M. Abdallah, L. Dubousset, F. Meuriot, E. P. Avice, J. Ourry et al., Effect of mineral sulphur availability on nitrogen and sulphur uptake and remobilization during the vegetative growth of Brassica napus L, Journal of Experimental Botany, vol.61, pp.2635-2646, 2010.

Z. Adam, I. Adamska, and K. Nakabayashi, Chloroplast and mitochondrial proteases in Arabidopsis. A proposed nomenclature, Plant Physiology, vol.125, pp.1912-1918, 2001.

Z. Ada, . Zalts-a-a, G. Si-a--villalo-o, S. , and W. , FtsH proteases in chloroplasts and cyanobacteria, Physiologia Plantarum, vol.123, pp.386-390, 2005.

P. Annamalai and S. Yanagihara, Identification and characterization of a heat-stress induced gene in cabbage Encodes a Künitz type protease inhibitor, Journal of Plant Physiology, vol.155, pp.226-233, 1999.

M. Arkoun, X. Sarda, L. Jannin, P. Laîné, E. P. et al., Hydroponics versus field lysimeter studies of urea, ammonium and nitrate uptake by oilseed rape(Brassica napus L.), Journal of Experimental Botany, vol.63, pp.5245-5258, 2012.

J. Avice and E. P. , Leaf senescence and nitrogen remobilization efficiency in oilseed rape (Brassica napus L.), Journal of Experimental Botany, vol.65, pp.3813-3824, 2014.

L. Avila-ospina, M. Moison, K. Yoshimoto, and C. Masclaux-daubresse, Autophagy, plant senescence, and nutrient recycling, Journal of Experimental Botany, vol.65, pp.3799-3811, 2014.
URL : https://hal.archives-ouvertes.fr/hal-01204073

S. Balazadeh, J. Schildhauer, W. L. Araújo, S. Munné-bosch, A. R. Fernie et al., Reversal of senescence by N resupply to N-starved Arabidopsis thaliana: transcriptomic and metabolomic consequences, Journal of Experimental Botany, vol.65, pp.3975-3992, 2014.

T. Balint and Z. Rengel, Nitrogen and sulfur uptake and remobilisation in canola genotypes with varied N-and S-use efficiency differ at vegetative and maturity stages, Crop and Pasture Science, vol.62, pp.299-312, 2011.

S. Basu, V. Ramegowda, A. Kumar, and A. Pereira, Plant adaptation to drought stress, 2016.

E. P. Beers, A. M. Jones, and A. W. Dickerman, The S8 serine, C1A cysteine and A1 aspartic protease families in Arabidopsis, Phytochemistry, vol.65, pp.43-58, 2004.

E. P. Beers, B. J. Woffenden, and C. Zhao, Plant proteolytic enzymes: possible roles during programmed cell death, Plant Molecular Biology, vol.44, pp.399-415, 2000.

F. Bellegarde, A. Gojon, and A. Martin, Signals and players in the transcriptional regulation of root responses by local and systemic N signaling in Arabidopsis thaliana, Journal of Experimental Botany, vol.68, pp.2553-2565, 2017.
URL : https://hal.archives-ouvertes.fr/hal-01534644

P. M. Berry, J. Spink, M. J. Foulkes, and P. J. White, The physiological basis of genotypic differences in nitrogen use efficiency in oilseed rape (Brassica napus L.), Field Crops Research, vol.119, pp.365-373, 2010.

R. Bhalerao, J. Keskitalo, and F. Sterky, Gene Expression in Autumn Leaves. Plant Physiology, vol.131, pp.430-442, 2003.

E. Boex-fontvieille, S. Rustgi, D. Von-wettstein, S. Reinbothe, and C. Reinbothe, , 2015.

B. Bollhöner, B. Zhang, S. Stael, N. Denancé, K. Overmyer et al., Post mortem function of AtMC9 in xylem vessel elements, New Phytologist, vol.200, pp.498-510, 2013.

A. Bouchet, A. Laperche, C. Bissuel-belaygue, R. Snowdon, N. Nesi et al., Nitrogen use efficiency in rapeseed. A review. Agronomy for Sustainable Development 36, 2016.
URL : https://hal.archives-ouvertes.fr/hal-01462752

M. M. Bradford, A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding, Analytical Biochemistry, vol.72, pp.248-254, 1976.

E. Breeze, E. Harrison, and S. Mchattie, High-resolution temporal profiling of transcripts during Arabidopsis leaf senescence reveals a distinct chronology of processes and regulation, The Plant Cell, vol.23, pp.873-894, 2011.

V. Buchanan-wollaston, The molecular biology of leaf senescence, Journal of Experimental Botany, vol.48, pp.181-199, 1997.

V. Buchanan-wollaston and C. Ainsworth, Leaf senescence in Brassica napus: cloning of senescence related genes by subtractive hybridisation, Plant Molecular Biology, vol.33, pp.821-834, 1996.

