Conclusions Directory UMM :Data Elmu:jurnal:E:Environmental and Experimental Botany:Vol44.Issue2.Oct2000:

5. Conclusions

Our findings explain the acceleration of senes- cence in annual monocarpic C 3 plants under CO 2 enrichment as a process driven by the differing effects of CO 2 on nutrients, in particular nitrogen, in vegetative and generative tissues. We postulate that CO 2 enhances flag leaf senescence in barley crops and probably leaf senescence in mono- carpic C 3 species in general by a sequence of several processes induced by CO 2 : 1 CO 2 reduces the nitrogen demand of green tissues by alter- ations of the photosynthetic apparatus, i.e. lower RubisCO contents, lower contents of enzymes of the PCO-cycle. Thus, nitrogen uptake does not keep pace with carbon acquisition during vegeta- tive growth. 2 CO 2 increases the number of diaspores. Thus, the sink size for nutrients during seed ripening is enlarged. 3 During grain filling or seed ripening in general there is a higher demand for nutrient redistribution to the seeds since CO 2 enrichment does not reduce the nitro- gen demand of the diaspores. Thus, a nitrogen deficiency is induced during the switch from vege- tative to generative growth. 4 This deficiency acts as a trigger to induce leaf senescence in order to release nutrients from vegetative tissues accord- ing to the nutrient salvage function of senescence, although the demand of the seeds can not be accomplished due to restricted nutrient pools. References Anonymous, 1995. Climate Change 1995. The Science of Climate Change. Summary for Policymakers. World Mete- orological OrganizationUnited Nations Environment Pro- gramme, pp. 1 – 56. Biswal, B., Biswal, U.C., 1999. Leaf senescence: physiology and molecular biology. Curr. Sci. 77, 775 – 782. Bleecker, A.B., 1998. The evolutionary basis of leaf senes- cence: method to the madness? Curr. Opin. Plant Biol. 1, 73 – 78. Chrost, B., Falk, J., Kernebeck, B., Mo¨lleken, H., Krupinska, K., 1999. Tocopherol biosynthesis in senescing chloroplasts — a mechanism to protect envelope membranes against oxidative stress and a prerequisite for lipid remobilization? In: Argyroudi-Akoyunoglou, J.H., Senger, H. Eds., The Chloroplast: from Molecular Biology to Biotechnology. Kluwer Academic Publishers, Dordrecht, pp. 171 – 176. Conroy, J.P., 1992. Influence of elevated atmospheric CO 2 concentrations on plant nutrition. Aust. J. Bot. 40, 445 – 456. Conroy, J.P., Hocking, P.J., 1993. Nitrogen nutrition of C 3 plants at elevated atmospheric CO 2 concentrations. Phys- iol. Plant. 89, 570 – 576. Cotrufo, M.F., Ineson, P., Scott, A., 1998. Elevated CO 2 reduces the nitrogen concentration of plant tissues. Global Change Biol. 4, 43 – 54. Fangmeier, A., Ja¨ger, H.-J., 1998. CO 2 enrichment, ozone, nitrogen fertilizer and wheat: physiological background of growth and yield responses. In: De Kok, L.J., Stulen, I. Eds., Responses of Plant Metabolism to Air Pollution and Global Change. Backhuys Publishers, Leiden, pp. 299 – 304. Fangmeier, A., Stein, W., Ja¨ger, H.-J., 1992. Advantages of an open-top chamber plant exposure system to assess the impact of atmospheric trace gases on vegetation. Angew. Bot. 66, 97 – 105. Fangmeier, A., Gru¨ters, U., Hertstein, U., Sandhage-Hof- mann, A., Vermehren, B., Ja¨ger, H.-J., 1996. Effects of elevated CO 2 , nitrogen supply and tropospheric ozone on spring wheat. I. Growth and yield. Environ. Pollut. 91, 381 – 390. Fangmeier, A., Gru¨ters, U., Ho¨gy, P., Vermehren, B., Ja¨ger, H.-J., 1997. Effects of elevated CO 2 , nitrogen supply and tropospheric ozone on spring wheat — II. Nutrients N, P, K, S, Ca, Mg, Fe, Mn, Zn. Environ. Pollut. 96, 43 – 59. Fangmeier, A., De Temmerman, L., Mortensen, L., Kemp, K., Burke, J.I., Mitchell, R.A.C., Van Oijen, M., Weigel, H.-J., 1999. Effects on nutrients and on grain quality in spring wheat crops grown under elevated CO 2 concentrations and stress conditions in the European, multiple-site experiment ‘ESPACE-wheat’. Eur. J. Agron. 10, 215 – 229. Gan, S., Amasino, R.M., 1997. Making sense of senescence — molecular genetic regulation and manipulation of leaf senescence. Plant Physiol. 113, 313 – 319. Garcia, R.L., Long, S.P., Wall, G.W., Osborne, C.P., Kimball, B.A., Nie, G.Y., Pinter, P.J., Lamorte, R.L., Wechsung, F., 1998. Photosynthesis and conductance of spring-wheat leaves: field response to continuous free-air atmospheric CO 2 enrichment. Plant Cell Environ. 21, 659 – 669. Harley, P.C., Sharkey, T.D., 1991. An improved model of C 3 photosynthesis at high CO 2 — reversed O 2 sensitivity explained by a lack of glycerate reentry into the chloro- plast. Photosynth. Res. 27, 169 – 178. Humbeck, K., Quast, S., Krupinska, K., 1996. Functional and molecular changes in the photosynthetic apparatus during senescence of flag leaves from field-grown barley plants. Plant Cell Environ. 