact on Septoria tritici blotch (STB) in France: accounting for climate model and disease model uncertainty, Agricultural and Forest Meteorology, vol.170, pp.242-252, 2013.
URL : https://hal.archives-ouvertes.fr/hal-01186800

P. Gregersen, P. Holm, and K. Krupinska, Leaf senescence and nutrient remobilisation in barley and wheat, Plant Biology, vol.10, issue.s1, pp.37-49, 2008.

P. L. Gregersen, A. Culetic, L. Boschian, and K. Krupinska, Plant senescence and crop productivity, Plant Molecular Biology, vol.82, issue.6, pp.603-622, 2013.

R. Lemoine, S. L. Camera, R. Atanassova, F. Dédaldéchamp, T. Allario et al., Source-to-sink transport of sugar and regulation by environmental factors, Frontiers in Plant Science, vol.4, pp.1-21, 2013.
URL : https://hal.archives-ouvertes.fr/hal-00870386

A. Liaw and M. Wiener, Classification and regression by Rrandom Forest, R News, vol.2, issue.3, pp.18-22, 2002.

K. Ono, I. Terashima, and A. Watanabe, Interaction between nitrogen deficit of a plant and nitrogen content in the old leaves, Plant and Cell Physiology, vol.37, issue.8, pp.1083-1089, 1996.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

D. L. Parrott, J. M. Martin, and A. M. Fischer, Analysis of barley (Hordeum vulgare) leaf senescence and protease gene expression: a family C1A cysteine protease is specifically induced under conditions characterized by high carbohydrate, but low to moderate nitrogen levels, New Phytologist, vol.187, issue.2, pp.313-331, 2010.

. R-core-team, R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, 2017.

, R Special Interest Group on Databases. DBI: R Database Interface, 2014.

R. Rakotomalala, Pratique de la régression linéaire multiple -Diagnostic et sélection de variables. Version 2.1, 2015.

C. Robert, B. Andrieu, C. Fournier, D. Gouache, P. Gate et al., Septo3D: un modèle pour analyser les e ets de la structure des couverts de blé sur les épidémies de septoriose, Conférence internationale, vol.9, p.page np, 2009.

D. Sarkar, Lattice: Multivariate Data Visualization with R, 2008.

D. Sarkar and F. Andrews, latticeExtra: Extra Graphical Utilities Based on Lattice, pp.6-28, 2016.

V. J. Shearman, R. Sylvester-bradley, R. K. Scott, and M. J. Foulkes, Physiological processes associated with wheat yield progress in the UK, Crop Science, vol.45, issue.1, pp.175-185, 2005.

U. Steinfort, B. Trevaskis, S. Fukai, K. L. Bell, and M. F. Dreccer, Vernalisation and photoperiod sensitivity in wheat: Impact on canopy development and yield components, Field Crops Research, vol.201, pp.108-121, 2017.

V. Svetnik, A. Liaw, C. Tong, and T. Wang, Multiple Classi er Systems: 5th International Workshop, Proceedings, chapter Application of Breiman's Random Forest to Modeling Structure-Activity Relationships of Pharmaceutical Molecules, pp.334-343, 2004.

L. Taiz and E. Zeiger, Plant physiology and development, chapter Plant senescence and cell death, pp.665-692, 2014.

H. Thomas and L. De-villiers, Gene expression in leaves of Arabidopsis thaliana induced to senesce by nutrient deprivation, Journal of Experimental Botany, vol.47, issue.12, pp.1845-1852, 1996.

C. Uauy, J. C. Brevis, and J. Dubcovsky, The high grain protein content gene Gpc-B1 accelerates senescence and has pleiotropic e ects on protein content in wheat, Journal of Experimental Botany, vol.57, issue.11, pp.2785-2794, 2006.

L. Weaver and R. Amasino, Senescence is induced in individually darkened Arabidopsis leaves, but inhibited in whole darkened plants, Plant Physiology, vol.127, pp.876-886, 2001.

J. C. Zadoks, T. T. Chang, and C. F. Konzak, A decimal code for the growth stages of cereals, Weed Research, vol.44, pp.415-421, 1974.

