Tansley Review No. 36 Excited leaves
10.1111/j.1469-8137.1992.tb02935.x
Crossref journal-article
Wiley
New Phytologist (311)
Abstract

SUMMARYPhotosynthesis is largely to do with energy transduction; the conversion of light energy into electrical energy into chemical energy. Precisely how much light energy is needed to bring about the reduction of one molecule of carbon dioxide and the release of one molecule of oxygen (the quantum requirement) is a matter of fundamental importance and one which has attracted much past controversy. This article concludes that a minimum quantum requirement of eight, as demanded by the Z‐scheme, is obviously consistent with much contemporary work which puts the measured value for C3 leaves close to nine. Moreover, while values of less than eight (obtained in some circumstances with micro‐organisms), are a reminder that nothing is beyond challenge they are not, in the absence of confirmation and extension, sufficiently compelling to demand rejection of either the Z‐scheme or current measuring procedures. This article also shows why, even if the underlying minimum requirement was now accepted beyond all reasonable doubt, there would still be very good reasons for continuing, indefinitely, to measure actual photosynthetic efficiency in the natural environment. It discusses some of the implications of the fact that all plants, if not stressed, appear to photosynthesize at the same rate in low light. It explains the role of fluorescence in its relation to quantum yield, the possibility that the rate of photosynthesis might be determined from fluorescence measurements alone, and that a combination of fluorescence and gas exchange measurements could provide new information about the manner in which ‘dark respiration’ is affected by light. It indicates how contemporary interest in all of these matters has focused attention on the necessity for safe dissipation of excitation energy by leaves and on the manner by which this might be achieved. CONTENTS Summary 325 I. Excitation 325 II. Quantum requirement 326 III. Learning from fluorescence 335 IV. Safely dissipated 340 Acknowledgements 342 References 342

Bibliography

WALKER, D. (1992). Tansley Review No. 36 Excited leaves. New Phytologist, 121(3), 325–345. Portico.

Authors 1
  1. DAVID WALKER (first)
References 151 Referenced 80
  1. {'key': 'e_1_2_1_2_1', 'first-page': '228', 'volume-title': 'Topics in Photosynthesis', 'author': 'Asada K.', 'year': '1987'} / Topics in Photosynthesis by Asada K. (1987)
  2. 10.1073/pnas.87.15.5930
  3. {'key': 'e_1_2_1_4_1', 'first-page': '159', 'volume-title': 'Photosynthesis I. Encyclopedia of Plant Physiology. New Series', 'author': 'Avron M.', 'year': '1987'} / Photosynthesis I. Encyclopedia of Plant Physiology. New Series by Avron M. (1987)
  4. 10.1104/pp.99.1.124
  5. 10.1111/j.1399-3054.1991.tb05101.x
  6. {'key': 'e_1_2_1_7_1', 'first-page': '91', 'volume-title': 'Photosynthetic Mechanisms and the Environment. Topics in Photosynthesis', 'author': 'Barber J.', 'year': '1985'} / Photosynthetic Mechanisms and the Environment. Topics in Photosynthesis by Barber J. (1985)
