KEY WORDS: CO2 and plants, CO2 and photosynthesis, CO2 and stomata, CO2 and respiration, plants and climate change ABSTRACTThe primary effect of the response of plants to rising atmospheric CO 2 (C a ) is to increase resource use efficiency. Elevated C a reduces stomatal conductance and transpiration and improves water use efficiency, and at the same time it stimulates higher rates of photosynthesis and increases light-use efficiency. Acclimation of photosynthesis during long-term exposure to elevated C a reduces key enzymes of the photosynthetic carbon reduction cycle, and this increases nutrient use efficiency. Improved soil-water balance, increased carbon uptake in the shade, greater carbon to nitrogen ratio, and reduced nutrient quality for insect and animal grazers are all possibilities that have been observed in field studies of the effects of elevated C a . These effects have major consequences for agriculture and native ecosystems in a world of rising atmospheric C a and climate change. CONTENTS

Carbon-use efficiency (CUE), the ratio of net primary production (NPP) to gross primary production (GPP), describes the capacity of forests to transfer carbon (C) from the atmosphere to terrestrial biomass. It is widely assumed in many landscape-scale carbon-cycling models that CUE for forests is a constant value of $ 0.5. To achieve a constant CUE, tree respiration must be a constant fraction of canopy photosynthesis. We conducted a literature survey to test the hypothesis that CUE is constant and universal among forest ecosystems. Of the 60 data points obtained from 26 papers published since 1975, more than half reported values of GPP that were not estimated independently from NPP; values of CUE calculated from independent estimates of GPP were greater than those calculated from estimates of GPP derived from NPP. The slope of the relationship between NPP and GPP for all forests was 0.53, but values of CUE varied from 0.23 to 0.83 for different forest types. CUE decreased with increasing age, and a substantial portion of the variation among forest types was caused by differences in stand age. When corrected for age the mean value of CUE was greatest for temperate deciduous forests and lowest for boreal forests. CUE also increased as the ratio of leaf mass-to-total mass increased. Contrary to the assumption of constancy, substantial variation in CUE has been reported in the literature. It may be inappropriate to assume that respiration is a constant fraction of GPP as adhering to this assumption may contribute to incorrect estimates of C cycles. A 20% error in current estimates of CUE used in landscape models (i.e. ranging from 0.4 to 0.6) could misrepresent an amount of C equal to total anthropogenic emissions of CO 2 when scaled to the terrestrial biosphere.

Estimates of forest net primary production (NPP) demand accurate estimates of root production and turnover. We assessed root turnover with the use of an isotope tracer in two forest free-air carbon dioxide enrichment experiments. Growth at elevated carbon dioxide did not accelerate root turnover in either the pine or the hardwood forest. Turnover of fine root carbon varied from 1.2 to 9 years, depending on root diameter and dominant tree species. These long turnover times suggest that root production and turnover in forests have been overestimated and that sequestration of anthropogenic atmospheric carbon in forest soils may be lower than currently estimated.

31I.31II.32III.34IV.35V.37VI.38VII.3941References41 Summary The rhizosphere priming effect (RPE) is a mechanism by which plants interact with soil functions. The large impact of the RPE on soil organic matter decomposition rates (from 50% reduction to 380% increase) warrants similar attention to that being paid to climatic controls on ecosystem functions. Furthermore, global increases in atmospheric CO2 concentration and surface temperature can significantly alter the RPE. Our analysis using a game theoretic model suggests that the RPE may have resulted from an evolutionarily stable mutualistic association between plants and rhizosphere microbes. Through model simulations based on microbial physiology, we demonstrate that a shift in microbial metabolic response to different substrate inputs from plants is a plausible mechanism leading to positive or negative RPEs. In a case study of the Duke Free‐Air CO2 Enrichment experiment, performance of the PhotoCent model was significantly improved by including an RPE‐induced 40% increase in soil organic matter decomposition rate for the elevated CO2 treatment – demonstrating the value of incorporating the RPE into future ecosystem models. Overall, the RPE is emerging as a crucial mechanism in terrestrial ecosystems, which awaits substantial research and model development.

A postulated role of the CN-resistant alternative respiratory pathway in plants is the maintenance of mitochondrial electron transport at low temperatures that would otherwise inhibit the main phosphorylating pathway and prevent the formation of toxic reactive oxygen species. This role is supported by the observation that alternative oxidase protein levels often increase when plants are subjected to growth at low temperatures. We used oxygen isotope fractionation to measure the distribution of electrons between the main and alternative pathways in mung bean (Vigna radiata) and soybean (Glycine max) following growth at low temperature. The amount of alternative oxidase protein in mung bean grown at 19°C increased over 2-fold in both hypocotyls and leaves compared with plants grown at 28°C but was unchanged in soybean cotyledons grown at 14°C compared with plants grown at 28°C. When the short-term response of tissue respiration was measured over the temperature range of 35°C to 9°C, decreases in the activities of both main and alternative pathway respiration were observed regardless of the growth temperature, and the relative partitioning of electrons to the alternative pathway generally decreased as the temperature was lowered. However, cold-grown mung bean plants that upregulated the level of alternative oxidase protein maintained a greater electron partitioning to the alternative oxidase when measured at temperatures below 19°C supporting a role for the alternative pathway in response to low temperatures in mung bean. This response was not observed in soybean cotyledons, in which high levels of alternative pathway activity were seen at both high and low temperatures.