M. M. Acreche and G. A. Slafer, Variation of grain nitrogen content in relation with grain yield in old and modern Spanish wheats grown under a wide range of agronomic conditions in a Mediterranean region, The Journal of Agricultural Science, vol.147, issue.6, pp.657-667, 2009.

K. A. Aisawi, M. P. Reynolds, R. P. Singh, and M. J. Foulkes, The physiological basis of the genetic progress in yield potential of CIMMYT spring wheat cultivars from, Crop Science, vol.55, pp.1749-1764, 1966.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

J. Burke and B. Dunne, Septoria tritici in winter wheat -to spray or not to spray? Irish farmer, pp.14-18, 2006.

D. F. Calderini, R. Savin, L. G. Abeledo, M. P. Reynolds, and G. A. Slafer, The importance of the period immediately preceding anthesis for grain weight determination in wheat, Euphytica, vol.119, issue.1-2, pp.199-204, 2001.

R. Carretero, R. A. Serrago, M. O. Bancal, A. E. Perelló, and D. J. Miralles, Absorbed radiation and radiation use e ciency as a ected by foliar diseases in relation to their vertical position into the canopy in wheat, Field Crops Research, vol.116, issue.1, pp.184-195, 2010.

P. Cheval, A. Siah, M. Bomble, A. D. Popper, P. Reignault et al., Evolution of QoI resistance of the wheat pathogen Zymoseptoria tritici in Northern France, Crop protection, vol.92, pp.131-133, 2017.

F. Collin, P. Bancal, J. Spink, P. Kock-appelgren, J. Smith et al., Wheat lines exhibiting variation in tolerance of Septoria tritici blotch di erentiated by grain source limitation, Field Crops Research, vol.217, pp.1-10, 2018.

J. Conway, D. Eddelbuettel, T. Nishiyama, S. K. Prayaga, and N. Ti-n, RPostgreSQL: R interface to the PostgreSQL database system, 2016.

P. S. Cornish, G. R. Baker, and G. M. Murray, Physiological responses of wheat (triticum aestivum) to infection with Mycosphaerella graminicola causing Septoria tritici blotch, Australian Journal of Agricultural Research, vol.41, issue.2, pp.317-327, 1990.

M. J. Crawley, The R Book, ects Models, pp.681-714, 2012.

F. M. Dupont and S. B. Altenbach, Molecular and biochemical impacts of environmental factors on wheat grain development and protein synthesis, Journal of Cereal Science, vol.38, issue.2, pp.133-146, 2003.

D. B. Egli and W. P. Bruening, Source-sink relationships, seed sucrose levels and seed growth rates in soybean, Annals of botany, vol.88, issue.2, pp.235-242, 2001.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

B. A. Fraaije, J. A. Lucas, W. S. Clark, and F. J. Burnett, QoI resistance development in populations of cereal pathogens in the UK, Proceedings of the BCPC International Congress, pp.689-694, 2003.

R. E. Gaunt and A. C. Wright, Disease-yield relationship in barley. II. Contribution of stored stem reserves to grain filling, Plant Pathology, vol.41, issue.6, pp.688-701, 1992.

R. V. Lenth, Least-squares means: The R package lsmeans, Journal of Statistical Software, vol.69, issue.1, pp.1-33, 2016.

Y. Ma, C. T. Mackown, and D. A. Van-sanford, Di erential e ects of partial spikelet removal and defoliation on kernel growth and assimilate partitioning among wheat cultivars, Field Crops Research, vol.47, issue.96, pp.16-16, 1996.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

J. Pinheiro, D. Bates, S. Debroy, D. Sarkar, and R. Team, Linear and Nonlinear Mixed E ects Models, 2017.

. R-core-team, R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, 2017.

, R Special Interest Group on Databases. DBI: R Database Interface, 2014.

M. Reynolds, A. Pask, and D. Mullan, Physiological breeding I: interdisciplinary approaches to improve crop adaptation, CIMMYT, 2012.