  7. {'key': 'e_1_2_1_8_1', 'first-page': '137', 'volume-title': 'Topics in Photosynthesis', 'author': 'Barber J.', 'year': '1987'} / Topics in Photosynthesis by Barber J. (1987)
  8. 10.1007/BF00033159
  9. 10.1104/pp.91.2.542
  10. 10.1007/BF00402983
  11. 10.1098/rstb.1895.0010
  12. 10.1098/rstb.1895.0011
  13. 10.1111/j.1751-1097.1991.tb03668.x
  14. 10.1016/0005-2728(81)90113-4
  15. 10.1007/BF00035202 / Photosynthesis Research / The analysis of photosynthetic performance in leaves under field conditions: a case study using Bruguiera mangroves by Cheeseman J. M. (1991)
  16. {'key': 'e_1_2_1_17_1', 'first-page': '41', 'article-title': 'Monitor and control of blood and tissue oxygen tension', 'volume': '2', 'author': 'Clark L. C.', 'year': '1956', 'journal-title': 'Transactions of the American Society for Artificial Internal Organs'} / Transactions of the American Society for Artificial Internal Organs / Monitor and control of blood and tissue oxygen tension by Clark L. C. (1956)
  17. {'key': 'e_1_2_1_18_1', 'first-page': '171', 'article-title': 'Partitioning of photosynthetic electron flow between CO2 and O2 reduction in a C3 leaf (Phaseolus vulgaris L.) at different CO2 concentrations and during drought stress', 'volume': '171', 'author': 'Cornic G.', 'year': '1991', 'journal-title': 'Planta'} / Planta / Partitioning of photosynthetic electron flow between CO2 and O2 reduction in a C3 leaf (Phaseolus vulgaris L.) at different CO2 concentrations and during drought stress by Cornic G. (1991)
  18. 10.1016/0038-092X(57)90052-X
  19. 10.1111/j.1469-8137.1981.tb07480.x
  20. 10.1104/pp.73.3.534
  21. 10.1071/PP9880163
  22. 10.1071/PP9880151
  23. 10.1104/pp.84.2.218
  24. 10.1104/pp.87.1.17
  25. 10.1104/pp.92.2.293
  26. {'key': 'e_1_2_1_27_1', 'first-page': '375', 'volume-title': 'Photosynthesis', 'author': 'Demmig‐Adams B.', 'year': '1989'} / Photosynthesis by Demmig‐Adams B. (1989)
  27. {'key': 'e_1_2_1_28_1', 'volume-title': 'C3, C4, Mechanisms, and Cellular and Environmental Regulation of Photosynthesis', 'author': 'Edwards G. E.', 'year': '1983'} / C3, C4, Mechanisms, and Cellular and Environmental Regulation of Photosynthesis by Edwards G. E. (1983)
  28. 10.1104/pp.59.1.86
  29. 10.1146/annurev.pp.09.060158.000245
  30. 10.1104/pp.30.6.504
  31. 10.1104/pp.13.1.157
  32. 10.2307/2436774
  33. 10.1002/j.1537-2197.1941.tb11010.x
  34. 10.1002/j.1537-2197.1943.tb14744.x
  35. 10.1007/978-3-642-67242-2_13
  36. 10.1016/0005-2728(88)90078-3
  37. {'key': 'e_1_2_1_38_1', 'first-page': '1', 'volume-title': 'Photosynthesis I. Encyclopedia of Plant Physiology, New Series', 'author': 'Forti G.', 'year': '1987'} / Photosynthesis I. Encyclopedia of Plant Physiology, New Series by Forti G. (1987)
  38. 10.1007/BF00035457
  39. 10.1016/0003-9861(53)90419-6
  40. 10.1016/S0304-4165(89)80016-9
  41. 10.1098/rspb.1991.0091
  42. {'key': 'e_1_2_1_43_1', 'first-page': '429', 'volume-title': 'Photosynthesis I. Encyclopedia of Plant Physiology. New Series', 'author': 'Good N. E.', 'year': '1977'} / Photosynthesis I. Encyclopedia of Plant Physiology. New Series by Good N. E. (1977)
  43. 10.1007/978-3-642-67242-2_12
  44. {'key': 'e_1_2_1_45_1', 'first-page': '505', 'volume-title': 'Biochemistry of Chloroplasts', 'author': 'Grant B. R.', 'year': '1967'} / Biochemistry of Chloroplasts by Grant B. R. (1967)
