Senescence and Nutrient Remobilization in Crop Plants

The Molecular and Physiological Basis of Nutrient Use Efficiency in Crops

Volumn , Issue , 2011, Pages 83-102

  Gregersen, Per L ^a  

^a AARHUS UNIVERSITY   (Denmark)

Author keywords

Cytosolic GS role in nitrogen remobilization, in knockout lines of maize; Efficient remobilization of nitrogen from senescing tissues, nitrogen use efficiency; Environmental modulation of senescence; Fatty acid degradation chloroplast endomembrane system breakdown; Nutrient remobilization core of senescence process; Senescence and nutrient remobilization, crop plants nutrient use efficiency (NUE), into harvested seed; Senescence in plants yellowing of leaves, and necrotization phase; Senescence process, dying off of vegetative plant parts and seed maturation; Senescence regulation, by changes in gene expression; Stay green phenotypes, yellowing of leaves and chlorophyll degradation

Indexed keywords

EID: 84872166550     PISSN: None     EISSN: None     Source Type: Book    
DOI: 10.1002/9780470960707.ch5     Document Type: Chapter

References (144)

1. Balazadeh , S. , Riano - Pachon , D.M. , & Mueller - Roeber , B. ( 2008 ) Transcription factors regulating leaf senescence in Arabidopsis thaliana . Plant Biology 10 , 63 - 75 . Available from: ISI:000258078400007.
2. Benbella, M. & Paulsen, G.M. ( 1998) Effi cacy of treatments for delaying senescence of wheat leaves: II, Senescence and grain yield under fi eld conditions . Agronomy Journal 90 ( 3 ), 332 - 338 . Available from: ISI:000074765600004.
3. Bernard , S.M. & Habash , D.Z. ( 2009 ) The importance of cytosolic glutamine synthetase in nitrogen assimilation and recycling . New Phytologist 182 ( 3 ), 608 - 620 . Available from: ISI:000265229400006.
4. Bernard , S.M. , Moller , A.L.B. , Dionisio , G. , et al. ( 2008 ) Gene expression, cellular localisation and function of glutamine synthetase isozymes in wheat ( Triticum aestivum L.) . Plant Molecular Biology 67 ( 1 - 2 ), 89 - 105 . Available from: ISI:000254965000007.
5. Birsin , M.A. , Adak , M.S. , Inal , A. , Aksu , A. , & Gunes , A. ( 2010 ) Mineral element distribution and accumulation patterns within two barley cultivars . Journal of Plant Nutrition 33 ( 2 ), 267 - 284 . Available at: http://www.informaworld.com/10.1080/019041 60903435391 .
6. Borraccino , G. , Mastropasqua , L. , Deleonardis , S. , & Dipierro , S. ( 1994 ) The role of the ascorbic - acid system in delaying the senescence of oat ( Avena sativa L) leaf segments . Journal of Plant Physiology 144 ( 2 ), 161 - 166 . Available from: ISI:A1994PC76300006.
7. Borrell , A.K. , Hammer , G.L. , & Henzell , R.G. ( 2000 ) Does maintaining green leaf area in sorghum improve yield under drought? II. Dry matter production and yield . Crop Science 40 ( 4 ), 1037 - 1048 . Available at: https://www.crops.org/publications/cs/ abstracts/40/4/1037 .
8. Buchanan - Wollaston , V. ( 1997 ) The molecular biology of leaf senescence . Journal of Experimental Botany 48 ( 307 ), 181 - 199 . Available from: ISI:A1997WL16900001.
9. Buchanan - Wollaston , V. , Earl , S. , Harrison , E. , et al. ( 2003 ) The molecular analysis of leaf senescence - a genomics approach . Plant Biotechnology Journal 1 ( 1 ), 3 - 22 . Available from: ISI:000188993900002.
10. Buchanan - Wollaston , V. , Page , T. , Harrison , E. , et al. ( 2005 ) Comparative transcriptome analysis reveals signifi cant differences in gene expression and signalling pathways between developmental and dark/ starvation - induced senescence in Arabidopsis . Plant Journal 42 ( 4 ), 567 - 585 . Available from: ISI:000228746000011.
11. Cai , H.M. , Zhou , Y. , Xiao , J.H. , Li , X.H. , Zhang , Q.F. , & Lian , X.M. ( 2009 ) Overexpressed glutamine synthetase gene modifi es nitrogen metabolism and abiotic stress responses in rice . Plant Cell Reports 28 ( 3 ), 527 - 537 . Available from: ISI:000263783 200017.
12. Cao , X. , Costa , L.M. , Biderre - Petit , C. , et al. ( 2007 ) Abscisic acid and stress signals induce viviparous1 expression in seed and vegetative tissues of maize . Plant Physiology 143 ( 2 ), 720 - 731 . Available at: http://www.plantphysiol.org/cgi/content/ abstract/143/2/720 .
13. Charlton , W.L. , Johnson , B. , Graham , I.A. , & Baker , A. ( 2005 ) Non - coordinate expression of peroxisome biogenesis, β - oxidation and glyoxylate cycle genes in mature Arabidopsis plants . Plant Cell Reports 23 ( 9 ), 647 - 653 . Available at: http:// dx.doi.org/10.1007/s00299 - 004 - 0879 - 7 .
14. Chiba , A. , Ishida , H. , Nishizawa , N.K. , Makino , A. , & Mae , T. ( 2003 ) Exclusion of Ribulose - 1,5 - bisphosphate Carboxylase/oxygenase from chloroplasts by specifi c bodies in naturally senescing leaves of wheat . Plant and Cell Physiology 44 ( 9 ), 914 - 921 . Available at: http://pcp.oxfordjournals.org/ cgi/content/abstract/44/9/914 .
15. Choi , H.I. , Hong , J.H. , Ha , J.O. , Kang , J.Y. , & Kim , S.Y. ( 2000 ) ABFs, a family of ABA - responsive element binding factors . The Journal of Biological Chemistry 275 ( 3 ), 1723 - 1730 . Available at: http:// www.jbc.org/content/275/3/1723.abstract .