D. Sarkar, Lattice: Multivariate Data Visualization with R, 2008.

J. Schafer, Annual review of phytopathology, vol.9, pp.235-252, 1971.

M. Schierenbeck, M. C. Fleitas, D. J. Miralles, and M. R. Simón, Does radiation interception or radiation use e ciency limit the growth of wheat inoculated with tan spot or leaf rust? Field Crops Research, vol.199, pp.65-76, 2016.

R. A. Serrago, R. Carretero, M. O. Bancal, and D. J. Miralles, Grain weight response to foliar diseases control in wheat (triticum aestivum l.), Field Crops Research, vol.120, issue.3, pp.352-359, 2011.
URL : https://hal.archives-ouvertes.fr/hal-01000496

V. J. Shearman, R. Sylvester-bradley, R. K. Scott, and M. J. Foulkes, Physiological processes associated with wheat yield progress in the UK, Crop Science, vol.45, issue.1, pp.175-185, 2005.

G. A. Slafer and R. Savin, Source-sink relationships and grain mass at di erent positions within the spike in wheat, Field Crops Research, vol.37, issue.94, pp.90080-90089, 1994.

R. J. Smith, Use and misuse of the reduced major axis for line-fitting, American Journal of Physical Anthropology, vol.140, issue.3, pp.476-486, 2009.

J. Voltas, I. Romagosa, and J. L. Araus, Grain size and nitrogen accumulation in sinkreduced barley under Mediterranean conditions, Field Crops Research, vol.52, pp.117-126, 1997.

J. C. Zadoks, T. T. Chang, and C. F. Konzak, A decimal code for the growth stages of cereals, Weed Research, vol.44, pp.415-421, 1974.

O. Ziv and Z. , Assessment of yield component losses caused in plants of spring wheat cultivars by selected isolates of Septoria tritici, Phytopathology, vol.68, issue.5, pp.791-796, 1978.

E. Zuckerman, A. Eshel, and Z. Eyal, Physiological aspects related to tolerance of spring wheat cultivars to Septoria tritici blotch, Phytopathology, vol.87, issue.1, pp.60-65, 1997.

P. Bancal, Positive contribution of stem growth to grain number per spike in wheat, Field Crops Research, vol.105, issue.1-2, pp.27-39, 2008.
URL : https://hal.archives-ouvertes.fr/hal-01191988

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

S. Demotes-mainard, G. Doussinault, and J. M. Meynard, E ects of low radiation and low temperature at meiosis on pollen viability and grain set in wheat, Agronomie, vol.15, pp.357-365, 1995.

S. Demotes-mainard, G. Doussinault, and J. M. Meynard, Abnormalities in the male developmental programme of winter wheat induced by climatic stess at meiosis, vol.15, pp.357-365, 1996.

R. A. Fischer and Y. M. Stockman, Kernel number per spike in wheat (Triticum aestivum l.): responses to preanthesis shading, Australian Journal of Plant Physiology, vol.7, issue.2, pp.169-180, 1980.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

D. Gouache, M. Bancal, and P. Bancal, Projet contrat de branche, Tolérance du blé tendre aux stress biotiques et abiotiques" C2008-02 blé tendre, 2012.

E. Justes, B. Mary, J. M. Meynard, J. M. Machet, and L. Thelierhuche, Determination of a critical nitrogen dilution curve for winter-wheat crops, Annals of Botany, vol.74, issue.4, pp.397-407, 1994.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

A. References, Wheat disease maagement guide. AHDB, 2016.

. Ahdb, Ahdb Recommended List -winter wheat 2016/17. tables, 2016.

D. Bates, M. Mächler, B. Bolker, and S. Walker, Fitting linear mixed-e ects models using lme4, Journal of Statistical Software, vol.67, issue.1, pp.1-48, 2015.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

J. Conway, D. Eddelbuettel, T. Nishiyama, S. K. Prayaga, and N. Ti-n, RPostgreSQL: R interface to the PostgreSQL database system, 2016.

M. J. Crawley, The R Book, ects Models, pp.681-714, 2012.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

D. Gouache, M. Bancal, B. De-solan, and P. Gate, Tolérance du blé tendre aux stress biotiques et abiotiques. Innovations Agronomiques, vol.35, pp.75-87, 2014.

. Hgca and . Hgca, , vol.16, 2015.