  45. 10.1104/pp.90.3.1029
  46. 10.1016/0005-2728(69)90116-9
  47. 10.1007/BF00403020
  48. {'key': 'e_1_2_1_49_1', 'first-page': '581', 'volume-title': 'Proceedings of the International Congress of Plant Physiology Neic Delhi', 'author': 'Heber U.', 'year': '1989'} / Proceedings of the International Congress of Plant Physiology Neic Delhi by Heber U. (1989)
  49. {'key': 'e_1_2_1_50_1', 'first-page': '17', 'volume-title': 'Perspectives in Biochemical and Genetic Regulation of Photosynthesis', 'author': 'Heber U.', 'year': '1990'} / Perspectives in Biochemical and Genetic Regulation of Photosynthesis by Heber U. (1990)
  50. {'key': 'e_1_2_1_51_1', 'first-page': '110', 'article-title': 'Photoinhibitory damage to chloroplasts under phosphate deficiency and alleviation of deficiency and damage by photorespiratory reactions', 'volume': '44', 'author': 'Heber U.', 'year': '1989', 'journal-title': 'Zeitschrift für Naturforschung'} / Zeitschrift für Naturforschung / Photoinhibitory damage to chloroplasts under phosphate deficiency and alleviation of deficiency and damage by photorespiratory reactions by Heber U. (1989)
  51. 10.1038/186136a0
  52. 10.1104/pp.91.1.331
  53. {'key': 'e_1_2_1_54_1', 'first-page': '135', 'volume-title': 'Photosynthetic Mechanisms and the Environment', 'author': 'Horton P.', 'year': '1985'} / Photosynthetic Mechanisms and the Environment by Horton P. (1985)
  54. {'key': 'e_1_2_1_55_1', 'first-page': '111', 'volume-title': 'Current Research in Photosynthesis', 'author': 'Horton P.', 'year': '1989'} / Current Research in Photosynthesis by Horton P. (1989)
  55. {'key': 'e_1_2_1_56_1', 'first-page': '259', 'article-title': 'Chlorophyll fluorescence transients', 'volume': '4', 'author': 'Horton P.', 'year': '1990', 'journal-title': 'Methods in Plant Biochemistry'} / Methods in Plant Biochemistry / Chlorophyll fluorescence transients by Horton P. (1990)
  56. {'key': 'e_1_2_1_57_1', 'first-page': '145', 'volume-title': 'Perspectives in Biochemical and Genetic Regulation of Photosynthesis', 'author': 'Horton P.', 'year': '1990'} / Perspectives in Biochemical and Genetic Regulation of Photosynthesis by Horton P. (1990)
  57. 10.1016/0014-5793(91)80819-O
  58. 10.1007/BF01516164
  59. 10.1098/rspb.1990.0066
  60. 10.1016/0005-2728(82)90117-7
  61. {'key': 'e_1_2_1_62_1', 'first-page': '1', 'article-title': 'A critical consideration of the quantum yield of Chlorella‐photosynthesis', 'volume': '13', 'author': 'Kok B.', 'year': '1948', 'journal-title': 'Enzymologia'} / Enzymologia / A critical consideration of the quantum yield of Chlorella‐photosynthesis by Kok B. (1948)
  62. 10.1016/0006-3002(49)90136-5
  63. {'key': 'e_1_2_1_64_1', 'first-page': '209', 'article-title': 'Photo‐induced interactions of metabolism of green plant cells', 'volume': '5', 'author': 'Kok B.', 'year': '1951', 'journal-title': 'Symposium for the Society of Experimental Biology'} / Symposium for the Society of Experimental Biology / Photo‐induced interactions of metabolism of green plant cells by Kok B. (1951)
  64. 10.1007/978-3-642-94798-8_22
  65. 10.1007/BF00033172
  66. 10.1071/PP9900579
