The objectives of this synthesis are (1) to review the factors that influence the ecological, geographical, and palaeoecological distributions of plants possessing C4 photosynthesis and (2) to propose a hypothesis/model to explain both the distribution of C4 plants with respect to temperature and CO2 and why C4 photosynthesis is relatively uncommon in dicotyledonous plants (hereafter dicots), especially in comparison with its widespread distribution in monocotyledonous species (hereafter monocots). Our goal is to stimulate discussion of the factors controlling distributions of C4 plants today, historically, and under future elevated CO2 environments. Understanding the distributions of C3/C4 plants impacts not only primary productivity, but also the distribution, evolution, and migration of both invertebrates and vertebrates that graze on these plants. Sixteen separate studies all indicate that the current distributions of C4 monocots are tightly correlated with temperature: elevated temperatures during the growing season favor C4 monocots. In contrast, the seven studies on C4 dicot distributions suggest that a different environmental parameter, such as aridity (combination of temperature and evaporative potential), more closely describes their distributions. Differences in the temperature dependence of the quantum yield for CO2 uptake (light-use efficiency) of C3 and C4 species relate well to observed plant distributions and light-use efficiency is the only mechanism that has been proposed to explain distributional differences in C3/C4 monocots. Modeling of C3 and C4 light-use efficiencies under different combinations of atmospheric CO2 and temperature predicts that C4-dominated ecosystems should not have expanded until atmospheric CO2 concentrations reached the lower levels that are thought to have existed beginning near the end of the Miocene. At that time, palaeocarbonate and fossil data indicate a simultaneous, global expansion of C4-dominated grasslands. The C4 monocots generally have a higher quantum yield than C4 dicots and it is proposed that leaf venation patterns play a role in increasing the light-use efficiency of most C4 monocots. The reduced quantum yield of most C4 dicots is consistent with their rarity, and it is suggested that C4 dicots may not have been selected until CO2 concentrations reached their lowest levels during glacial maxima in the Quaternary. Given the intrinsic light-use efficiency advantage of C4 monocots, C4 dicots may have been limited in their distributions to the warmest ecosystems, saline ecosystems, and/or to highly disturbed ecosystems. All C4 plants have a significant advantage over C3 plants under low atmospheric CO2 conditions and are predicted to have expanded significantly on a global scale during full-glacial periods, especially in tropical regions. Bog and lake sediment cores as well as pedogenic carbonates support the hypothesis that C4 ecosystems were more extensive during the last glacial maximum and then decreased in abundance following deglaciation as atmospheric CO2 levels increased.

C 4 photosynthesis, atmospheric CO 2 , and climate

In a world of increasing atmospheric CO2, there is intensified interest in the ecophysiology of photosynthesis and more attention is being given to other aspects of carbon exchange and storage in natural ecosystems. For example, how much will the photosynthesis of terrestrial and aquatic vegetation change as global CO2 increases? Are there major ecosystems, such as the boreal forests, which may become important sinks of CO2 and slow down the effects of anthropogenic CO2 emissions on climate? This volume reviews the progress which has been made in understanding photosynthesis in the past few decades at several levels of integration, from the molecular level to canopy, ecosystem and global scales.

Ecophysiology of Photosynthesis

Although trees have responded to global warming in the past - to temperatures higher than they are now - the rate of change predicted in the 21st century is likely to be unprecedented. Greenhouse gas emissions could cause a 3-6°C increase in mean land surface temperature at high and temperate latitudes. Despite this, few experiments have isolated the effects of temperature for this scenario on trees and forests. This review focuses on tree and forest responses at boreal and temperate latitudes, ranging from the cellular to the ecosystem level. Adaptation to varying temperatures revolves around the trade-off between utilizing the full growing season and minimizing frost damage through proper timing of hardening in autumn and dehardening in spring. But the evolutionary change in these traits must be sufficiently rapid to compensate for the temperature changes. Many species have a positive response to increased temperature - but how close are we to the optima? Management is critical for a positive response of forest growth to a warmer climate, and selection of the best species for the new conditions will be of vital importance. Contents Summary 369 I. Introduction 370 II. Photosynthesis and respiration 370 III. Soil organic matter decomposition and mineralization 373 IV. Phenology and frost hardiness 376 V. Whole tree experimental responses to warming 380 VI. Changes in species distribution at warmer temperatures 381 VII. Adaptation and evolution 383 VIII. Ecosystem level responses to warming 387 Acknowledgements 390 References 390 Appendix I. Temperature response functions 399.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1469-8137.2001.00057.x

Tree and forest functioning in response to global warming

Modelling Potential Crop Growth Processes

1. Introduction.- 2. Tree Form and Stem Taper.- 3. Tree-stem Volume Equations.- 4. Tree Weight and Biomass Estimation.- 5. Quantifying Tree Crowns.- 6. Growth Functions.- 7. Evaluating Site Quality.- 8. Quantifying Stand Density.- 9. Indices of Individual-tree Competition.- 10. Modeling Forest Stand Development.- 11. Whole-stand Models for Even-aged Stands.- 12. Diameter-distribution Models for Even-aged Stands.- 13. Size-class Models for Even-aged Stands.- 14. Individual-tree Models for Even-aged Stands.- 15. Growth and Yield Models for Uneven-aged Stands.- 16. Modeling Response to Silvicultural Treatments.- 17. Modeling Wood Characteristics.- 18. Model Implementation and Evaluation.-

Modeling Forest Trees and Stands

Part 1 General topics: dynamic modelling some subjects of general importance transport processes temperature effects of plant and crop processes growth functions biological switches development. Part 2 Plant and crop physiology: light relations in canopies leaf photosynthesis whole-plant respiration and growth energetics biochemical and chemical approaches to plant growth efficiency partitioning during vegetative growth transpiration by a crop canopy crop water relations crop responses root growth. Part 3 Plant morphology: branching phyllotaxis.

Ian R. Johnson

Papers

Plant and Crop Modelling: A Mathematical Approach to Plant and Crop Physiology