16. Choudhury , A. & Lahiri , A. ( 2011 ) Comparative analysis of abscisic acid - regulated transcriptomes in Arabidopsis . Plant Biology 13 ( 1 ), 28 - 35 .
17. Criado , M.V. , Caputo , C. , Roberts , I.N. , Castro , M.A. , & Barneix , A.J. ( 2009 ) Cytokinin - induced changes of nitrogen remobilization and chloroplast ultrastructure in wheat ( Triticum aestivum ) . Journal of Plant Physiology 166 ( 16 ), 1775 - 1785 . Available from: ISI:000271524900008.
18. Doelling , J.H. , Walker , J.M. , Friedman , E.M. , Thompson , A.R. , & Vierstra , R.D. ( 2002 ) The APG8/12 - activating enzyme APG7 is required for proper nutrient recycling and senescence in Arabidopsis thaliana . Journal of Biological Chemistry 277 ( 36 ), 33105 - 33114 . Available at: http://www.jbc.org/content/277/36/33105.abstract .
19. Dreccer , M.F. , Grashoff , C. , & Rabbinge , R. ( 1997 ) Source - sink ratio in barley ( Hordeum vulgare L.) during grain fi lling: effects on senescence and grain protein concentration . Field Crops Research 49 ( 2 - 3 ), 269 - 277 . Available at: http:// www.sciencedirect.com/science/article/B6T6M - 3W3FGNS - H/2/8356558215b1cbf9e2cbcd06 6213e81c .
20. Ehdaie , B. , Alloush , G.A. , Madore , M.A. , & Waines , J.G. ( 2006 ) Genotypic variation for stem reserves and mobilization in wheat: II. Postanthesis changes in internode water - soluble carbohydrates . Crop Science 46 ( 5 ), 2093 - 2103 . Available at: https:// www.crops.org/publications/cs/abstracts/46/5/ 2093 .
21. Feller , U. & Fischer , A. ( 1994 ) Nitrogen - metabolism in senescing leaves . Critical Reviews in Plant Sciences 13 ( 3 ), 241 - 273 . Available from: ISI:A1994PB67600003.
22. Feller , U. , Anders , I. , & Mae , T. ( 2008 ) Rubiscolytics: fate of Rubisco after its enzymatic function in a cell is terminated . Journal of Experimental Botany 59 ( 7 ), 1615 - 1624 . Available at: http:// jxb.oxfordjournals.org/cgi/content/ abstract/59/7/1615 .
23. Feller , U.K. , Soong , T.S. , & Hageman , R.H. ( 1977 ) Leaf proteolytic activities and senescence during grain development of fi eld - grown corn (Zea mays L) . Plant Physiology 59 ( 2 ), 290 - 294 . Available at: http://www.plantphysiol.org/cgi/content/ abstract/59/2/290 .
24. Foulkes , M.J. , Hawkesford , M.J. , Barraclough , P.B. , et al. ( 2009 ) Identifying traits to improve the nitrogen economy of wheat: recent advances and future prospects . Field Crops Research 114 ( 3 ), 329 - 342 . Available at: http://www.sciencedirect.com/science/ article/B6T6M - 4XDKCBJ - 2/2/08bfa420 2beb8251819f4efc69336077 .
25. Foyer , C.H. , Parry , M. , & Noctor , G. ( 2003) Markers and signals associated with nitrogen assimilation in higher plants. Journal of Experimental Botany 54 ( 382), 585 - 593. Available from: ISI:000180649800016.
26. Frink , C.R. , Waggoner , P.E. , & Ausubel , J.H. ( 1999 ) Nitrogen fertilizer: retrospect and prospect . Proceedings of the National Academy of Sciences of the United States of America 96 ( 4 ), 1175 - 1180 . Available at: http://www.pnas.org/content/96/4/ 1175.short .
27. Fujii , H. & Zhu , J.K. ( 2009 ) Arabidopsis mutant defi cient in 3 abscisic acid - activated protein kinases reveals critical roles in growth, reproduction, and stress . Proceedings of the National Academy of Sciences 106 ( 20 ), 8380 - 8385 . Available at: http://www.pnas.org/ content/106/20/8380.abstract .
28. Gan , S. & Amasino , R.M. ( 1995 ) Inhibition of leaf senescence by autoregulated production of cytokinin . Science 270 ( 5244 ), 1986 - 1988 . Available at: http://www.sciencemag.org/cgi/content/ abstract/270/5244/1986 .
29. Garg , O.P. & Kapoor , V. ( 1972 ) Retardation of leaf senescence by ascorbic acid . Journal of Experimental Botany 23 ( 3 ), 699 - 703 . Available at: http://jxb.oxfordjournals.org/cgi/content/ abstract/23/3/699 .
30. Garnett , T.P. & Graham , R.D. ( 2005 ) Distribution and remobilization of iron and copper in wheat . Annals of Botany 95 ( 5 ), 817 - 826 . Available at: http:// aob.oxfordjournals.org/cgi/content/ abstract/95/5/817 .
31. Gepstein , S. , Sabehi , G. , Carp , M.J. , et al. ( 2003 ) Large - scale identifi cation of leaf senescence - associated genes . Plant Journal 36 ( 5 ), 629 - 642 . Available from: ISI:000186532400005.
32. Gergoff , G. , Chaves , A. , & Bartoli , C.G. ( 2010 ) Ethylene regulates ascorbic acid content during dark - induced leaf senescence . Plant Science 178 ( 2 ), 207 - 212 . Available from: ISI:0002748777 00018.
33. Goldbach , H. , Goldbach , E. , & Michael , G. ( 1977 ) Transport of abscisic acid from leaves to grains in wheat and barley plants . Naturwissenschaften 64 ( 9 ), 488 - 489 . Available at: http:// dx.doi.org/10.1007/BF00446269 .