V. Buchanan-wollaston, S. Earl, E. Harrison, E. Mathas, S. Navabpour et al., The molecular analysis of leaf senescence -a genomics approach, Plant Biotechnology Journal, vol.1, pp.3-22, 2003.

V. Buchanan-wollaston, T. Page, and E. Harrison, Comparative transcriptome analysis reveals significant differences in gene expression and signalling pathways between developmental and dark/starvation-induced senescence in Arabidopsis, The Plant Journal: For Cell and Molecular Biology, vol.42, pp.567-585, 2005.

C. A. C-carrión, M. L. Costa, D. E. Martínez, C. Mohr, K. Humbeck et al., , 2013.

C. A. Carrión, D. E. Martínez, M. L. Costa, and J. J. Guiamet, Senescence-Associated Vacuoles, a specific lytic compartment for degradation of chloroplast proteins? Plants, vol.3, pp.498-512, 2014.

A. Cartelat, Z. G. Cerovic, and Y. Goulas, Optically assessed contents of leaf polyphenolics and chlorophyll as indicators of nitrogen deficiency in wheat (Triticum aestivum L.), Field Crops Research, vol.91, pp.35-49, 2005.

B. Chalhoub, F. Denoeud, and S. Liu, Early allopolyploid evolution in the post-Neolithic Brassica napus oilseed genome, Science, vol.345, pp.950-953, 2014.
URL : https://hal.archives-ouvertes.fr/hal-01208705

S. Chauhan, S. Srivalli, A. R. Nautiyal, and R. Khanna-chopra, , 2009.

J. Chen, J. J. Burke, J. Velten, and Z. Xin, FtsH11 protease plays a critical role in Arabidopsis thermotolerance, The Plant Journal: For Cell and Molecular Biology, vol.48, pp.73-84, 2006.

M. Chen, A. Maodzeka, L. Zhou, A. E. Wang, Z. Jiang et al., Removal of DELLA repression promotes leaf senescence in Arabidopsis, Plant Science, vol.219, issue.220, pp.26-34, 2014.

L. Chen, S. Xiang, Y. Chen, D. Li, and D. Yu, Arabidopsis WRKY45 interacts with the DELLA protein RGL1 to positively regulate age-triggered leaf senescence, Molecular Plant, vol.10, pp.1174-1189, 2017.

W. Chi, X. Sun, and L. Zhang, The roles of chloroplast proteases in the biogenesis and maintenance of photosystem II, Biochimica et Biophysica Acta (BBA) -Bioenergetics, vol.1817, pp.239-246, 2012.

A. Chiba, H. Ishida, N. K. Nishizawa, A. Makino, and T. Mae,

N. V. Chichkova, J. Shaw, and R. A. Galiullina, Phytaspase, a relocalisable cell death promoting plant protease with caspase specificity, The EMBO Journal, vol.29, pp.1149-1161, 2010.

K. Cho, L. Mabasa, A. W. Fowler, D. M. Walsh, and C. S. Park, Canola oil inhibits breast cancer cell growth in cultures and in vivo and acts synergistically with chemotherapeutic drugs, Lipids, vol.45, pp.777-784, 2010.

P. Chouard, Vernalization and its relations to dormancy, Annual Review of Plant Physiology, vol.11, pp.191-238, 1960.

D. Chrobok, S. R. Law, and B. Brouwer, Dissecting the metabolic role of mitochondria during developmental leaf senescence, Plant Physiology, vol.172, pp.2132-2153, 2016.
URL : https://hal.archives-ouvertes.fr/hal-01494737

S. J. Clough and A. F. Bent, Floral dip: a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana, The Plant Journal: For Cell and Molecular Biology, vol.16, pp.735-743, 1998.

C. Colnenne, J. M. Meynard, R. Reau, E. Justes, and A. Merrien, Determination of a critical nitrogen dilution curve for winter oilseed rape, Annals of Botany, vol.81, pp.311-317, 1998.

S. A. Coupe, B. K. Sinclair, L. M. Watson, J. A. Heyes, and J. R. Eason,

, Ide tifi atio of deh d atio -espo si e

N. M. Crawford and B. G. Forde, Molecular and Developmental Biology of Inorganic Nitrogen Nutrition. The Arabidopsis Book, vol.1, 2002.

N. M. Crawford and A. Glass, Molecular and physiological aspects of nitrate uptake in plants, Trends in Plant Science, vol.3, pp.389-395, 1998.

P. L. Curci, A. Cigliano, R. Zuluaga, D. L. Janni, M. Sanseverino et al., Transcriptomic response of durum wheat to nitrogen starvation, Scientific Reports, vol.7, 2017.

S. Gan and R. M. Amasino, Inhibition of Leaf Senescence by Autoregulated Production of Cytokinin, Science, vol.270, pp.1986-1988, 1995.

