Tansley Review No. 36 Excited leaves

Photosynthesis 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.