34. Gong , Y.H. , Zhang , J. , Gao , J.F. , Lu , J.Y. , & Wang , J.R. ( 2005 ) Slow export of photoassimilate from stay - green leaves during late grain - fi lling stage in hybrid winter wheat ( Triticum aestivum L.) . Journal of Agronomy and Crop Science 191 ( 4 ), 292 - 299 . Available from: ISI:000230728000007.
35. Gregersen , P.L. & Holm , P.B. ( 2007 ) Transcriptome analysis of senescence in the fl ag leaf of wheat ( Triticum aestivum L.) . Plant Biotechnology Journal 5 ( 1 ), 192 - 206 . Available from: ISI:000242963700018.
36. Gregersen , P.L. , Holm , P.B. , & Krupinska , K. ( 2008 ) Leaf senescence and nutrient remobilisation in barley and wheat . Plant Biology 10 , 37 - 49 . Available from: ISI:000258078400005.
37. Guo , Y. , Cai , Z. , & Gan , S. ( 2004 ) Transcriptome of Arabidopsis leaf senescence . Plant, Cell & Environment 27 ( 5 ), 521 - 549 . Available from: ISI:000221231600001.
38. Guo , Y.F. & Gan , S.S. ( 2006 ) AtNAP, a NAC family transcription factor, has an important role in leaf senescence . Plant Journal 46 ( 4 ), 601 - 612 . Available from: ISI:000237098000006.
39. Gut , H. & Matile , P. ( 1988 ) Apparent induction of key enzymes of the glyoxylic acid cycle in senescent barley leaves . Planta 176 ( 4 ), 548 - 550 . Available at: http://dx.doi.org/10.1007/BF00397663 .
40. Hafsi , M. , Mechmeche , W. , Bouamama , L. , Djekoune , A. , Zaharieva , M. , & Monneveux , P. ( 2000 ) Flag leaf senescence, as evaluated by numerical image analysis, and its relationship with yield under drought in durum wheat . Journal of Agronomy and Crop Science 185 ( 4 ), 275 - 280 . Available from: ISI:000165669300009.
41. Harding , S.A. , Guikema , J.A. , & Paulsen , G.M. ( 1990 ) Photosynthetic decline from high temperature stress during maturation of wheat: I. Interaction with senescence processes . Plant Physiology 92 ( 3 ), 648 - 653 . Available at: http://www.plantphysiol.org/cgi/ content/abstract/92/3/648 .
42. Hayashi , H. & Chino , M. ( 1990 ) Chemical composition of phloem sap from the uppermost internode of the rice plant . Plant and Cell Physiology 31 ( 2 ), 247 - 251 . Available at: http://pcp.oxfordjournals.org/cgi/ content/abstract/31/2/247 .
43. Heidlebaugh , N.M. , Trethewey , B.R. , Jukanti , A.K. , Parrott , D.L. , Martin , J.M. , & Fischer , A.M. ( 2008 ) Effects of a barley ( Hordeum vulgare ) chromosome 6 grain protein content locus on whole - plant nitrogen reallocation under two different fertilisation regimes . Functional Plant Biology 35 ( 7 ), 619 - 632 . Available from: ISI:000258592100009.
44. Helsel , D.B. & Frey , K.J. ( 1978 ) Grain yield variations in oats associated with differences in leaf area duration among oat lines . Crop Science 18 ( 5 ), 765 - 769 . Available at: https://www.crops.org/publications/cs/ abstracts/18/5/CS0180050765 .
45. Hill , J. , Robson, A.D. , & Loneragan, J.F. ( 1979) The effect of copper supply on the senescence and the retranslocation of nutrients of the oldest leaf of wheat . Annals of Botany 44 ( 3), 279 - 287. Available at: http:// aob.oxfordjournals.org/cgi/content/abstract/44/3/279.
46. Himelblau , E. & Amasino , R.M. ( 2001 ) Nutrients mobilized from leaves of Arabidopsis thaliana during leaf senescence . Journal of Plant Physiology 158 ( 10 ), 1317 - 1323 . Available at: http:// www.sciencedirect.com/science/article/B7GJ7 - 4DPXFGH - 5R/2/363dfa7c687 66d2d60ed83b76622430e .
47. Hirel , B. , Le Gouis , J. , Ney , B. , & Gallais , A. ( 2007 ) The challenge of improving nitrogen use effi ciency in crop plants: towards a more central role for genetic variability and quantitative genetics within integrated approaches . Journal of Experimental Botany 58 ( 9 ), 2369 - 2387 . Available from: ISI:000248846400010.
48. Hodges , M. ( 2002 ) Enzyme redundancy and the importance of 2 - oxoglutarate in plant ammonium assimilation . Journal of Experimental Botany 53 ( 370 ), 905 - 916 . Available from: ISI:000174853500014.
49. Hörtensteiner , S. ( 2009 ) Stay - green regulates chlorophyll and chlorophyll - binding protein degradation during senescence . Trends in Plant Science 14 ( 3 ), 155 - 162 . Available from: ISI:000264681000007.
50. Hörtensteiner , S. & Feller , U. ( 2002 ) Nitrogen metabolism and remobilization during senescence . Journal of Experimental Botany 53 ( 370 ), 927 - 937 . Available from: ISI:000174853500016.
51. Howarth , J.R. , Parmar , S. , Jones , J. , et al. ( 2008 ) Co - ordinated expression of amino acid metabolism in response to N and S defi ciency during wheat grain fi lling . Journal of Experimental Botany 59 ( 13 ), 3675 - 3689 . Available from: ISI:000259973800016.
52. Humbeck , K. , Quast , S. , & Krupinska , K. ( 1996 ) Functional and molecular changes in the photosynthetic apparatus during senescence of fl ag leaves from fi eld - grown barley plants . Plant, Cell & Environment 19 ( 3 ), 337 - 344 . Available at: http://www.blackwell- synergy.com/doi/abs/ 10.1111/j.1365 - 3040.1996.tb00256.x .