19, 337 – 344. Jacob, J., Greitner, C., Drake, B.G., 1995. Acclimation of photosynthesis in relation to Rubisco and nonstructural carbohydrate contents and in situ carboxylase activity in Scirpus olneyi grown at elevated CO 2 in the field. Plant Cell Environ. 18, 875 – 884. Kelly, M.O., Davies, P.J., 1988. The control of whole plant senescence. CRC Crit. Rev. Plant Sci. 7, 139 – 173. Kleber-Janke, T., Krupinska, K., 1997. Isolation of cDNA clones for genes showing enhanced expression in barley leaves during dark-induced senescence as well as during senescence under field conditions. Planta 203, 332 – 340. Ko¨rner, C., 1995. Towards a better experimental basis for upscaling plant responses to elevated CO 2 and climate warming. Plant Cell Environ. 18, 1101 – 1110. Lowry, O.H., Rosebrough, N.J., Farr, A.L., Randall, R.J., 1951. Protein measurement with the folin phenol reagent. J. Biol. Chem. 193, 265 – 275. Manderscheid, R., Bender, J., Ja¨ger, H.-J., Weigel, H.-J., 1995. Effects of season long CO 2 enrichment on cereals. II. Nutrient concentrations and grain quality. Agric. Ecosyst. Environ. 54, 175 – 185. McKee, I.F., Woodward, F.I., 1994. CO 2 enrichment responses of wheat: interactions with temperature, nitrate and phos- phate. New Phytol. 127, 447 – 453. Miller, A., Tsai, C.H., Hemphill, D., Endres, M., Rodermel, S., Spalding, M., 1997. Elevated CO 2 effects during leaf on- togeny — a new perspective on acclimation. Plant Physiol. 115, 1195 – 1200. Moore, B.D., Cheng, S.-H., Sims, D., Seeman, J.R., 1999. The biochemical and molecular basis for photosynthetic acclima- tion to elevated atmospheric CO 2 . Plant Cell Environ. 22, 567 – 582. Noode´n, L.D., Leopold, A.C., 1978. Phytohormones and the endogeneous regulation of senescence and abscission. In: Letham, D.S, Goodwin, P.B., Higgins, T.J.V. Eds., Phyto- hormones and Related Compounds: a Comprehensive Trea- tise. Elsevier, Amsterdam, pp. 329 – 369. Noode´n, L.D., Guiamet, J.J., John, I., 1997. Senescence mech- anisms. Physiol. Plant. 101, 746 – 753. Poorter, H., Roumet, C., Campbell, B.D., 1996. Interspecific variation in the growth response of plants to elevated CO 2 : a search for functional types. In: Ko¨rner, C., Bazzaz, F.A. Eds., Carbon Dioxide, Populations, and Communities. Academic Press, San Diego, pp. 375 – 412. Sage, R.F., Sharkey, T.D., Seemann, J.R., 1989. Acclimation of photosynthesis to elevated CO 2 in five C 3 species. Plant Physiol. 89, 590 – 596. Sharkey, T.D., 1988. Estimating the rate of photorespiration in leaves. Physiol. Plant. 73, 147 – 152. Sicher, R.C., Bunce, J.A., 1997. Relationship of photosynthetic acclimation to changes of Rubisco activity in field-grown winter wheat and barley during growth in elevated carbon dioxide. Photosynth. Res. 52, 27 – 38. Sicher, R.C., Bunce, J.A., 1998. Evidence that premature senescence affects photosynthetic decline of wheat flag leaves during growth in elevated carbon dioxide. Int. J. Plant Sci. 159, 798 – 804. Smart, C.M., 1994. Gene expression during leaf senescence. New Phytol. 126, 419 – 448. Stitt, M., Krapp, A., 1999. The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background. Plant Cell Environ. 22, 583 – 621. Theobald, J.C., Mitchell, R.A.C., Parry, M.A.J., Lawlor, D.W., 1998. Estimating the excess investment in ribulose-1,5-bis- phosphate carboxylaseoxygenase in leaves of spring wheat grown under elevated CO 2 . Plant Physiol. 118, 945 – 955. Thompson, J.E., Froese, C.D., Madey, E., Smith, M.D., Hong, Y.W., 1998. Lipid metabolism during plant senescence. Prog. Lipid Res. 37, 119 – 141. Tottman, D.R., Broad, H., 1987. The decimal code for the growth stages of cereals, with illustrations. Ann. Appl. Biol. 110, 441 – 454. Van Kraalingen, D.W.G., 1990. Effects of CO 2 enrichment on nutrient-deficient plants. In: Goudriaan, J., Van Keulen, H., Van Laar, H.H. Eds., The Greenhouse Effect and Primary Productivity in European Agroecosystems. Pudoc, Wa- geningen, pp. 42 – 45. Vermehren, B., Fangmeier, A., Ja¨ger, H.-J., 1998. Influnce of elevated CO 2 on nitrogen economy of wheat. In: Peter, D., Maracchi, G., Ghazi, A. Eds., Climate Change Impact on Agriculture and Forestry. European Commission, Brussels, pp. 497 – 505. Webber, A.N., Nie, G.Y., Long, S.P., 1994. Acclimation of photosynthetic proteins to rising atmospheric CO 2 . Photo- synth. Res. 39, 413 – 425. Weigel, H.J., Manderscheid, R., Ja¨ger, H.-J., Mejer, G.J., 1994. Effects of season-long CO 2 enrichment on cereals. I. Growth performance and yield. Agric. Ecosyst. Environ. 48, 231 – 240. Wilson, J.B., 1997. An evolutionary perspective on the ‘death hormone’ hypothesis in plants. Physiol. Plant. 99, 511 – 516. Yen, C.H., Yang, C.H., 1998. Evidence for programmed cell death during leaf senescence in plants. Plant Cell Physiol. 39, 922 – 927. .