  67. 10.1071/PP9910267
  68. Krall J. P.&Edwards G. E.(July 1992).Relationship between photosystem II activity and CO2fixation in leaves.Physiologia Plantarum(in the press). (10.1111/j.1399-3054.1992.tb01328.x)
  69. 10.1071/PP9910369
  70. 10.1016/0005-2728(73)90019-4
  71. {'key': 'e_1_2_1_72_1', 'first-page': '145', 'volume-title': 'Photoinhibition', 'author': 'Krause G. H.', 'year': '1987'} / Photoinhibition by Krause G. H. (1987)
  72. 10.1007/BF00028527
  73. 10.1146/annurev.pp.42.060191.001525
  74. {'key': 'e_1_2_1_75_1', 'volume-title': 'Kinetics of Photosynthesis and Photorespiration in C3 Plants', 'author': 'Laisk A.', 'year': '1977'} / Kinetics of Photosynthesis and Photorespiration in C3 Plants by Laisk A. (1977)
  75. 10.1098/rstb.1989.0017
  76. 10.1098/rspb.1989.0057
  77. 10.1007/BF00403593
  78. {'key': 'e_1_2_1_79_1', 'article-title': 'Steady state, induction and oscillatory kinetics of photosynthetic electron transport related to donor side oxidation and acceptor side reduction of photosystem I in sunflower leaves. Evidence for photosynthetic control and functional Cyt b/f‐PSI‐FNR complexes', 'author': 'Laisk A.', 'year': '1992', 'journal-title': 'Photosynthetica'} / Photosynthetica / Steady state, induction and oscillatory kinetics of photosynthetic electron transport related to donor side oxidation and acceptor side reduction of photosystem I in sunflower leaves. Evidence for photosynthetic control and functional Cyt b/f‐PSI‐FNR complexes by Laisk A. (1992)
  79. {'key': 'e_1_2_1_80_1', 'first-page': '554', 'article-title': 'Oscillations in photosynthesis are initiated and supported by imbalances in the supply of ATP and NADPH to the Calvin cycle', 'volume': '182', 'author': 'Laisk A.', 'year': '1991', 'journal-title': 'Planta'} / Planta / Oscillations in photosynthesis are initiated and supported by imbalances in the supply of ATP and NADPH to the Calvin cycle by Laisk A. (1991)
  80. 10.1098/rspb.1989.0058
  81. {'key': 'e_1_2_1_82_1', 'first-page': '203', 'volume-title': 'Primary Processes in Photosynthesis', 'author': 'Lavorell J.', 'year': '1977'} / Primary Processes in Photosynthesis by Lavorell J. (1977)
  82. 10.1007/978-94-009-0511-5_668
  83. {'key': 'e_1_2_1_84_1', 'first-page': '175', 'volume-title': '‘Photosynthesis’ New Comprehensive Biochemistry', 'author': 'MacDonald F. D.', 'year': '1987'} / ‘Photosynthesis’ New Comprehensive Biochemistry by MacDonald F. D. (1987)
  84. 10.1016/S0167-7306(08)60135-0
  85. 10.1016/0003-9861(51)90082-3
  86. 10.1016/0003-9861(51)90012-4
  87. {'key': 'e_1_2_1_88_1', 'first-page': '178', 'volume-title': 'Physiology of Stomata', 'author': 'Meidner H.', 'year': '1968'} / Physiology of Stomata by Meidner H. (1968)
  88. 10.1016/0005-2728(84)90036-7
  89. 10.1515/znc-1989-3-415
  90. 10.1016/0005-2728(90)90063-A
  91. 10.1016/S0005-2728(05)80143-4
  92. 10.1146/annurev.pp.35.060184.002215
  93. 10.1016/0304-4173(81)90006-9
  94. 10.1111/j.1469-8137.1987.tb04688.x
  95. 10.1016/0005-2728(88)90128-4
  96. 10.1104/pp.88.4.1125
  97. 10.1104/pp.88.1.158
  98. 10.1111/j.1469-8137.1986.tb00794.x
  99. 10.1002/jctb.503300105