53. Huynh , L.N. , VanToai , T. , Streeter , J. , & Banowetz , G. ( 2005 ) Regulation of fl ooding tolerance of SAG12:ipt Arabidopsis plants by cytokinin . Journal of Experimental Botany 56 ( 415 ), 1397 - 1407 . Available at: http://jxb.oxfordjournals.org/cgi/ content/abstract/56/415/1397 .
54. Ishida , H. , Yoshimoto , K. , Izumi , M. , et al. ( 2008 ) Mobilization of rubisco and stroma - localized fl uorescent proteins of chloroplasts to the vacuole by an ATG gene - dependent autophagic process . Plant Physiology 148 ( 1 ), 142 - 155 . Available at: http:// www.plantphysiol.org/cgi/content/abstract/148/ 1/142 .
55. Jahn , T.P. , Møller , A.L.B. , Zeuthen , T. , et al. ( 2004 ) Aquaporin homologues in plants and mammals transport ammonia . FEBS Letters 574 ( 1 - 3 ), 31 - 36 . Available at: http://www.sciencedirect.com/science/ article/B6T36 - 4D2XB26 - 1/2/7cd9297f5f4b99b48 2a2722d6a2075a5 .
56. Joppa , L.R. , Du, C. , Hart, G.E. , & Hareland, G.A. ( 1997) Mapping gene(s) for grain protein in tetraploid wheat ( Triticum turgidum L.) using a population of recombinant inbred chromosome lines . Crop Science 37 ( 5), 1586 - 1589. Available at: https://www.crops.org/ publications/cs/abstracts/37/5/CS0370051586.
57. Jukanti , A. & Fischer , A. ( 2008 ) A high - grain protein content locus on barley ( Hordeum vulgare ) chromosome 6 is associated with increased fl ag leaf proteolysis and nitrogen remobilization . Physiologia Plantarum 132 ( 4 ), 426 - 439 . Available at: http:// onlinelibrary.wiley.com/doi/10.1111/j.1399 - 3054. 2007.01044.x/abstract .
58. Jukanti , A.K. , Heidlebaugh , N.M. , Parrott , D.L. , Fischer , I.A. , McInnerney , K. , & Fischer , A.M. ( 2008 ) Comparative transcriptome profi ling of near - isogenic barley ( Hordeum vulgare ) lines differing in the allelic state of a major grain protein content locus identifi es genes with possible roles in leaf senescence and nitrogen reallocation . New Phytologist 177 ( 2 ), 333 - 349 . Available from: ISI:000251766300008.
59. Kato , Y. & Sakamoto , W. ( 2009 ) Protein quality control in chloroplasts: a current model of D1 protein degradation in the photosystem II repair cycle . Journal of Biochemistry 146 ( 4 ), 463 - 469 . Available from: ISI:000270709400003.
60. Kato , Y. , Yamamoto , Y. , Murakami , S. , & Sato , F. ( 2005 ) Post - translational regulation of CND41 protease activity in senescent tobacco leaves . Planta 222 ( 4 ), 643 - 651 . Available at: http://dx.doi. org/10.1007/s00425 - 005 - 0011 - 4 .
61. Kichey , T. , Heumez , E. , Pocholle , D. , et al. ( 2006 ) Combined agronomic and physiological aspects of nitrogen management in wheat highlight a central role for glutamine synthetase . New Phytologist 169 ( 2 ), 265 - 278 . Available from: ISI:00023448 2900006.
62. Kichey , T. , Hirel , B. , Heumez , E. , Dubois , F. , & Le Gouis , J. ( 2007 ) In winter wheat ( Triticum aestivum L.), post - anthesis nitrogen uptake and remobilisation to the grain correlates with agronomic traits and nitrogen physiological markers . Field Crops Research 102 ( 1 ), 22 - 32 . Available from: ISI:000246533800003.
63. Kim , J.H. , Woo , H.R. , Kim , J. , et al. ( 2009 ) Trifurcate feed - forward regulation of age - dependent cell death involving miR164 in Arabidopsis . Science 323 ( 5917), 1053 - 1057. Available from: ISI:000263478 400040.
64. Kwon , S. & Park , O. ( 2008 ) Autophagy in plants . Journal of Plant Biology 51 ( 5 ), 313 - 320 . Available at: http://dx.doi.org/10.1007/BF03036132 .
65. Lam , E. , Kato , N. , & Lawton , M. ( 2001 ) Programmed cell death, mitochondria and the plant hypersensitive response . Nature 411 ( 6839 ), 848 - 853 . Available from: ISI:000169246400062.
66. Lim , P.O. , Kim , H.J. , & Gil Nam , H. ( 2007 ) Leaf senescence . Annual Review of Plant Biology 58 ( 1), 115 - 136. from: http://dx.doi.org/10.1146/annurev.arplant. 57.032905.105316 .
67. Lin , W. & Wittenbach , V.A. ( 1981 ) Subcellular localization of proteases in wheat and corn mesophyll protoplasts . P lant Physiology 67 ( 5 ), 969 - 972 . Available at: http://www.plantphysiol.org/cgi/ content/abstract/67/5/969 .
68. Liu , F.L. , Jensen , C.R. , & Andersen , M.N. ( 2005 ) A review of drought adaptation in crop plants: changes in vegetative and reproductive physiology induced by ABA - based chemical signals . Australian Journal of Agricultural Research 56 ( 11 ), 1245 - 1252 . Available from: ISI:000233570500015.
69. Lupton , F.G.H. , Oliver , R.H. , & Ruckenbauer , P. ( 1974 ) An analysis of the factors determining yields in crosses between semi - dwarf and taller wheat varieties . The Journal of Agricultural Science 82 ( 03 ), 483 - 496 . Available at: http:// journals.cambridge.org/action/displayAbstract ?fromPage=online & aid=4705320 & fulltextType=R A & fi leId=S0021859600051388 .
70. Ma , Y. , Szostkiewicz , I. , Korte , A. , et al. ( 2009 ) Regulators of PP2C phosphatase activity function as abscisic acid sensors . Science 324 ( 5930 ), 1064 - 1068 . Available at: http://www.sciencemag.org/cgi/ content/abstract/324/5930/1064 .