R. V. Lenth, Least-squares means: The R package lsmeans, Journal of Statistical Software, vol.69, issue.1, pp.1-33, 2016.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

R. Piessens, E. De-doncker-kapenga, C. Uberhuber, and D. Kahaner, Quadpack: a Subroutine Package for Automatic Integration, pp.978-3540125531, 1983.

, References R Core Team. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, 2016.

, R Special Interest Group on Databases. DBI: R Database Interface, 2014.

, RAGT. Blé tendre d'hiver. Leaflet, 2016.

D. Sarkar, Lattice: Multivariate Data Visualization with R, 2008.

V. J. Shearman, R. Sylvester-bradley, R. K. Scott, and M. J. Foulkes, Physiological processes associated with wheat yield progress in the UK, Crop Science, vol.45, issue.1, pp.175-185, 2005.

J. C. Zadoks, T. T. Chang, and C. F. Konzak, A decimal code for the growth stages of cereals, Weed Research, vol.44, pp.415-421, 1974.

I. J. Bingham and C. F. Topp, Potential contribution of selected canopy traits to the tolerance of foliar disease by spring barley, Plant Pathology, vol.58, issue.6, pp.1010-1020, 2009.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

A. Borrell, G. Hammer, and E. Van-oosterom, Stay-green: A consequence of the balance between supply and demand for nitrogen during grain filling?, Annals of Applied Biology, vol.138, issue.1, pp.91-95, 2001.

H. Braun and T. Payne, Physiological breeding I: interdisciplinary approaches to improve crop adaptation, chapter Mega-environment breeding, pp.6-17, 2012.

J. Deswarte, K. Beauchene, G. Arjaure, S. Jezequel, G. Meloux et al., Platform development for drought tolerance evaluation of wheat in france, Procedia Environmental Sciences, vol.29, pp.93-94, 2015.

A. Distelfeld, R. Avni, and A. M. Fischer, Senescence, nutrient remobilization, and yield in wheat and barley, Journal of experimental botany, vol.65, pp.3783-3798, 2014.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

D. Gouache, M. Bancal, and P. Bancal, Projet contrat de branche, Tolérance du blé tendre aux stress biotiques et abiotiques" C2008-02 blé tendre, 2012.

D. Gouache, M. Bancal, B. De-solan, and P. Gate, Tolérance du blé tendre aux stress biotiques et abiotiques. Innovations Agronomiques, vol.35, pp.75-87, 2014.

S. J. Inglese and N. D. Paul, Tolerance of Senecio vulgaris to infection and disease caused by native and alien rust fungi, Phytopathology, vol.96, issue.7, pp.718-726, 2006.

J. L. Monteith and C. J. Moss, Climate and the e ciency of crop production in Britain, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, pp.277-294, 1977.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

N. D. Paveley, R. Sylvester-bradley, R. K. Scott, J. Craigon, and W. Day, Steps in predicting the relationship of yield on fungicide dose, Phytopathology, vol.91, issue.7, pp.708-716, 2001.

G. J. Rebetzke, J. A. Jimenez-berni, W. D. Bovill, D. M. Deery, and R. A. James, Highthroughput phenotyping technologies allow accurate selection of stay-green, Journal of Experimental Botany, vol.67, issue.17, pp.4919-4924, 2016.

M. Reynolds, M. J. Foulkes, G. A. Slafer, P. Berry, M. A. Parry et al., Raising yield potential in wheat, Journal of Experimental Botany, vol.60, issue.7, pp.1899-1918, 2009.
URL : https://hal.archives-ouvertes.fr/hal-00964196

J. Schafer, Annual review of phytopathology, vol.9, pp.235-252, 1971.

J. D. Scholes and J. F. Farrar, Increased rates of photosynthesis in localized regions of a barley leaf infected with brown rust, New Phytologist, vol.104, issue.4, pp.601-612, 1986.

V. J. Shearman, R. Sylvester-bradley, R. K. Scott, and M. J. Foulkes, Physiological processes associated with wheat yield progress in the UK, Crop Science, vol.45, issue.1, pp.175-185, 2005.