  100. 10.1002/jctb.280330105 / Journal of Chemical Technology and Biotechnolog / A tubular bioreactor for photosynthetic production of biomass from carbon dioxide: design and performance by Pirt S. J. (1983)
  101. 10.1098/rspb.1984.0007
  102. 10.1007/978-3-642-66505-9_6
  103. 10.1016/0005-2728(90)90062-9
  104. 10.1016/0014-5793(89)81723-5
  105. 10.1007/BF00035539
  106. 10.1016/S0005-2728(05)80221-X
  107. 10.1007/BF00041833
  108. 10.1007/BF00033169
  109. {'key': 'e_1_2_1_110_1', 'first-page': '173', 'article-title': 'Contrasting pH‐Optima of light‐driven O2‐ and H2O2, –reduction in spinach chloroplasts as measured via chlorophyll fluorescence quenching', 'volume': '46', 'author': 'Schreiber U.', 'year': '1991', 'journal-title': 'Zeitschrift für Naturforschung'} / Zeitschrift für Naturforschung / Contrasting pH‐Optima of light‐driven O2‐ and H2O2, –reduction in spinach chloroplasts as measured via chlorophyll fluorescence quenching by Schreiber U. (1991)
  110. 10.1007/BF00024185
  111. 10.1098/rspb.1990.0099
  112. {'key': 'e_1_2_1_113_1', 'volume-title': 'Trends in Photosynthesis Research', 'author': 'Seaton G. G. R.', 'year': '1990'} / Trends in Photosynthesis Research by Seaton G. G. R. (1990)
  113. 10.1111/j.1399-3054.1988.tb09205.x
  114. 10.1007/BF00395423
  115. 10.1104/pp.75.1.95
  116. {'key': 'e_1_2_1_117_1', 'first-page': '9', 'volume-title': 'Photosynthesis and Physiology of the Whole Plant', 'author': 'Sivak M. N.', 'year': '1985'} / Photosynthesis and Physiology of the Whole Plant by Sivak M. N. (1985)
  117. 10.1016/0003-9861(85)90304-2
  118. 10.1007/BF00395152
  119. 10.1111/j.1365-3040.1985.tb01679.x
  120. 10.1111/j.1469-8137.1986.tb00826.x
  121. {'key': 'e_1_2_1_122_1', 'first-page': '105', 'volume-title': 'Hungarian‐USA Binational Symposium on Photosynthesis', 'author': 'Sivak M. N.', 'year': '1987'} / Hungarian‐USA Binational Symposium on Photosynthesis by Sivak M. N. (1987)
  122. {'key': 'e_1_2_1_123_1', 'first-page': '319', 'volume-title': 'Perspectives in Biochemical and Genetic Regulation of Photosynthesis', 'author': 'Stokes D.', 'year': '1990'} / Perspectives in Biochemical and Genetic Regulation of Photosynthesis by Stokes D. (1990)
  123. 10.1038/1991247a0
  124. 10.1093/oxfordjournals.aob.a087034
  125. {'issue': '6', 'key': 'e_1_2_1_126_1', 'first-page': '1023', 'article-title': 'Intra‐leaf and intracellular gradients in chloroplast ultrastructure of dorsiventral leaves illuminated from the adaxial or abaxial side during their development', 'volume': '27', 'author': 'Terashima I.', 'year': '1986', 'journal-title': 'Plant Cell Physiology'} / Plant Cell Physiology / Intra‐leaf and intracellular gradients in chloroplast ultrastructure of dorsiventral leaves illuminated from the adaxial or abaxial side during their development by Terashima I. (1986)
  126. 10.1007/978-94-009-4384-1_20
  127. 10.1111/j.1469-8137.1973.tb02027.x
  128. {'key': 'e_1_2_1_129_1', 'first-page': '1', 'volume-title': 'Energy, Plants & Man', 'author': 'Walker D. A.', 'year': '1979'} / Energy, Plants & Man by Walker D. A. (1979)
  129. 10.1007/BF00383899
  130. 10.1098/rstb.1989.0013
  131. Walker D. A.(1990).The Use of the Oxygen Electrode and Fluorescence Probes in Simple Measurements of Photosynthesis 2nd impression pp.1–203.