71. Martin , A. , Lee , J. , Kichey , T. , et al. ( 2006 ) Two cytosolic glutamine synthetase isoforms of maize are specifi cally involved in the control of grain production . Plant Cell 18 ( 11 ), 3252 - 3274 . Available from: ISI:000243093700030.
72. Martinez , D.E. , Bartoli , C.G. , Grbic , V. , & Guiamet , J.J. ( 2007 ) Vacuolar cysteine proteases of wheat ( Triticum aestivum L.) are common to leaf senescence induced by different factors . Journal of Experimental Botany 58 ( 5 ), 1099 - 1107 . Available at: http://jxb.oxfordjournals.org/cgi/content/abstract/ 58/5/1099 .
73. Martinez , D.E. , Costa , M.L. , & Guiamet , J.J. ( 2008 ) Senescence - associated degradation of chloroplast proteins inside and outside the organelle . Plant Biology 10 , 15 - 22 . Available from: ISI:000258078400003.
74. Matile , P. , Hortensteiner , S. , Thomas , H. , & Krautler , B. ( 1996 ) Chlorophyll breakdown in senescent leaves . Plant Physiology 112 ( 4 ), 1403 - 1409 . Available at: http://www.plantphysiol.org/content/ 112/4/1403.full.pdf+html?sid=9b2b77c4 - 8c56 - 46bd - 8633 - 001686f3c20e .
75. Mauk , C.S. & Nooden , L.D. ( 1992 ) Regulation of mineral redistribution in pod - bearing soybean explants . Journal of Experimental Botany 43 ( 256 ), 1429 - 1440 . Available from: ISI:A1992JZ 53100006.
76. Miao , Y. , Laun , T. , Zimmermann , P. , & Zentgraf , U. ( 2004 ) Targets of the WRKY53 transcription factor and its role during leaf senescence in Arabidopsis . Plant Molecular Biology 55 ( 6 ), 853 - 867 . Available from: ISI:000226091100007.
77. Monaghan , J.M. , Snape , J.W. , Chojecki , A.J.S. , & Kettlewell , P.S. ( 2001 ) The use of grain protein deviation for identifying wheat cultivars with high grain protein concentration and yield . Euphytica 122 , 309 - 317 .
78. Moon , J. , Parry , G. , & Estelle , M. ( 2004 ) The ubiquitin - proteasome pathway and plant development . Plant Cell 16 ( 12 ), 3181 - 3195 . Available from: ISI:000225780700003.
79. Nakano , R. , Ishida , H. , Makino , A. , & Mae , T. ( 2006 ) In vivo fragmentation of the large subunit of ribulose - 1,5 - bisphosphate carboxylase by reactive oxygen species in an intact leaf of cucumber under chilling - light conditions . Plant and Cell Physiology 47 ( 2), 270 - 276. Available at: http://pcp.oxfordjournals. org/cgi/content/abstract/47/2/270 .
80. Nakashima , K. , Kiyosue , T. , YamaguchiShinozaki , K. , & Shinozaki , K. ( 1997 ) A nuclear gene, erd1 encoding a chloroplast - targeted Clp protease regulatory subunit homolog is not only induced by water stress but also developmentally up - regulated during senescence in Arabidopsis thaliana . Plant Journal 12 ( 4 ), 851 - 861 . Available from: ISI:A1997YF 45400011.
81. Nambara, E. & Marion- Poll, A. ( 2003) ABA action and interactions in seeds . Trends in Plant Science 8 ( 5 ), 213 - 217 . Available at: http:// www.sciencedirect.com/science/article/B6TD1 - 48CFPX8 - 2/2/01a1510ecd757727 aff3a118c3a7976a .
82. Noodén , L.D. ( 1988 ) The phenomena of senescence and aging . In: Senescence and Aging in Plants (eds. L.D. Noodén & A.C. Leopold ), pp. 1 - 50 , Academic Press , San Diego, CA .
83. Okatan , Y. , Kahanak , G.M. , & Nooden , L.D. ( 1981 ) Characterization and kinetics of soybean maturation and monocarpic senescence . Physiologia Plantarum 52 ( 2 ), 330 - 338 . Available from: ISI:A1981ME 32400031.
84. Olmos , S. , Distelfeld , A. , Chicaiza , O. , et al. ( 2003 ) Precise mapping of a locus affecting grain protein content in durum wheat . Theoretical and Applied Genetics 107 ( 7 ), 1243 - 1251 . Available from: ISI:000186401200012.
85. Otegui , M.S. , Noh , Y.S. , Martinez , D.E. , et al. ( 2005 ) Senescence - associated vacuoles with intense proteolytic activity develop in leaves of Arabidopsis and soybean . Plant Journal 41 ( 6 ), 831 - 844 . Available from: ISI:000227350700004.
86. Palta , J.A. & Fillery , I.R.P. ( 1995 ) N - Application enhances remobilization and reduces losses of preanthesis - N in wheat grown on a duplex soil . Australian Journal of Agricultural Research 46 ( 3 ), 519 - 531 . Available from: ISI:A1995QV27500005.
87. Park , S.Y. , Yu , J.W. , Park , J.S. , et al. ( 2007 ) The senescence - induced staygreen protein regulates chlorophyll degradation . The Plant Cell 19 ( 5 ), 1649 - 1664 . Available at: http://www.plantcell.org/ content/19/5/1649.abstract .
88. Park , S.Y. , Fung , P. , Nishimura , N. , et al. ( 2009 ) Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins . Science 324 ( 5930 ), 1068 - 1071 . Available at: http://www.sciencemag.org/cgi/content/abstract/ 324/5930/1068 .
89. Pearson , J.N. & Rengel , Z. ( 1994 ) Distribution and remobilization of Zn and Mn during grain development in wheat . Journal of Experimental Botany 45 ( 12 ), 1829 - 1835 . Available at: http:// jxb.oxfordjournals.org/cgi/content/ abstract/45/12/1829 .