S. Gan and R. M. Amasino, Making Sense of Senescence (Molecular Genetic Regulation and Manipulation of Leaf Senescence), Plant Physiology, vol.113, pp.313-319, 1997.

L. Gent and B. G. Forde, How do plants sense their nitrogen status, Journal of Experimental Botany, vol.68, pp.2531-2539, 2017.

A. Girondé, L. Dubousset, J. Trouverie, E. P. Avice, and J. ,

A. Girondé, P. Etienne, and J. Trouverie,

A. Girondé, M. Poret, and P. Etienne, A profiling approach of the natural variability of foliar, Botany, vol.66, pp.2461-2473, 2015.

A. Girondé, M. Poret, E. P. Trouverie, J. Bouchereau, A. et al., A Comparative Study of Proteolytic Mechanisms during Leaf Senescence of Four Genotypes of Winter Oilseed Rape Highlighted Relevant Physiological and Molecular Traits for NRE Improvement, vol.5, p.1, 2016.

D. Golldack, O. V. Popova, and K. Dietz, Mutation of the matrix metalloproteinase At2-MMP inhibits growth and causes late flowering and early senescence in Arabidopsis, The Journal of Biological Chemistry, vol.277, pp.5541-5547, 2002.

J. Gombert, F. L. Dily, J. Lothier, E. P. Rossato, L. Allirand et al., Effect of nitrogen fertilization on nitrogen dynamics in oilseed rape using 15N-labeling field experiment, Journal of Plant Nutrition and Soil Science, vol.173, pp.875-884, 2010.
URL : https://hal.archives-ouvertes.fr/hal-01168762

G. and V. , SAG2 and SAG12 protein expression in senescing Arabidopsis plants, Physiologia Plantarum, vol.119, pp.263-269, 2003.

D. C. Greenbaum, A. Baruch, M. Grainger, Z. Bozdech, K. F. Medzihradszky et al., A Role for the Protease Falcipain 1 in Host Cell Invasion by the Human Malaria Parasite, Science, vol.298, pp.2002-2006, 2002.

P. L. Gregersen, Senescence and nutrient remobilization in crop plants. The Molecular and Physiological Basis of Nutrient Use Efficiency in Crops, pp.83-102, 2011.

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

P. L. Gregersen and P. B. Holm, Transcriptome analysis of senescence in the flag leaf of wheat, Triticum aestivum L.). Plant Biotechnology Journal, vol.5, pp.192-206, 2007.

A. Guiboileau, L. Avila-ospina, K. Yoshimoto, F. Soulay, M. Azzopardi et al., Physiological and metabolic consequences of autophagy deficiency for the management of nitrogen and protein resources in Arabidopsis leaves depending on nitrate availability, The New Phytologist, vol.199, pp.683-694, 2013.
URL : https://hal.archives-ouvertes.fr/hal-01001035

A. Guiboileau, R. Sormani, C. Meyer, and C. Masclaux-daubresse, Senescence and death of plant organs: Nutrient recycling and developmental regulation, Comptes Rendus Biologies, vol.333, pp.382-391, 2010.
URL : https://hal.archives-ouvertes.fr/hal-01203896

A. Guiboileau, K. Yoshimoto, F. Soulay, M. Bataillé, J. Avice et al., Autophagy machinery controls nitrogen remobilization at the whole-plant level under both limiting and ample nitrate conditions in Arabidopsis, The New Phytologist, vol.194, pp.732-740, 2012.
URL : https://hal.archives-ouvertes.fr/hal-01004199

Y. Guo, Z. Cai, and S. Gan, Transcriptome of Arabidopsis leaf senescence, Plant, Cell & Environment, vol.27, pp.521-549, 2004.

Y. Guo and S. Gan, Leaf senescence: signals, execution, and regulation, Current Topics in Developmental Biology, vol.71, pp.83-112, 2005.

Y. Guo and S. Gan, AtNAP, a NAC family transcription factor, has an important role in leaf senescence, The Plant Journal: For Cell and Molecular Biology, vol.46, pp.601-612, 2006.

H. Ishida, K. Yoshimoto, M. Izumi, D. Reisen, Y. Yano et al., , 2008.

J. Lequeu, F. Simon-plas, J. Fromentin, E. P. Petitot, A. Blein et al.,

W. Li, H. Zhang, and X. Li, Intergrative metabolomic and transcriptomic analyses unveil nutrient remobilization events in leaf senescence of tobacco, Scientific Reports, vol.7, p.12126, 2017.

P. O. Lim, H. J. Kim, and H. G. Nam, Leaf Senescence, Annual Review of Plant Biology, vol.58, pp.115-136, 2007.

J. Lin and S. Wu, Molecular events in senescing Arabidopsis leaves, The Plant Journal: For Cell and Molecular Biology, vol.39, pp.612-628, 2004.