  132. {'key': 'e_1_2_1_133_1', 'first-page': '93', 'volume-title': 'Regulation of Carbon Partitioning in Photosynthetic Tissues', 'author': 'Walker D. A.', 'year': '1985'} / Regulation of Carbon Partitioning in Photosynthetic Tissues by Walker D. A. (1985)
  133. {'key': 'e_1_2_1_134_1', 'first-page': '829', 'article-title': 'Can phosphate limit photosynthetic carbon assimilation in vivo', 'volume': '23', 'author': 'Walker D. A.', 'year': '1985', 'journal-title': 'Physiologie Vegetate'} / Physiologie Vegetate / Can phosphate limit photosynthetic carbon assimilation in vivo by Walker D. A. (1985)
  134. 10.1016/0968-0004(86)90136-2
  135. 10.1098/rstb.1986.0039
  136. {'key': 'e_1_2_1_137_1', 'first-page': '227', 'article-title': 'Energetik der Photosynthese', 'volume': '15', 'author': 'Warburg O.', 'year': '1952', 'journal-title': 'Die Naturwissenschaften'} / Die Naturwissenschaften / Energetik der Photosynthese by Warburg O. (1952)
  137. 10.1126/science.128.3315.68
  138. {'key': 'e_1_2_1_139_1', 'first-page': '410', 'article-title': 'The maximum efficiency of photosynthesis', 'volume': '25', 'author': 'Warburg O.', 'year': '1950', 'journal-title': 'Archives of Biochemistry'} / Archives of Biochemistry / The maximum efficiency of photosynthesis by Warburg O. (1950)
  139. 10.1515/znb-1952-0301
  140. 10.1002/j.1537-2197.1969.tb09747.x
  141. 10.1515/zpch-1923-10614 / Zeitschift für Physikalische Chemie / Uber den Einfluss der Wellenlange auf den Energieumsalz bei der Kohlensaure‐assimilation by Warburg O. (1923)
  142. 10.1007/978-94-009-3535-8_131
  143. 10.1016/0005-2728(87)90190-3
  144. {'key': 'e_1_2_1_145_1', 'first-page': '307', 'volume-title': 'Current Research in Photosynthesis', 'author': 'Weis E.', 'year': '1989'} / Current Research in Photosynthesis by Weis E. (1989)
  145. {'key': 'e_1_2_1_146_1', 'first-page': '772', 'article-title': 'Participation of the Mehler Reaction and catalase in the oxygen exchange of chloroplast preparations', 'volume': '22', 'author': 'Whitehouse D.G.', 'year': '1971', 'journal-title': 'Journal of Experimental Biology'} / Journal of Experimental Biology / Participation of the Mehler Reaction and catalase in the oxygen exchange of chloroplast preparations by Whitehouse D.G. (1971)
  146. 10.1016/0304-4173(79)90008-9
  147. 10.1146/annurev.pp.39.060188.002533
  148. 10.1111/j.1438-8677.1991.tb00231.x
  149. 10.1351/pac197951030639
  150. {'key': 'e_1_2_1_151_1', 'volume-title': 'Photosynthesis, Photorespiration and Plant Productivity', 'author': 'Zelitch I.', 'year': '1971'} / Photosynthesis, Photorespiration and Plant Productivity by Zelitch I. (1971)
  151. {'key': 'e_1_2_1_152_1', 'first-page': '123', 'article-title': 'Pathways of carbon fixation in green plants', 'volume': '44', 'author': 'Zelitch I.', 'year': '1985', 'journal-title': 'Annual Review of Plant Physiology'} / Annual Review of Plant Physiology / Pathways of carbon fixation in green plants by Zelitch I. (1985)
Dates
Type When
Created 19 years, 4 months ago (April 29, 2006, 11:18 a.m.)
Deposited 2 years ago (Aug. 31, 2023, 11:33 p.m.)
Indexed 3 weeks, 6 days ago (Aug. 6, 2025, 8:05 a.m.)
Issued 33 years, 2 months ago (July 1, 1992)
Published 33 years, 2 months ago (July 1, 1992)
Published Online 19 years, 4 months ago (April 28, 2006)
Published Print 33 years, 2 months ago (July 1, 1992)
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@article{WALKER_1992, title={Tansley Review No. 36 Excited leaves}, volume={121}, ISSN={1469-8137}, url={http://dx.doi.org/10.1111/j.1469-8137.1992.tb02935.x}, DOI={10.1111/j.1469-8137.1992.tb02935.x}, number={3}, journal={New Phytologist}, publisher={Wiley}, author={WALKER, DAVID}, year={1992}, month=jul, pages={325–345} }