90. Pennisi , E. ( 2009 ) Stressed out over a stress hormone . Science 324 ( 5930 ), 1012 - 1013 . Available at: http:// www.sciencemag.org/content/324/5930/1012. summary?sid=72c7d455 - b883 - 41e7 - 960e - 2d411961acd7 .
91. Pracharoenwattana , I. & Smith , S.M. ( 2008 ) When is a peroxisome not a peroxisome? Trends in Plant Science 13 ( 10 ), 522 - 525 . Available from: ISI:000260645400004.
92. Raab , S. , Drechsel , G. , Zarepour , M. , et al. ( 2009 ) Identifi cation of a novel E3 ubiquitin ligase that is required for suppression of premature senescence in Arabidopsis . P lant Journal 5 9 ( 1 ), 39 - 51 . Available from: ISI:000267428400004.
93. Rawson , H.M. , Hindmarsh , J.H. , Fischer , R.A. , & Stockman , Y.M. ( 1983 ) Changes in leaf photosynthesis with plant ontogeny and relationships with yield per ear in wheat cultivars and 120 - progeny . Australian Journal of Plant Physiology 10 ( 6 ), 503 - 514 . Available from: ISI:A1983SC29800003.
94. Rivero , R.M. , Shulaev , V. , & Blumwald , E. ( 2009 ) Cytokinin - dependent photorespiration and the protection of photosynthesis during water defi cit . Plant Physiology 150 ( 3 ), 1530 - 1540 . Available at: http://www.plantphysiol.org/cgi/content/ abstract/150/3/1530 .
95. Robatzek, S. & Somssich, I.E. ( 2001) A new member of the Arabidopsis WRKY transcription factor family, AtWRKY6, is associated with both senescence - and defence- related processes. Plant Journal 28 ( 2), 123 - 133. Available from: ISI:000172099500001.
96. Robatzek , S. & Somssich , I.E. ( 2002 ) Targets of AtWRKY6 regulation during plant senescence and pathogen defense . Genes & Development 16 ( 9 ), 1139 - 1149 . Available from: ISI:000175556800011.
97. Sakakibara , H. , Takei , K. , & Hirose , N. ( 2006 ) Interactions between nitrogen and cytokinin in the regulation of metabolism and development . Trends in Plant Science 11 ( 9 ), 440 - 448 . Available at: http://www.sciencedirect.com/science/article/ B6TD1 - 4KKNNHJ - 1/2/665741e23d139642aae073 9c055a3091 .
98. Santner , A. & Estelle , M. ( 2010 ) The ubiquitin - proteasome system regulates plant hormone signaling . Plant Journal 61 ( 6 ), 1029 - 1040 . Available from: ISI:000275394300012.
99. Schelbert , S. , Aubry , S. , Burla , B. , et al. ( 2009 ) Pheophytin pheophorbide hydrolase (pheophytinase) is involved in chlorophyll breakdown during leaf senescence in Arabidopsis . The Plant Cell 21 ( 3 ), 767 - 785 . Available at: http://www.plantcell. org/cgi/content/abstract/21/3/767 .
100. Schippers, J.H.M. , Jing, H.- C. , Hille, J. , & Dijkwel, P.P. ( 2007) Developmental and hormonal control of leaf senescence . In: Senescence Processes in Plants (ed. S. Gan ), pp. 145 - 170, Blackwell Publ., Oxford
101. Schjoerring , J.K. , Kyllingsbaek , A. , Mortensen , J.V. , & Byskovnielsen , S. ( 1993 ) Field investigations of ammonia exchange between barley plants and the atmosphere. 1. Concentration profi les and fl ux densities of ammonia . Plant, Cell & Environment 16 ( 2 ), 161 - 167 . Available from: ISI:A1993KV93700005.
102. Seki , M. , Ishida , J. , Narusaka , M. , et al. ( 2002 ) Monitoring the expression pattern of around 7000 Arabidopsis genes under ABA treatments using a full - length cDNA microarray . Functional & Integrative Genomics 2 ( 6 ), 282 - 291 . Available at: http://dx.doi.org/10.1007/s10142 - 002 - 0070 - 6 .
103. Shen , H. , Yin , Y. , Chen , F. , Xu , Y. , & Dixon , R. ( 2009 ) A bioinformatic analysis of NAC genes for plant cell wall development in relation to lignocellulosic bioenergy production . BioEnergy Research 2 ( 4 ), 217 - 232 . Available from: CABI:20093331713.
104. Simmonds , N.W. ( 1995 ) The relation between yield and protein in cereal grain . Journal of the Science of Food and Agriculture 67 ( 3 ), 309 - 315 . Available from: ISI:A1995QM87300005.
105. Simpson , R.J. & Dalling , M.J. ( 1981 ) Nitrogen redistribution during grain - growth in wheat ( Triticum aestivum L). 3. Enzymology and transport of amino - acids from senescing fl ag leaves . Planta 151 ( 5 ), 447 - 456 . Available from: ISI:A1981LP 99900006.
106. Simpson , R.J. , Lambers , H. , & Dalling , M.J. ( 1983 ) Nitrogen redistribution during grain - growth in wheat ( Triticum aestivum L). 4. Development of a quantitative model of the translocation of nitrogen to the grain . Plant Physiology 71 ( 1 ), 7 - 14 . Available from: ISI:A1983PY81500002.
107. Snape , J.W. , Butterworth , K. , Whitechurch , E. , & Worland , A.J. ( 2001 ) Waiting for fi ne times: genetics of fl owering time in wheat . Euphytica 119 ( 1 ), 185 - 190 . Available at: http://dx.doi.org/10.1023/A: 1017594422176 .
108. Spano , G. , Di Fonzo , N. , Perrotta , C. , et al. ( 2003 ) Physiological characterization of " stay green " mutants in durum wheat . Journal of Experimental Botany 54 ( 386 ), 1415 - 1420 . Available from: ISI:000182630300013.