L. Liu, Y. Zhou, G. Zhou, R. Ye, L. Zhao et al., Identification of early senescence-associated genes in rice flag leaves, Plant Molecular Biology, vol.67, pp.37-55, 2008.

K. J. Livak and T. D. Schmittgen, Analysis of relative gene expression data using real-time quantitative PCR a d the ???CT Method, Methods, vol.25, pp.402-408, 2001.

K. N. Lohman, S. Gan, M. C. John, and R. M. Amasino, Molecular analysis of natural leaf senescence in Arabidopsis thaliana, Physiologia Plantarum, vol.92, pp.322-328, 1994.

O. Loudet, S. Chaillou, C. Camilleri, D. Bouchez, and F. Daniel-vedele, Bay-0 x Shahdara recombinant inbred line population: a powerful tool for the genetic dissection of complex traits in Arabidopsis. TAG. Theoretical and applied genetics, Theoretische und angewandte Genetik, vol.104, pp.1173-1184, 2002.

H. Lu, B. Chandrasekar, J. Oeljeklaus, J. C. Misas-villamil, Z. Wang et al., Subfamily-specific fluorescent probes for cysteine proteases display dynamic protease activities during seed germination, Plant Physiology, vol.168, pp.1462-1475, 2015.

R. Lu-i?ski, L. Misztal, S. Sa-a-dakie-i-z, J. Ko-ski, and G. , Involvement of Deg5 protease in woundingrelated disposal of PsbF apoprotein, Plant Physiology and Biochemistry, vol.49, pp.311-320, 2011.

R. Lu-i?ski, L. Misztal, S. Sa-a-dakie-i-z, J. Ko-ski, and G. , The thylakoid protease Deg2 is involved in stress-related degradation of the photosystem II light-harvesting protein Lhcb6 in Arabidopsis thaliana, The New Phytologist, vol.192, pp.74-86, 2011.

P. Malagoli, P. Lainé, L. Deunff, E. Rossato, L. Ney et al., , 2004.

P. Malagoli, P. Laine, L. Rossato, and A. Ourry, Dynamics of nitrogen uptake and mobilization in fieldgrown winter oilseed rape (Brassica napus) from stem extension to Hharvest I. global N flows between vegetative and reproductive tissues in relation to leaf fall and their residual N, Annals of Botany, vol.95, pp.853-861, 2005.

P. Malagoli, P. Lainé, L. Rossato, and A. Ourry, Dynamics of nitrogen uptake and mobilization in fieldgrown winter oilseed rape (Brassica napus) From stem extension to harvest. II. An 15N-labelling-based simulation model of N partitioning between vegetative and reproductive tissues, Annals of Botany, vol.95, pp.1187-1198, 2005.
URL : https://hal.archives-ouvertes.fr/hal-01422741

G. Marino and C. Funk, Matrix metalloproteinases in plants: a brief overview, Physiologia Plantarum, vol.145, pp.196-202, 2011.

D. E. Martínez, C. G. Bartoli, V. Grbic, and J. J. Guiamet, Vacuolar cysteine proteases of wheat (Triticum aestivum L.) are common to leaf senescence induced by different factors, Journal of Experimental Botany, vol.58, pp.1099-1107, 2007.

D. E. Martinez, M. L. Borniego, N. Battchikova, E. Aro, E. Tyystjärvi et al., SASP, a SenescenceAssociated Subtilisin Protease, is involved in reproductive development and determination of silique number in Arabidopsis, Journal of Experimental Botany, vol.66, pp.161-174, 2015.

D. E. Martínez, M. L. Costa, F. M. Gomez, M. S. Otegui, and J. J. Guiamet, Se es e e-asso iated a uoles a e involved in the degradation of chloroplast proteins in tobacco leaves, The Plant Journal, vol.2, pp.196-206

D. E. Martínez and J. J. Guiamet, Senescence-related changes in the leaf apoplast, Journal of Plant Growth Regulation, vol.33, pp.44-55, 2014.

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

C. Masclaux-daubresse, Q. Chen, and M. Havé, Regulation of nutrient recycling via autophagy, Current Opinion in Plant Biology, vol.39, pp.8-17, 2017.
URL : https://hal.archives-ouvertes.fr/hal-01605933

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, pp.1141-1157, 2010.
URL : https://hal.archives-ouvertes.fr/hal-01203920

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

C. H. Mcallister, M. Wolansky, and A. G. Good, The impact on nitrogen-efficient phenotypes when aspartate aminotransferase is expressed tissue-specifically in Brassica napus, New Negatives in Plant Science 3-4, pp.1-9, 2016.

A. Merrien and A. Pouzet,

, P i ipau fa teu s li ita t les e de e ts de olza d hi e da s les o ditio s

Y. Miao, T. Laun, P. Zimmermann, and U. Zentgraf, Targets of the WRKY53 transcription factor and its role during leaf senescence in Arabidopsis, Plant Molecular Biology, vol.55, pp.853-867, 2004.

