Ecological Studies, Vol. 178Analysis and Synthesis

Edited by

M.M. Caldwell, Logan, USAG. Heldmaier, Marburg, GermanyO.L. Lange, Würzburg, GermanyH.A. Mooney, Stanford, USAE.-D. Schulze, Bayreuth, GermanyU. Sommer, Kiel, Germany

Ecological Studies

Volumes published since 1992 are listed at the end of this book.

William K. SmithThomas C. VogelmannChrista CritchleyEditors

Photosynthetic AdaptationChloroplast to Landscape

With 94 illustrations

William K. Smith Thomas C. VogelmannDepartment of Biology 109 Carrigan DriveWakeforest University Burlington, VT 05405-0086Winston-Salem, NC 27109USAsmithwk@wfu.edu

Christa CritchleyDepartment of BotanyUniversity of QueenslandBrisbane 4072Australiac.critchley@botany.uq.edu.au

Cover illustration: Schematic representation of the structural/spatial hierarchy of organizational com-plexity in the plant kingdom (see Figure 1.1 on page 5).

Library of Congress Cataloging-in-Publication DataPhotosynthetic adaptation : chloroplast to landscape / William K. Smith, Thomas C.

Vogelmann, Christa Critchley.p. cm. — (Ecological studies)

Includes bibliographical references and index.ISBN 0-387-22079-8 (hc : alk. paper)

1. Chloroplasts. 2. Photosynthesis. 3. Plants—Adaptation. I. Smith, William K. (WilliamKirby), 1944- II. Vogelmann, Thomas Craig. III. Critchley, Christa. IV. Series.QK725.P48 2004571.6�592—dc22 2004049195

ISBN 0-387-22079-8 Printed on acid-free paper.

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Contents

Preface viiContributors ix

Part 1. Introduction

1. Background and Objectives 3William K. Smith, Thomas C. Vogelmann, and Christa Critchley

Part 2. Sunlight Capture

2. Chloroplast to Leaf 15John R. Evans, Thomas C. Vogelmann, William E. Williams, and Holly L. Gorton

3. Leaf to Landscape 42Alessandro Cescatti and Ülo Niinemets

Part 3. Sunlight Processing

4. Chloroplast to Leaf 89Neil R. Baker, Donald R. Ort, Jeremy Harbinson, and John Whitmarsh

v

Part 4. CO2 Capture

5. Chloroplast to Leaf 107John R. Evans, Ichiro Terashima, Yuko Hanba, and Francesco Loreto

6. Leaf to Landscape 133Mathew Williams, F. Ian Woodward, Dennis D. Baldocchi, andDavid Ellsworth

Part 5. CO2 Processing

7. Chloroplast to Leaf 171Thomas D. Sharkey, Sean E. Weise, Andrew J. Standish, and Ichiro Terashima

8. Leaf to Landscape 207David S. Ellsworth, Ülo Niinemets, and Peter B. Reich

Part 6. Environmental Constraints

9. Chloroplast to Leaf 231Francesco Loreto, Neil R. Baker, and Donald R. Ort

10. Leaf to Landscape 262Stanley D. Smith, Elke Naumberg, Ülo Niinemets, and Matthew J. Germino

Part 7. Overview

11. Summary and Future Perspectives 297William K. Smith, Park S. Nobel, William A. Reiners, Thomas C. Vogelmann, and Christa Critchley

Index 311

vi Contents

Preface

Across the broad structural and spatial hierarchy of organization found in theplant kingdom, chloroplasts are progressively packaged into more complex lev-els of organization, generating newly emerged limitations to photosynthesis ateach level, as well as corresponding adaptive responses. Most likely, the adap-tive advantage of size and height for enhanced sunlight competition also gener-ated selective pressures for adaptations that maximized, for one example, the ef-ficiency of light capture expressed per unit of biomass invested. The primaryobjective of this book is to identify and evaluate quantitatively fundamental adap-tations at each level of the organizational hierarchy (i.e., chloroplast, cell, tissue,leaf, branch, canopy, stand, landscape). Thus, the present volume has been or-ganized into two sections, the capture and processing of sunlight and CO2 acrosstwo structural/spatial hierarchies (chloroplast-to-leaf and leaf-to-landscape). Themost general objective was to identify which levels of the organizational hierar-chy generates the greatest potential for influencing photosynthetic carbon gainin the plant kingdom. Within this framework, the potential effectiveness of meta-bolic (C4 versus C3) versus structural/developmental adaptations (e.g., sun/shadeleaves) on the photosynthetic process is addressed specifically, as well as theirintegration. Such an understanding of the integrated mechanisms of plant formand physiology will be critical for making future decisions about the genomicmanipulation of agricultural species to improve production efficiency and yield,as well as current policies in agroecology and land-use management. Similarly,incorporation of global change impacts on sources, sinks, and sequestration of

vii

photosynthetic carbon will also be critical for understanding future impacts onagricultural and native species.