109. Tabuchi , M. , Sugiyama , K. , Ishiyama , K. , et al. ( 2005 ) Severe reduction in growth rate and grain fi lling of rice mutants lacking OsGS1;1, a cytosolic glutamine synthetase1;1 . Plant Journal 42 ( 5 ), 641 - 651 . Available from: ISI:000229180400003.
110. Tabuchi, M. , Abiko, T. , & Yamaya, T. ( 2007) Assimilation of ammonium ions and reutilization of nitrogen in rice ( Oryza sativa L). Journal of Experimental Botany 58 ( 9), 2319 - 2327. Available at: http://jxb. oxfordjournals.org/cgi/content/abstract/58/9/2319.
111. Tauris , B. , Borg , S. , Gregersen , P.L. , & Holm , P.B. ( 2009 ) A roadmap for zinc traffi cking in the developing barley grain based on laser capture microdissection and gene expression profi ling . Journal of Experimental Botany 60 ( 4 ), 1333 - 1347 . Available at: http://jxb.oxfordjournals.org/cgi/content/abstract/ 60/4/1333 .
112. Thomas , H. ( 1994 ) Aging in the plant and animal kingdoms? The role of cell death . Reviews in Clinical Gerontology 4 ( 01 ), 5 - 20 . Available at: http:// journals.cambridge.org/action/displayAbstract ?fromPage=online & aid=2184512 & fulltextType=R V & fi leId=S0959259800002227 .
113. Thomas , H. & Howarth , C.J. ( 2000 ) Five ways to stay green . Journal of Experimental Botany 51 , 329 - 337 . Available from: ISI:000085386600003.
114. Thomas , H. & Stoddart , J.L. ( 1980 ) Leaf senescence . Annual Review of Plant Physiology and Plant Molecular Biology 31 , 83 - 111 . Available from: ISI:A1980JU72700004.
115. Thompson , J.E. , Froese , C.D. , Madey , E. , Smith , M.D. , & Yuwen , H. ( 1998 ) Lipid metabolism during plant senescence . Progress in Lipid Research 37 ( 2 - 3 ), 119 - 141 . Available at: http://www.sciencedirect. com/science/article/B6TBP - 3TXDH37 - 2/2/9bad60fe8b61c093334c0d6465677888 .
116. Tollenaar , M. & Wu , J. ( 1999 ) Yield improvement in temperate maize is attributable to greater stress tolerance . Crop Science 39 ( 6 ), 1597 - 1604 . Available at: https://www.crops.org/publications/cs/abstracts/ 39/6/1597 .
117. Uauy , C. , Brevis , J.C. , & Dubcovsky , J. ( 2006a ) The high grain protein content gene Gpc - B1 accelerates senescence and has pleiotropic effects on protein content in wheat . Journal of Experimental Botany 57 ( 11 ), 2785 - 2794 . Available from: ISI:000239 901500030.
118. Uauy , C. , Distelfeld , A. , Fahima , T. , Blechl , A. , & Dubcovsky , J. ( 2006b ) A NAC gene regulating senescence improves grain protein, zinc, and iron content in wheat . Science 314 ( 5803 ), 1298 - 1301 . Available from: ISI:000242215800043.
119. Van der Graaff , E. , Schwacke , R. , Schneider , A. , Desimone , M. , Flugge , U.I. , & Kunze , R. ( 2006 ) Transcription analysis of Arabidopsis membrane transporters and hormone pathways during developmental and induced leaf senescence . Plant Physiology 141 ( 2 ), 776 - 792 . Available from: ISI:000238168800051.
120. van Doorn , G. & Woltering , E. ( 2004 ) Senescence and programmed cell death: substance or semantics? Journal of Experimental Botany 55 , 2147 - 2153 . Available at: http://jxb.oxfordjournals.org/content/ 55/406/2147.abstract .
121. van Oosterom , E.J. , Chapman , S.C. , Borrell , A.K. , Broad , I.J. , & Hammer , G.L. ( 2010 ) Functional dynamics of the nitrogen balance of sorghum. II. Grain fi lling period . Field Crops Research 115 ( 1 ), 29 - 38 . Available at: http://www.sciencedirect.com/ science/article/B6T6M - 4XK9J3K - 2/2/ b51461b7947f768a531219e9e9cd0fef .
122. Verdier , J. & Thompson , R.D. ( 2008 ) Transcriptional regulation of storage protein synthesis during dicotyledon seed fi lling . Plant and Cell Physiology 49 ( 9), 1263 - 1271. Available at: http://pcp.oxfordjournals. org/cgi/content/abstract/49/9/1263 .
123. Vierstra , R.D. ( 2009 ) The ubiquitin - 26S proteasome system at the nexus of plant biology . Nature Reviews. Molecular Cell Biology 10 ( 6 ), 385 - 397 . Available at: http://dx.doi.org/10.1038/nrm2688 .
124. Wada , S. , Ishida , H. , Izumi , M. , et al. ( 2009 ) Autophagy plays a role in chloroplast degradation during senescence in individually darkened leaves . Plant Physiology 149 ( 2 ), 885 - 893 . Available at: http://www.plantphysiol.org/cgi/content/abstract/ 149/2/885 .
125. Waters , S.P. , Peoples , M.B. , Simpson , R.J. , & Dalling , M.J. ( 1980 ) Nitrogen redistribution during grain - growth in wheat (Triticum - Aestivum L). 1. Peptide - hydrolase activity and protein breakdown in the fl ag leaf, glumes and stem . Planta 148 ( 5 ), 422 - 428 . Available from: ISI:A1980JW58600002.
126. Waters , B.M. , Uauy , C. , Dubcovsky , J. , & Grusak , M.A. ( 2009 ) Wheat (Triticum aestivum) NAM proteins regulate the translocation of iron, zinc, and nitrogen compounds from vegetative tissues to grain . Journal of Experimental Botany 60 ( 15 ), 4263 - 4274 . Available at: http://jxb.oxfordjournals.org/cgi/ content/abstract/60/15/4263 .