S. Michaeli, A. Honig, H. Levanony, H. Peled-zehavi, and G. Galili, Arabidopsis ATG8-INTERACTING PROTEIN1 is involved in autophagy-dependent vesicular trafficking of plastid proteins to the vacuole, The Plant Cell, vol.26, pp.4084-4101, 2014.

A. Migge, E. Carrayol, B. Hirel, and T. W. Becker, Leaf-specific overexpression of plastidic glutamine synthetase stimulates the growth of transgenic tobacco seedlings, Planta, vol.210, pp.252-260, 2000.

J. Moon, G. Parry, and E. M. , The ubiquitin-proteasome pathway and plant development, The Plant Cell, vol.16, pp.3181-3195, 2004.

K. Mo-is, A. Ke-ess, S. Page, T. Joh, C. F. Mu-ph et al., Salicylic acid has a role in regulating gene expression during leaf senescence, The Plant Journal, vol.23, pp.677-685, 2001.

V. V. Mosolov and T. A. Valueva, Proteinase inhibitors and their function in plants: A review, Applied Biochemistry and Microbiology, vol.41, pp.227-246, 2005.

R. Muñoz-huerta, R. Guevara-gonzalez, L. Contreras-medina, I. Torres-pacheco, J. Prado-olivarez et al., A review of methods for sensing the nitrogen status in plants: advantages, disadvantages and recent advances, Sensors, vol.13, pp.10823-10843, 2013.

S. Murakami, Y. Kondo, T. Nakano, and F. Sato, Protease activity of CND41, a chloroplast nucleoid DNAbinding protein, isolated from cultured tobacco cells, FEBS Letters, vol.468, pp.15-18, 2000.

T. Murata, F. Toda, K. Uchino, and E. Yakushiji, Water-soluble chlorophyll protein of Brassica oleracea var. Botrys (cauliflower), Biochimica Et Biophysica Acta, vol.245, pp.208-215, 1971.

P. Nacry, E. Bouguyon, and A. Gojon, Nitrogen acquisition by roots: physiological and developmental mechanisms ensuring plant adaptation to a fluctuating resource, Plant and Soil, vol.370, pp.1-29, 2013.
URL : https://hal.archives-ouvertes.fr/hal-00921012

T. Nakano, S. Murakami, T. Shoji, S. Yoshida, Y. Yamada et al., A novel protein with DNA binding activity from tobacco chloroplast nucleoids, The Plant Cell, vol.9, pp.1673-1682, 1997.

N. Nesi, R. Delourme, M. Brégeon, C. Falentin, and R. M. , Genetic and molecular approaches to improve nutritional value of Brassica napus L. seed, Comptes Rendus Biologies, vol.331, pp.763-771, 2008.

G. R. Newman, Use and abuse of LR White, The Histochemical Journal, vol.19, pp.118-120, 1987.

M. Nishimura, M. Hayashi, A. Kato, K. Yamaguchi, and S. Mano, Functional transformation of microbodies in higher plant cells, Cell Structure and Function, vol.21, pp.387-393, 1996.

Y. S. Noh and R. M. Amasino, Regulation of developmental senescence is conserved between Arabidopsis and Brassica napus, Plant Molecular Biology, vol.41, pp.195-206, 1999.

Y. Noh and R. M. Amasino, Identification of a promoter region responsible for the senescence-specific expression of SAG12, Plant Molecular Biology, vol.41, pp.181-194, 1999.

C. Noquet, J. Avice, L. Rossato, P. Beauclair, M. Henry et al., Effects of altered source-sink relationships on N allocation and vegetative storage protein accumulation in Brassica napus L, Plant Science, vol.166, pp.1007-1018, 2004.

J. Nyikako, A. Schierholt, B. Kessel, and H. C. Becker, Genetic variation in nitrogen uptake and utilization, 2014.

S. A. Oh, J. Park, G. I. Lee, K. H. Paek, S. K. Park et al., Identification of three genetic loci controlling leaf senescence in Arabidopsis thaliana, The Plant Journal, vol.12, pp.527-535, 2003.

M. S. Otegui, Vacuolar degradation of chloroplast components: autophagy and beyond, Journal of Experimental Botany, vol.69, pp.741-750, 2018.

M. S. Otegui, Y. Noh, D. E. Martínez, V. Petroff, M. G. et al., Senescence-associated vacuoles with intense proteolytic activity develop in leaves of Arabidopsis and soybean, The Plant Journal, vol.41, pp.831-844, 2005.

A. Prins, V. Heerden, D. , P. Olmos, E. Kunert et al., Cysteine proteinases regulate chloroplast protein content and composition in tobacco leaves: a model for dynamic interactions with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) vesicular bodies, Journal of Experimental Botany, vol.59, 1935.