Essential to answering the fundamental questions posed above is the improvedcapability for measuring photosynthetic light and CO2 capture and processingacross a broad range of structural and spatial scales, for example, from the chloro-plast and cell (e.g., biochemistry, nanomicroscopy) to inside a leaf (e.g., fiberoptic microprobes, fluorescence, laser beams, and photoacoustics), to the wholeleaf, crown and whole plant (e.g., gas exchange chambers, fluorescence, stableisotope techniques), and to the community and landscape level (e.g., tower andaircraft eddy covariance, remote sensing). The above “footprints,” or spatial res-olution for measuring photosynthetic properties, range from nanometers to kilo-meters. Moreover, these measurement capabilities are now producing the com-parative data necessary for evaluating comprehensively the adaptive significanceof the emerging properties upscale that may require the concerted evolution ofphotosynthetic traits at lower levels of organization. These measurement capa-bilities also enable comparisons with adaptations in photosynthetic physiologyacross the broad scale of plant structural organization and complexity.

A workshop was held in conjunction with the Twelfth International Congresson Photosynthesis in Brisbane, Australia (August 18 to 23, 2001) to organize thecontent and contributors in this volume. Funding was provided by the NationalScience Foundation (Ecology and Evolutionary Physiology Program), the U.S.Department of Agriculture (Competitive Grants Program, Plant Biochemistry),and the U.S. Department of Energy (Global Change Program).

William K. SmithThomas C. Vogelmann

Christa Chritchley

viii Preface

Contributors

Neil R. Baker Department of Biological Sciences, University of Essex, Colchester C04 3SQ,United Kingdom

Dennis D. Baldocchi Ecosystem Science Division, Departmentof Environmental Science, Policy andManagement, 151 Hilgard Hall, University of California, Berkeley,Berkeley, CA 94720, USA

Alessandro Cescatti Centro di Ecologia Alpina, I-38040Viote del Monte Bondone, Italy

Christa Critchley Department of Botany, The Universityof Queensland, Brisbane, QLD 4072,Australia

David S. Ellsworth School of Natural Resources and Environment, University of Michigan,Dana Building, 430 E. University, AnnArbor, MI 48109-1115, USA

ix

John R. Evans Environmental Biology Group, ResearchSchool of Biological Sciences, Australian National University, GPOBox 475 Canberra, ACT 2601, Australia

Matthew J. Germino Department of Biology, Idaho State University, Pocatello, ID 83209-8007,USA

Holly L. Gorton Department of Biology, St. Marys College, St. Marys City, MD 20686-3001, USA

Yuko Hanba Research Institute for Bioresources,Okayama University, Kurashiki, 710-0046, Japan

Jeremy Harbinson Horticultural Production Chains (HPC)Group, Marijkeweg 22 (building number527), Wageningen University, 6709 PGWageningen, The Netherlands

Francesco Loreto Consiglio Nazionale delle Ricerche(CNR), Istituto di Biochimica ed Ecofisiologia Vegetali (IBEV), ViaSalaria Km. 29,300, 00016 Monterotondo Scalo (Roma), Italy

Elke Naumburg MWH Energy and Infrastructure, Inc.,760 Whalers Way, Suite A-100, Fort Collins, CO 80525, USA

Ülo Niinemets Department of Plant Physiology, University of Tartu, Riia 23,51011,Tartu, Estonia

Park S. Nobel Department of Organismic Biology,Ecology and Evolution, 621 YoungDrive South, University of California,Los Angeles, Los Angeles, California90095-1606, USA

Donald R. Ort Department of Plant Biology, Universityof Illinois, Urbana, IL 61801, USA

x Contributors

Peter B. Reich Department of Forest Resources, University of Minnesota, 1530 N. Cleveland Avenue, St. Paul, MN 55108,USA

Thomas D. Sharkey Department of Botany, 430 LincolnDrive, University of Wisconsin, Madison, WI 53706, USA

Stanley D. Smith Department of Biological Sciences, University of Nevada, Las Vegas, LasVegas, NV 89154-4004, USA

William K. Smith Department of Biology, Wake ForestUniversity, Winston-Salem, NC 27109-7325, USA

Andrew J. Standish University of Wisconsin-Madison, Department of Botany, 430 Lincoln Dr.,Madison, WI 53706, USA

Ichiro Terashima Department of Biology, Graduate Schoolof Science, Osaka University, 1-16Machikaneyama-cho, Toyonaka, Osaka,560-0043, Japan

Thomas C. Vogelmann Botany and Agricultural Biochemistry,University of Vermont, Burlington, VT05405-0086, USA

Sean E. Weise University of Wisconsin-Madison, Department of Botany, 430 Lincoln Dr.,Madison, WI 53706, USA

John Whitmarsh National Institutes of Health, NationalInstitute of General Medical Sciences,45 Center Drive, Room 2AS55F,Bethesda, Maryland 20892-6200, USA

Mathew Williams Institute of Ecology and Resource Management, University of Edinburgh,Darwin Building, Mayfield Rd, Edinburgh, EH9 3JU, United Kingdom

Contributors xi

William E. Williams Department of Biology, St. Marys College, St. Marys City, MD 20686-3001, USA

Ian Woodward Department of Animal and Plant Sciences, University of Sheffield,Sheffield, S10 2TN, United Kingdom

xii Contributors