127. Wilkinson , S. & Davies , W.J. ( 2002 ) ABA - based chemical signalling: the co - ordination of responses to stress in plants . Plant, Cell & Environment 25 ( 2 ), 195 - 210 . Available from: ISI:000174061500007.
128. Wittenbach , V.A. ( 1979 ) Ribulose bisphosphate carboxylase and proteolytic activity in wheat leaves from anthesis through senescence . Plant Physiology 64 ( 5 ), 884 - 887 . Available from: ISI:A1979HV 88000043.
129. Wolfe , D.W. , Henderson , D.W. , Hsiao , T.C. , & Alvino , A. ( 1988 ) Interactive water and nitrogen effects on senescence of maize. I. Leaf area duration, nitrogen distribution, and yield . Agronomy Journal 80 ( 6 ), 859 - 864 . Available at: https://www.agronomy.org/ publications/aj/abstracts/80/6/AJ0800060859 .
130. Woo , H.R. , Chung , K.M. , Park , J.H. , et al. ( 2001 ) ORE9, an F - Box protein that regulates leaf senescence in Arabidopsis . The Plant Cell 13 ( 8 ), 1779 - 1790 . Available at: http://www.plantcell.org/cgi/ content/abstract/13/8/1779 .
131. Xie , Z. , Jiang , D. , Dai , T. , Jing , Q. , & Cao , W. ( 2004 ) Effects of exogenous ABA and cytokinin on leaf photosynthesis and grain protein accumulation in wheat ears cultured in vitro . Plant Growth Regulation 44 ( 1 ), 25 - 32 . Available at: http:// dx.doi.org/10.1007/s10725 - 004 - 1880 - 4 .
132. Yang , Z.L. & Ohlrogge , J.B. ( 2009 ) Turnover of fatty acids during natural senescence of Arabidopsis, brachypodium, and switchgrass and in Arabidopsis beta - oxidation mutants . Plant Physiology 150 ( 4 ), 1981 - 1989 . Available from: ISI:000268692200032.
133. Yang , J.C. & Zhang , J.H. ( 2006 ) Grain fi lling of cereals under soil drying . New Phytologist 169 ( 2 ), 223 - 236 . Available from: ISI:000234482900002.
134. Yang , J. , Zhang , J. , Wang , Z. , Zhu , Q. , & Liu , L. ( 2001 ) Water defi cit - induced senescence and its relationship to the remobilization of pre - stored carbon in wheat during grain fi lling . Agronomy Journal 93 ( 1 ), 196 - 206 . Available at: https://www.agronomy.org/ publications/aj/abstracts/93/1/196 .
135. Yang , J.C. , Zhang , J.H. , Wang , Z.Q. , Zhu , Q.S. , & Liu , L.J. ( 2003 ) Involvement of abscisic acid and cytokinins in the senescence and remobilization of carbon reserves in wheat subjected to water stress during grain fi lling . Plant, Cell & Environment 26 ( 10 ), 1621 - 1631 . Available from: ISI:00018553 6000003.
136. Yoo , S.C. , Cho , S.H. , Zhang , H. , et al. ( 2007 ) Quantitative trait loci associated with functional stay - green SNU - SG1 in rice . Molecules and Cells 24 ( 1 ), 83 - 94 . Available at: http://www.scopus.com/ inward/record.url?eid=2 - s2.0 - 35048855476 & par tnerID=40 & md5=3638e8d644722a1e2418628237 8a7c6f .
137. Yoshida , S. , Ito , M. , Callis , J. , Nishida , I. , & Watanabe , A. ( 2002 ) A delayed leaf senescence mutant is defective in arginyl - tRNA: protein arginyltransferase, a component of the N - end rule pathway in Arabidopsis . P lant Journal 32 ( 1 ), 129 - 137 . Available from: ISI:000178374100011.
138. Zacarias , L. & Reid , M.S. ( 1990 ) Role of growth - regulators in the senescence of Arabidopsis thaliana leaves . Physiologia Plantarum 80 ( 4 ), 549 - 554 . Available from: ISI:A1990ER94300009.
139. Zapata , J.M. , Guera , A. , Esteban - Carrasco , A. , Martin , M. , & Sabater , B. ( 2005 ) Chloroplasts regulate leaf senescence: delayed senescence in transgenic ndhF - defective tobacco . Cell Death and Differentiation 12 ( 10 ), 1277 - 1284 . Available at: http:// dx.doi.org/10.1038/sj.cdd.4401657 .
140. Zapata , J.M. , Gasulla , F. , Esteban - Carrasco , A. , Barreno , E. , & Guera , A. ( 2007 ) Inactivation of a plastid evolutionary conserved gene affects PSII electron transport, life span and fi tness of tobacco plants . New Phytologist 174 ( 2 ), 357 - 366 . Available from: ISI:000245413900018.
141. Zelisko , A. , Garcia - Lorenzo , M. , Jackowski , G. , Jansson, S. , & Funk, C. ( 2005) 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 of the United States of America 102 ( 38 ), 13699 - 13704 . Available from: ISI:000232115100062.
142. Zentgraf , U. , Laun , T. , & Miao , Y. ( 2002 ) The complex regulation of WRKY53 during leaf senescence of Arabidopsis thaliana . European Journal of Cell Biology 89 ( 2 - 3 ), 133 - 137 . Available at: http://www.sciencedirect.com/science/article/ B7GJ2 - 4XWMGG1 - 1/2/ d4cd48b1af888fc622e3d56a87bd7774 .
143. Zentgraf , U. , Laun , T. , & Miao , Y. ( 2010 ) The complex regulation of WRKY53 during leaf senescence of Arabidopsis thaliana . European Journal of Cell Biology 89 ( 2 - 3 ), 133 - 137 . Available from: ISI:000275308300003.
144. Zhong , G.V. & Burns , J.K. ( 2003 ) Profi ling ethylene - regulated gene expression in Arabidopsis thaliana by microarray analysis . Plant Molecular Biology 53 ( 1 ), 117 - 131 . Available from: ISI:000187 356900010.