P. U?i-ská, A. Shi-do, T. Niesse, S. , K. Ha-i et al., Major Cys, 2017.

B. F. Quirino, Y. Noh, E. Himelblau, and R. M. Amasino, , 2000.

Y. Rahbé, C. Deraison, M. Bonadé-bottino, C. Girard, C. Nardon et al., Effects of the cysteine protease inhibitor oryzacystatin (OC-I) on different aphids and reduced performance of Myzus persicae on OC-I expressing transgenic oilseed rape, Plant Science, vol.164, pp.441-450, 2003.

G. Rathke, T. Behrens, and W. Diepenbrock, Integrated nitrogen management strategies to improve seed yield, oil content and nitrogen efficiency of winter oilseed rape (Brassica napus L.): A review, Ecosystems & Environment, vol.117, pp.80-108, 2006.

C. Rautengarten, D. Steinhauser, D. Büssis, A. Stintzi, A. Schaller et al., Inferring hypotheses on functional relationships of genes: Analysis of the Arabidopsis thaliana subtilase gene family, PLOS Computational Biology, vol.1, p.40, 2005.

N. D. Rawlings, A. J. Barrett, and R. Finn, Twenty years of the MEROPS database of proteolytic enzymes, their substrates and inhibitors, Nucleic Acids Research, vol.44, pp.343-350, 2016.

C. Reinbothe, A. Springer, I. Samol, and S. Reinbothe, Plant oxylipins: role of jasmonic acid during programmed cell death, defence and leaf senescence, The FEBS Journal, vol.276, pp.4666-4681, 2009.

M. P. Reviron, N. Vartanian, M. Sallantin, J. C. Huet, J. C. Pernollet et al., Characterization of a novel protein induced by progressive or rapid drought and salinity in Brassica napus leaves, Plant Physiology, vol.100, pp.1486-1493, 1992.

K. H. Richau, F. Kaschani, M. Verdoes, T. C. Pansuriya, S. Niessen et al., Subclassification and biochemical analysis of plant papain-like cysteine proteases displays subfamily-specific characteristics, 2012.

A. E. Richmond and A. Lang, Effect of kinetin on protein content and survival of detached xanthium leaves, Science, vol.125, pp.650-651, 1957.

L. A. R??o, P. Del, G. M. Sa-dalio, L. M. Se-illa, F. Co-pas et al., The activated oxygen role of peroxisomes in senescence, Plant Physiology, vol.116, pp.1195-1200, 1998.

I. N. Roberts, C. Caputo, M. V. Criado, and C. Funk, Senescence-associated proteases in plants, Physiologia Plantarum, vol.145, pp.130-139, 2012.

I. N. Roberts, C. Caputo, M. Kade, M. V. Criado, and A. J. Barneix, Subtilisin-like serine proteases involved in N remobilization during grain filling in wheat, Acta Physiologiae Plantarum, vol.33, 1997.

I. Roberts, P. F. Murray, S. Passeron, and A. J. Barneix, The activity of the 20S proteasome is maintained in detached wheat leaves during senescence in darkness, Plant Physiology and Biochemistry, vol.40, pp.161-166, 2002.

I. N. Roberts, S. Passeron, and A. J. Barneix, The two main endoproteases present in dark-induced senescent wheat leaves are distinct subtilisin-like proteases, Planta, vol.224, pp.1437-1447, 2006.

I. N. Roberts, C. G. Veliz, M. V. Criado, A. Signorini, E. Simonetti et al., Identification and expression analysis of 11 subtilase genes during natural and induced senescence of barley plants, Journal of Plant Physiology, vol.211, pp.70-80, 2017.

E. Rojo, J. Zouhar, C. Carter, V. Kovaleva, and N. V. Raikhel, A unique mechanism for protein processing and degradation in Arabidopsis thaliana, Proceedings of the National Academy of Sciences of the United States of America, vol.100, pp.7389-7394, 2003.

L. Rossato, P. Lainé, and A. Ourry, Nitrogen storage and remobilization in Brassica napus L. during the growth cycle: nitrogen fluxes within the plant and changes in soluble protein patterns, Journal of Experimental Botany, vol.52, pp.1655-1663, 2001.

L. Rossato, J. H. Macduff, P. Laine, L. Deunff, E. Ourry et al., Nitrogen storage and remobilization in Brassica napus L. during the growth cycle: effects of methyl jasmonate on nitrate uptake, senescence, growth, and VSP accumulation, Journal of Experimental Botany, vol.53, pp.1131-1141, 2002.

J. W. Rouse, Monitoring vegetation systems in the Great Plains with ERTS, pp.309-317, 1974.

P. Runeberg-roos and M. Saarma, Phytepsin, a barley vacuolar aspartic proteinase, is highly expressed during autolysis of developing tracheary elements and sieve cells, The Plant Journal: For Cell and Molecular Biology, vol.15, pp.139-145, 1998.

