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Box 9E. 1 Continued FIGURE 2. The C-S-R triangle model (Grime 1979). The strategies at the three corners are C, competiti- winning species; S, stress-tolerating s- cies; R,ruderalspecies. Particular species can engage in any mixture of these three primary strategies, and the m- ture is described by their position within the triangle. comment briefly on some other dimensions that Grime's (1977) triangle (Fig. 2) (see also Sects. 6. 1 are not yet so well understood. and 6. 3 of Chapter 7 on growth and allocation) is a two-dimensional scheme. A C-S axis (Com- tition-winning species to Stress-tolerating spe- Leaf Economics Spectrum cies) reflects adaptation to favorable vs. unfavorable sites for plant growth, and an R- Five traits that are coordinated across species are axis (Ruderal species) reflects adaptation to leaf mass per area (LMA), leaf life-span, leaf N disturbance. concentration, and potential photosynthesis and dark respiration on a mass basis. In the five-trait Trait-Dimensions space,79%ofallvariation worldwideliesalonga single main axis (Fig. 33 of Chapter 2A on photo- A recent trend in plant strategy thinking has synthesis; Wright et al. 2004). Species with low been trait-dimensions, that is, spectra of varia- LMA tend to have short leaf life-spans, high leaf tion with respect to measurable traits. Compared nutrient concentrations, and high potential rates of mass-based photosynthesis. These species with category schemes, such as Raunkiaer's, trait occur at the ''quick-return'' end of the leaf e- dimensions have the merit of capturing cont- nomics spectrum.

Plant Physiological Ecology

Contents Foreword (by David T. Clarkson) About the authors Foreword to the first edition by David T. Clarkson) Acknowledgments Abbreviations 1. ASSUMPTIONS AND APPROACHES Introduction-History, Assumptions, and Approaches 1 What is Ecophysiology? 2 The Roots of Ecophysiology 3 Physiological Ecology and the Distribution of Organisms 4 Time Scale of Plant Responses to Environment 5 Conceptual and Experimental Approaches 6 New Directions in Ecophysiology 7 The Structure of the Book References 2. PHOTOSYNTHESIS, RESPIRATION, AND LONG-DISTANCE TRANSPORT 2A. PHOTOSYNTHESIS 1 Introduction 2 General Characteristics of the Photosynthetic Apparatus 2.1 The 'Light' and 'Dark' Reactions of Photosynthesis 2.1.1 Absorption of Photons 2.1.2 Fate of the Excited Chlorophyll 2.1.3 Membrane-bound Photosynthetic Electron Transport and Bioenergetics 2.1.4 Photosynthetic Carbon Reduction 2.1.5 Oxygenation and Photorespiration 2.2 Supply and Demand of CO2 in the Photosynthetic Process 2.2.1 The CO2-response Curve 2.2.2 Supply of CO2 - Stomatal and Boundary layer Conductances 2.2.3 The Internal Conductance 3 Response of Photosynthesis to Light 3.1 Characterization of the Light Climate under a Leaf Canopy 3.2 Physiological, Biochemical, and Anatomical Differences between Sun and Shade Leaves 3.2.1 The Light-response Curve of Sun and Shade Leaves 3.2.2 Anatomy and Ultrastructure of Sun and Shade Leaves 3.2.3 Biochemical Differences between Shade and Sun Leaves 3.2.4 The Light-response Curve of Sun and Shade Leaves Revisited 3.2.5 The Environmental signal for Shade Acclimation inChloroplasts 3.3 Effects of Excess Irradiance 3.3.1 Photoinhibition - Protection by Carotenoids of the Xanthophyll Cycle 3.3.2 Chloroplast Movement in Response to Changes in Irradiance 3.4 Responses to Variable Irradiance 3.4.1 Photosynthetic Induction 3.4.2 Light Activation of Rubisco 3.4.3 Post-illumination CO2 Assimilation and Sunfleck Utilization Efficiency 3.4.4 Metabolite Pools in Sun and Shade Leaves 3.4.5 Net Effect of Sunflecks on Carbon Gain and Growth 4 Partitioning of the Products of Photosynthesis and Regulation by 'feedback' 4.1 Partitioning within the Cell 4.2 Regulation of the Rate of Photosynthesis by feedback 4.3 Sugar-induced Repression of Genes Encoding for Calvin-cycle Enzymes 4.4 Ecological impacts Mediated by source-sink Interactions 5 Responses to Availability of Water 5.1 Regulation of Stomatal Opening 5.2 The A-Ci Curve as Affected by Water Stress 5.3 Carbon isotope Discrimination in Relation to Water-use Efficiency 5.4 Other sources of Variation in Carbon isotope ratios in C3 Plants 6 Effects of Nutrient Supply on Photosynthesis 6.1 The Photosynthesis-nitrogen Relationship 6.2 Interactions of Nitrogen, Light and Water 6.3 Photosynthesis, Nitrogen, and Leaf Life-span 7 Photosynthesis and Leaf Temperature: Effects and Adaptations 7.1 Effects of High Temperatures on Photosynthesis 7.2 Effects of Low Temperatures on Photosynthesis 8 Effects of Air Pollutants on Photosynthesis 9 C4 Plants 9.1 Introduction 9.2 Biochemical and Anatomical Aspects 9.3 Physiology of C4 Photosynthesis 9.4 Intercellular and Intracellular Transport of Metabolites of the C4 Pathway 9.5 Photosynthetic Nitrogen-use Efficiency, Water-use Efficienc