T. R. Sinclair and P. D. Jamieson, Grain number, wheat yield, and bottling beer: an analysis, Field Crops Research, vol.98, issue.1, pp.60-67, 2006.

G. A. Slafer, R. Savin, and V. O. Sadras, Coarse and fine regulation of wheat yield components in response to genotype and environment, Field Crops Research, vol.157, issue.0, pp.71-83, 2014.

H. Thomas and H. Ougham, The stay-green trait, Journal of Experimental Botany, vol.65, issue.14, pp.3889-3900, 2014.

O. Ziv and Z. , Assessment of yield component losses caused in plants of spring wheat cultivars by selected isolates of Septoria tritici, Phytopathology, vol.68, issue.5, pp.791-796, 1978.

E. Zuckerman, A. Eshel, and Z. Eyal, Physiological aspects related to tolerance of spring wheat cultivars to Septoria tritici blotch, Phytopathology, vol.87, issue.1, pp.60-65, 1997.

. Serrago, essentiel des dégâts est donc fonction des symptômes qui s'étendent sur les feuilles supérieures. Ceux-ci limitent la photosynthèse et par conséquent l'assimilation de carbone durant la période de remplissage du grain, 2009.

. Bingham, Tout d'abord, les réserves carbonées, essentiellement localisées dans les tiges, constitueraient une source alternative mobilisable lorsque la photosynthèse est limitée par la maladie, Actuellement, deux principales voies sont explorées pour accroître la tolérance, 1985.

. Bingham, Enfin, le niveau de disponibilité en assimilats par grain conditionnerait les pertes observées (van den Berg et al., 2017) : s'il est pléthorique les pertes seront nulles ou faibles ; s'il est limitant, la réduction des sources par les maladies se traduira par une diminution du PMG, un élargissement des surfaces photosynthétiques, 1997.

, Le manuscrit se compose d'une revue de la littérature scientifique, p.1

, visant à définir les concepts sur lesquels repose la tolérance du blé à la septoriose, décrire les processus physiologiques invoqués dans le manuscrit, et identifier les manques scientifiques et les enjeux qui justifient les objectifs de ce travail de thèse. Les méthodologies déployées et l'articulation des études menées sont présentées dans le Chapitre, vol.2, p.43

, Les quatre chapitres suivants (chapitres 3 -6, p.53) constituent les synthèses des quatre études misent en oeuvre lors de ce projet. Enfin, une large discussion des résultats est proposée au Chapitre, où les faits saillants de la thèse sont mis en lumière, ressitués dans un cadre général afin d'appréhender la portée du travail réalisé mais aussi de suggérer de nouvelles hypothèses et perspectives à l'aune de ces nouveaux résultats, vol.7, p.205

. Bancal, Cette piste privilégiée a été l'objet d'une première 1. Variables importantes ? Tout d'abord, parce qu'on a voulu étudier l'impact potentiel d'un grand nombre de variables (climatiques, génétiques) sur la sénescence ou le PMG, une méthode de machine learning a été employée : les modèles Random Forest, Un retard de la sénescence foliaire allonge la durée de la photosynthèse durant le remplissage des grains, accroît la disponibilité des ressources et constitue un trait potentiel de tolérance, 2001.

, ils peuvent identifier les variables importantes et il est même possible de les utiliser pour fournir des prédictions, mais on n'a pas accès à la nature de la relation entre la variable réponse et les variables explicatives (comment varie y en fonction de x ?, Par conséquent, des modèles de régressions multiples ont été ajustés estimant les relations linéaires entre sénescence ou PMG et les variables explicatives importantes selon les modèles Random Forest

, En e et, le grain est composé de 10 à 15% de protéines. L'essentiel du flux d'azote vers le grain après floraison est alimenté par la remobilisation depuis les organes végétatifs (e.g. tiges, feuilles). Or, l'essentiel de l'azote est également associé aux fonctions photosynthétiques, 1989.

. Zhang, L'objectif principal de l'essai était d'étudier le lien entre tolérance et métabolisme azoté. Un essai en conditions contrôlées (serre + semi-hydroponie), l'azote et du carbone après floraison sont donc liés, de leur interaction dépend le rendement en grains, 2014.