S. Rustgi, E. Boex-fontvieille, C. Reinbothe, D. Von-wettstein, and S. Reinbothe, Serpin1 and WSCP differentially regulate the activity of the cysteine protease RD21 during plant development in Arabidopsis thaliana, Proceedings of the National Academy of Sciences of the United States of America, vol.114, pp.2212-2217, 2017.

S. Rustgi, E. Boex-fontvieille, C. Reinbothe, D. Von-wettstein, and S. Reinbothe, The complex world of plant protease inhibitors: Insights into a Kunitz-type cysteine protease inhibitor of Arabidopsis thaliana, Communicative & Integrative Biology, vol.11, 2017.

S. A. Ruuska, D. C. Lewis, G. Kennedy, R. T. Furbank, C. Jenkins et al., , 2008.

V. Safavi-rizi, J. Franzaring, A. Fangmeier, and R. Kunze, Divergent N Deficiency-Dependent Senescence and Transcriptome Response in Developmentally Old and Young Brassica napus Leaves, Frontiers in Plant Science, vol.9, 2018.

H. Satoh, A. Uchida, K. Nakayama, and M. Okada, Water-soluble chlorophyll protein in Brassicaceae plants is a stress-induced chlorophyll-binding protein, Plant and Cell Physiology, vol.42, pp.906-911, 2001.

A. Schaller, A cut above the rest: the regulatory function of plant proteases, Planta, vol.220, pp.183-197, 2004.

S. Schiltz, K. Gallardo, M. Huart, L. Negroni, N. Sommerer et al., Proteome reference maps of vegetative tissues in pea. An investigation of nitrogen mobilization from leaves during seed filling, Plant Physiology, vol.135, pp.2241-2260, 2004.

J. K. Schjoerring, J. Bock, L. Gammelvind, C. R. Jensen, and M. Vo, Nitrogen incorporation and remobilization in different shoot components of field-grown winter oilseed rape (Brassica napus), Plant and Soil, vol.177, pp.255-264, 1995.

K. Schmidt, C. Fufezan, A. Krieger-liszkay, H. Satoh, and H. Paulsen, Recombinant water-soluble chlorophyll protein from Brassica oleracea Var. Botrys binds various chlorophyll derivatives, Biochemistry, vol.42, pp.7427-7433, 2003.

C. A. Schneider, W. S. Rasband, and K. W. Eliceiri, NIH Image to ImageJ: 25 years of image analysis, Nature Methods, vol.9, pp.671-675, 2012.

H. Schuhmann and I. Adamska, Deg proteases and their role in protein quality control and processing in different subcellular compartments of the plant cell, Physiologia Plantarum, vol.145, pp.224-234, 2012.

G. Le, T. Beh-e-s, A. Ulas, F. Wiesle, and W. J. Ho-st, Agronomic traits contributing to nitrogen efficiency of winter oilseed rape cultivars, Field Crops Research, vol.124, pp.114-123, 2011.

M. A. Seltmann, N. E. Stingl, J. K. Lautenschlaeger, M. Krischke, M. J. Mueller et al., Differential impact of lipoxygenase 2 and jasmonates on natural and stress-induced senescence in Arabidopsis, Plant Physiology, vol.152, pp.1940-1950, 2010.

U. Singh, Integrated nitrogen fertilization for intensive and sustainable agriculture, Journal of Crop Improvement, vol.15, pp.259-288, 2006.

S. Singh, M. K. Giri, P. K. Singh, A. Siddiqui, and A. K. Nandi, Down-regulation of OsSAG12-1 results in enhanced senescence and pathogen-induced cell death in transgenic rice plants, Journal of Biosciences, vol.38, pp.583-592, 2013.

S. Singh, A. Singh, and A. K. Nandi, The rice OsSAG12-2 gene codes for a functional protease that negatively regulates stress-induced cell death, Journal of Biosciences, vol.41, pp.445-453, 2016.

J. Smalle and R. D. Vierstra, The ubiquitin 26s proteasome proteolytic pathway, Annual Review of Plant Biology, vol.55, pp.555-590, 2004.

C. Sorin, M. Musse, M. F. Bouchereau, A. Leport, and L. ,

C. Spitzer, F. Li, R. Buono, H. Roschzttardtz, T. Chung et al.,

, The endosomal protein CHARGED MULTIVESICULAR BODY PROTEIN1 regulates the autophagic turnover of plastids in Arabidopsis, The Plant Cell, vol.27, pp.391-402

D. Stessman, A. Miller, M. Spalding, and S. Rodermel, Regulation of photosynthesis during Arabidopsis leaf development in continuous light, Photosynthesis Research, vol.72, pp.27-37, 2002.

P. Subash-babu and A. A. Alshatwi,

J. A. Sullivan, K. Shirasu, and X. W. Deng, The diverse roles of ubiquitin and the 26S proteasome in the life of plants, Nature Reviews. Genetics, vol.4, pp.948-958, 2003.

