Canopy conundrums: building on the Biosphere 2 experience to scale measurements of inner and outer canopy photoprotection from the leaf to the landscape
Caroline J. Nichol A , Roland Pieruschka B , Kotaro Takayama C , Britta Förster D , Zbigniew Kolber E , Uwe Rascher B , John Grace A , Sharon A. Robinson F , Barry Pogson D and Barry Osmond D F GA School of GeoSciences, University of Edinburgh, West Mains Road, Edinburgh EH9 3JN, Scotland, UK.
B Institute for Bio- and Geosciences IBG 2: Plant Sciences, Forschungszentrum Jülich, 52425 Jülich, Germany.
C Laboratory of Physiological Green Systems, Department of Biomechanical Systems, Faculty of Agriculture, Ehime University, 3-5-7, Tarumi, Matsuyama 790-8566, Japan.
D Plant Sciences Division, Research School of Biology, Australian National University, Canberra, ACT 0200, Australia.
E Ocean Sciences, University of California Santa Cruz, 1156 High Street, Santa Cruz, CA 95064, USA.
F Institute for Conservation Biology and Ecosystem Management, School of Biological Sciences, University of Wollongong, Wollongong, NSW 2522, Australia.
G Corresponding author. Email: osmond@grapevine.net.au
Functional Plant Biology 39(1) 1-24 https://doi.org/10.1071/FP11255
Submitted: 12 November 2011 Accepted: 2 December 2011 Published: 12 January 2012
Abstract
Recognising that plant leaves are the fundamental productive units of terrestrial vegetation and the complexity of different environments in which they must function, this review considers a few of the ways in which these functions may be measured and potentially scaled to the canopy. Although canopy photosynthetic productivity is clearly the sum of all leaves in the canopy, we focus on the quest for ‘economical insights’ from measurements that might facilitate integration of leaf photosynthetic activities into canopy performance, to better inform modelling based on the ‘insights of economics’. It is focussed on the reversible downregulation of photosynthetic efficiency in response to light environment and stress and summarises various xanthophyll-independent and dependent forms of photoprotection within the inner and outer canopy of woody plants. Two main themes are developed. First, we review experiments showing the retention of leaves that grow old in the shade may involve more than the ‘payback times’ required to recover the costs of their construction and maintenance. In some cases at least, retention of these leaves may reflect selection for distinctive properties that contribute to canopy photosynthesis through utilisation of sun flecks or provide ‘back up’ capacity following damage to the outer canopy. Second, we report experiments offering hope that remote sensing of photosynthetic properties in the outer canopy (using chlorophyll fluorescence and spectral reflectance technologies) may overcome problems of access and provide integrated measurements of these properties in the canopy as a whole. Finding appropriate tools to scale photosynthesis from the leaf to the landscape still presents a challenge but this synthesis identifies some measurements and criteria in the laboratory and the field that improve our understanding of inner and outer canopy processes.
Additional keywords: avocado, chlorophyll fluorescence, LIFT, lutein-epoxide cycle, NPQ, PAM photoacclimation, photoinhibition, PRI, violaxanthin cycle.
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