, L'essai en serre s'appuie sur quatre hypothèses

, ? la tolérance reposant sur des équilibres source/puits, il est possible de produire en serre des couverts semblables à ceux généralement obtenus au champ

, ? les variations génotypiques observées au champ se traduisent par des variations génotypiques cohérentes en serre

, ? les traits des génotypes associés à une faible limitation du remplissage des grains par les sources sont des traits de tolérance potentiels

, ? les variations de tolérance à la septoriose dépendent du génotype et sont liées au métabolisme azoté

, Au stade GS44, immédiatement après un échantillon initial, l'alimentation azotée a été stoppée pour la moitié des plantes et suivie par une inoculation au pinceau des deux feuilles supérieures par une solution de spores de Septoria tritici. Ensuite, une analyse de la croissance a été réalisée, en suivant l'évolution des surfaces vertes, les masses de biomasses sèches, Quatre génotypes contrastés pour la tolérance parmi les six utilisés dans l'essai au champ 2014-15 (C2015) ont été exploités ici

. Demotes-mainard, L'inoculation n'a pas généré de maladie mais a induit un égrainage physiologique expliqué par l'avortement de fleurs probablement associé à l'extinction des lampes que la méthode d'inoculation nécessitait, 1980.

, (C2015), l'égrainage a été le principal facteur générant une gamme étendue d'équilibres source/puits. En C2015, les puits (les épis) ont été diminués à l'échelle du brin et de la culture par le traitement d'égrainage alors que le rapport source/puits a été, Les trois expérimentations ont généré de nouvelles données sources/puits aux échelles du brin, du grain ou de la culture, pp.2014-2029

, la gamme d'équilibres source/puits a été accentuée par la modalité d'arrêt de la nutrition azotée à GS44 en réduisant la disponibilité des sources durant la période post-épiaison (IFVI) sans modifier le nombre de grains par épi. Dans les deux essais C2015 et S2015 les modalités de contraste de septoriose (fongicide et inoculation, respectivement) n'ont pas permis d'obtenir une sévérité de maladie su sante pour qu'elles soient utilisées en tant que telles, Au contraire, les traitements fongicides appliqués dans l'essai au champ 2015-2016 (C2016) ont abouti à de forts contrastes entre microparcelles saines et malades

, En C2016, compte tenu des IFVI élevés, les rendements obtenus étaient eux aussi élevés et proches du plateau théorique conformément aux attentes pour des cultivars à haut potentiel en conditions de cultures intensives (Gouache References le PMG. C'est bien la prise en compte de E qui a permis l'identification de traits, tels que la date d'épiaison, les sources (IFVI) obtenues en S2015 étaient comparables aux données obtenue en C2015

, Pourtant, la tolérance serait également une propriété émanant de l'environnement : certains environnements étant plus propices à l'expression de tolérance que d'autres. Par exemple, si la précocité épiaison (probable trait de tolérance) varie significativement avec le géno-References AHDB. Ahdb Recommended List -winter wheat 2016/17. tables, La tolérance a été étudiée ici dans l'objectif d'identifier des traits de tolérance, 2016.

I. J. Bingham, D. R. Walters, M. J. Foulkes, and N. D. Paveley, Crop traits and tolerance of wheat and barley to foliar disease, Annals of Applied Biology, vol.154, pp.159-173, 2009.

A. Blum, Improving wheat grain filling under stress by stem reserve mobilisation, Wheat: Prospects for Global Improvement, pp.135-141, 1997.

L. Breiman, Random forests, Machine Learning, vol.45, pp.5-32, 2001.

J. Burke and B. Dunne, Septoria tritici in winter wheat -to spray or not to spray? Irish farmer, pp.14-18, 2006.

N. A. Cobb, Contributions to an economic knowledge of Australian rusts (Uredineae), chapter Chapter, Agricultural Gazette of New South Wales, pp.239-250, 1894.

M. J. Crawley, The R Book, ects Models, pp.681-714, 2012.

S. Demotes-mainard, G. Doussinault, and J. M. Meynard, E ects of low radiation and low temperature at meiosis on pollen viability and grain set in wheat, Agronomie, vol.15, pp.357-365, 1995.