X. Sun, T. Fu, N. Chen, J. Guo, J. Ma et al., The stromal chloroplast Deg7 protease participates in the repair of photosystem II after photoinhibition in Arabidopsis, Plant Physiology, vol.152, pp.1263-1273, 2010.

X. Sun, M. Ouyang, J. Guo, J. Ma, C. Lu et al., The thylakoid protease Deg1 is involved in photosystem-II assembly in Arabidopsis thaliana, The Plant Journal, vol.62, pp.240-249, 2010.

L. M. Weaver, S. Gan, B. Quirino, and R. M. Amasino, A comparison of the expression patterns of several senescence-associated genes in response to stress and hormone treatment, Plant Molecular Biology, vol.37, pp.455-469, 1998.

B. Wei, W. Zhang, J. Chao, T. Zhang, T. Zhao et al., Functional analysis of the role of hydrogen sulfide in the regulation of dark-induced leaf senescence in Arabidopsis, Scientific Reports, vol.7, 2017.

F. Wiesler, T. Behrens, and W. J. Horst, Nitrogen efficiency of contrasting rape ideotypes. Developments in Plant and Soil Sciences. Plant Nutrition, pp.60-61, 2001.

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, pp.391-399, 2006.

D. Winter, B. Vinegar, H. Nahal, R. Ammar, G. V. Wilson et al.,

N. Wojciechowska, E. Sobieszczuk-nowicka, and A. Bagniewska-zadworna, Plant organ senescenceregulation by manifold pathways, Plant Biology, 2017.

Y. Xu and B. Huang, Effects of foliar-applied ethylene inhibitor and synthetic cytokinin on creeping bentgrass to enhance heat tolerance, Crop Science, vol.49, pp.1876-1884, 2009.

Y. Yakushiji, E. Uchino, K. Sugimura, Y. Shiratori, I. Takamiya et al., Isolation of water-soluble chlorophyll protein from the leaves of Chenopodium album, Biochimica et Biophysica Acta, vol.75, pp.293-298, 1963.

K. Yamada, R. Matsushima, M. Nishimura, and I. Hara-nishimura, A slow maturation of a cysteine protease with a granulin domain in the vacuoles of senescing Arabidopsis leaves, Plant Physiology, vol.127, pp.1626-1634, 2001.

L. Yang, S. Mickelson, D. See, T. K. Blake, and A. M. Fischer, Genetic analysis of the function of major leaf proteases in barley (Hordeum vulgare L.) nitrogen remobilization, Journal of Experimental Botany, vol.55, pp.2607-2616, 2004.

Z. Zavaleta-mancera, H. A. Thomas, B. J. Thomas, H. Scott, and I. M. , Regreening of senescent Nicotiana leaves: II. Redifferentiation of plastids, Journal of Experimental Botany, vol.50, pp.1683-1689, 1999.

A. ?elisko, M. Ga-Ía-lorenzo, G. Jackowski, S. Jansson, and C. Funk, AtFtsH6 is involved in the degradation of the light-harvesting complex II during high-light acclimation and senescence, Proceedings of the National Academy of Sciences, vol.102, pp.13699-13704, 2005.

D. Zhang, Y. Kato, L. Zhang, M. Fujimoto, N. Tsutsumi et al., The FtsH protease heterocomplex in Arabidopsis: dispensability of Type-B protease activity for proper chloroplast development, The Plant Cell, vol.22, pp.3710-3725, 2010.

X. Zhang, S. Liu, and T. Takano, Two cysteine proteinase inhibitors from Arabidopsis thaliana, AtCYSa and AtCYSb, increasing the salt, drought, oxidation and cold tolerance, Plant Molecular Biology, vol.68, pp.131-143, 2008.

Y. Zhang, Z. Liu, X. Wang, J. Wang, K. Fan et al., DELLA proteins negatively regulate dark-induced, 2018.

M. Zhang, C. Zhang, Y. Jiang, Y. Strasser, R. J. Yuan et al., Changes in chloroplast ultrastructure, fatty acid components of thylakoid membrane and chlorophyll a fluorescence transient in flag leaves of a super-high-yield hybrid rice and its parents during the reproductive stage, Journal of Plant Physiology, vol.167, pp.277-285, 2010.

H. Zhang and C. Zhou, Signal transduction in leaf senescence, Plant Molecular Biology, vol.82, pp.539-545, 2013.

M. Zienkiewicz, A. Ferenc, W. Wasilewska, and E. Romanowska, High light stimulates Deg1-dependent cleavage of the minor LHCII antenna proteins CP26 and CP29 and the PsbS protein in Arabidopsis thaliana, Planta, vol.235, pp.279-288, 2012.

P. Zimmermann and U. Zentgraf, The correlation between oxidative stress and leaf senescence during plant development, Cellular & Molecular Biology Letters, vol.10, pp.515-534, 2005.