S. Demotes-mainard, G. Doussinault, and J. M. Meynard, Abnormalities in the male developmental programme of winter wheat induced by climatic stess at meiosis, vol.15, pp.357-365, 1996.

R. T. Ellis, Tolerance to the maize rust Puccinia polysora underw, Nature, vol.174, p.1021, 1954.

J. R. Evans, Photosynthesis and nitrogen relationships in leaves of C3 plants, Oecologia, vol.78, pp.9-19, 1989.

R. A. Fischer and Y. M. Stockman, Kernel number per spike in wheat (Triticum aestivum l.): responses to preanthesis shading, Australian Journal of Plant Physiology, vol.7, issue.2, pp.169-180, 1980.

M. J. Foulkes, N. D. Paveley, A. Worland, S. J. Welham, J. Thomas et al., Major genetic changes in wheat with potential to a ect disease tolerance, Phytopathology, vol.96, pp.680-688, 2006.

D. Gouache, M. Bancal, B. De-solan, and P. Gate, Tolérance du blé tendre aux stress biotiques et abiotiques. Innovations Agronomiques, vol.35, pp.75-87, 2014.

. Hgca and . Hgca, , vol.16, 2015.

S. J. Inglese and N. D. Paul, Tolerance of Senecio vulgaris to infection and disease caused by native and alien rust fungi, Phytopathology, vol.96, issue.7, pp.718-726, 2006.

E. Justes, B. Mary, J. M. Meynard, J. M. Machet, and L. Thelierhuche, Determination of a critical nitrogen dilution curve for winter-wheat crops, Annals of Botany, vol.74, issue.4, pp.397-407, 1994.

T. Kramer, B. H. Gildemacher, M. Van-der, J. E. Ster, and . Parlevliet, Tolerance of spring barley cultivars to leaf rust, Puccinia hordei, Euphytica, vol.29, issue.2, pp.209-216, 1980.

S. R. Parker, S. Welham, N. Paveley, J. M. Foulkes, and R. K. Scott, Tolerance of Septoria leaf blotch in winter wheat, Plant Pathology, vol.53, issue.1, pp.1-10, 2004.

N. D. Paveley, R. Sylvester-bradley, R. K. Scott, J. Craigon, and W. Day, Steps in predicting the relationship of yield on fungicide dose, Phytopathology, vol.91, issue.7, pp.708-716, 2001.

, RAGT. Blé tendre d'hiver. Leaflet, 2016.

J. Schafer, Annual review of phytopathology, vol.9, pp.235-252, 1971.

R. A. Serrago, R. Carretero, M. Bancal, and D. J. Miralles, Foliar diseases a ect the eco-physiological attributes linked with yield and biomass in wheat (Triticum aestivum L.), European Journal of Agronomy, vol.31, issue.4, pp.195-203, 2009.

F. Van-den, N. D. Berg, I. J. Paveley, F. Bingham, . Van-den et al., Physiological traits determining yield tolerance of wheat to foliar diseases. Phytopathology, 2017.

M. Zilberstein, A. Blum, and Z. Eyal, Chemical desiccation of wheat plants as a simulator of postanthesis speckled leaf blotch stress, Phytopathology, vol.75, issue.2, pp.226-230, 1985.

E. Zuckerman, A. Eshel, and Z. Eyal, Physiological aspects related to tolerance of spring wheat cultivars to Septoria tritici blotch, Phytopathology, vol.87, issue.1, pp.60-65, 1997.

T. B. Adhikari, J. M. Anderson, and S. B. Goodwin, Identification and molecular mapping of a gene in wheat conferring resistance to Mycosphaerella graminicola, Phytopathology, vol.93, issue.9, pp.1158-1164, 2003.

T. B. Adhikari, J. R. Cavaletto, J. Dubcovsky, J. O. Gieco, A. R. Schlatter et al., Molecular mapping of the Stb4 gene for resistance to Septoria tritici blotch in wheat, Phytopathology, 2004.

. Ahdb, Wheat disease maagement guide. AHDB, 2016.

. Ahdb, Ahdb Recommended List -winter wheat 2016/17. tables, 2016.

. Ahdb and . Ahdb, , pp.2017-2025, 2017.

K. Ando, R. Grumet, K. Terpstra, and J. D. Kelly, Manipulation of plant architecture to enhance crop disease control. CABI, 2(26):8 pp, 2007.

J. Angus, R. Jones, and J. Wilson, A comparison of barley cultivars with di erent leaf inclinations, Crop and Pasture Science, vol.23, pp.945-957, 1972.

I. Aranjuelo, L. Cabrera-bosquet, R. Morcuende, J. C. Avice, S. Nogués et al., Does ear C sink strength contribute to overcoming photosynthetic acclimation of wheat plants exposed to elevated CO2?, Journal of Experimental Botany, vol.62, issue.11, pp.3957-3969, 2011.

I. Aranjuelo, L. Cabrera-bosquet, J. L. Araus, and S. Noguès, Carbon and nitrogen partitioning during the post-anthesis period is conditioned by N fertilisation and sink strength in three cereals, Plant Biology, vol.15, issue.1, pp.135-143, 2013.

L. Arraiano, A. Worland, C. Ellerbrook, and J. Brown, Chromosomal location of a gene for resistance to Septoria tritici blotch (Mycosphaerella graminicola) in the hexaploid wheat 'Synthetic 6x, TAG Theoretical and Applied Genetics, vol.103, issue.5, pp.758-764, 2001.

R. B. Austin, J. Bingham, R. D. Blackwell, L. T. Evans, M. A. Ford et al., Genetic improvements in winter wheat yields since 1900 and associated physiological changes, The Journal of Agricultural Science, vol.94, issue.3, pp.675-689, 1980.

R. Baccar, C. Fournier, T. Dornbusch, B. Andrieu, D. Gouache et al., Modelling the e ect of wheat canopy architecture as a ected by sowing density on Septoria tritici epidemics using a coupled epidemic × virtual plant model, Annals of Botany, vol.108, issue.6, pp.1179-1194, 2011.

B. Baltazar, A. Scharen, and W. Kronstad, Association between dwarfing genes 'rht1'and 'rht2'and resistance to Septoria tritici blotch in winter wheat (Triticum aestivum L. em Thell). Theoretical and applied genetics, vol.79, pp.422-426, 1990.

M. Bancal, R. Roche, and P. Bancal, Late foliar diseases in wheat crops decrease nitrogen yield through N uptake rather than through variations in N remobilization, Annals of Botany, vol.102, issue.4, pp.579-590, 2008.
URL : https://hal.archives-ouvertes.fr/hal-01192195

P. Bancal, Positive contribution of stem growth to grain number per spike in wheat, Field Crops Research, vol.105, issue.1-2, pp.27-39, 2008.
URL : https://hal.archives-ouvertes.fr/hal-01191988

P. Bancal, Data: Précocité et surface de feuille étendard, 2017.

A. Barbottin, C. Lecomte, C. Bouchard, and M. H. Roy, Nitrogen remobilization during grain filling in wheat: Genotypic and environmental e ects, Crop Science, vol.45, issue.3, pp.1141-1150, 2005.

A. J. Barneix, Physiology and biochemistry of source-regulated protein accumulation in the wheat grain, Journal of plant physiology, vol.164, issue.5, pp.581-590, 2007.

L. Bastiaans, Ratio between virtual and visual lesion size as a measure to describe reduction in leaf photosynthesis of rice due to leaf blast, Phytopathology, vol.81, pp.611-615, 1991.

L. Bastiaans, E ects of leaf blast on growth and production of a rice crop. 1. Determining the mechanism of yield reduction. Netherland journal of plant pathology, vol.99, pp.323-334, 1993.

D. Bates, M. Mächler, B. Bolker, and S. Walker, Fitting linear mixed-e ects models using lme4, Journal of Statistical Software, vol.67, issue.1, pp.1-48, 2015.

R. and B. Slimane, E ets de la septoriose foliaire sur la sénescence et les ux d'azote pendant le remplissage des grains chez le blé tendre, AgroParisTech, 2010.