Contents

 Summary 1

I. What is FACE? 2
II. Materials and methods 2
III. Photosynthetic carbon uptake 3
IV. Acclimation of photosynthesis 6
V. Growth, above-ground production and yield 8
VI. So, what have we learned? 10

 Acknowledgements 11


 References 11


Appendix 1. References included in the database for meta-analyses  14


 Appendix 2. Results of the meta-analysis of FACE effects 18



Summary
Free-air CO2 enrichment (FACE) experiments allow study of the effects of elevated [CO2] on plants and ecosystems grown under natural conditions without enclosure. Data from 120 primary, peer-reviewed articles describing physiology and production in the 12 large-scale FACE experiments (475–600 ppm) were collected and summarized using meta-analytic techniques. The results confirm some results from previous chamber experiments: light-saturated carbon uptake, diurnal C assimilation, growth and above-ground production increased, while specific leaf area and stomatal conductance decreased in elevated [CO2]. There were differences in FACE. Trees were more responsive than herbaceous species to elevated [CO2]. Grain crop yields increased far less than anticipated from prior enclosure studies. The broad direction of change in photosynthesis and production in elevated [CO2] may be similar in FACE and enclosure studies, but there are major quantitative differences: trees were more responsive than other functional types; C4 species showed little response; and the reduction in plant nitrogen was small and largely accounted for by decreased Rubisco. The results from this review may provide the most plausible estimates of how plants in their native environments and field-grown crops will respond to rising atmospheric [CO2]; but even with FACE there are limitations, which are also discussed.

What have we learned from 15 years of free-air CO2 enrichment (FACE)? A meta-analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2.

The effects of land use change on soil carbon stocks are of concern in the context of international policy agendas on greenhouse gas emissions mitigation. This paper reviews the literature for the influence of land use changes on soil C stocks and reports the results of a meta analysis of these data from 74 publications. The meta analysis indicates that soil C stocks decline after land use changes from pasture to plantation (−10%), native forest to plantation (−13%), native forest to crop (−42%), and pasture to crop (−59%). Soil C stocks increase after land use changes from native forest to pasture (+ 8%), crop to pasture (+ 19%), crop to plantation (+ 18%), and crop to secondary forest (+ 53%). Wherever one of the land use changes decreased soil C, the reverse process usually increased soil carbon and vice versa. As the quantity of available data is not large and the methodologies used are diverse, the conclusions drawn must be regarded as working hypotheses from which to design future targeted investigations that broaden the database. Within some land use changes there were, however, sufficient examples to explore the role of other factors contributing to the above conclusions. One outcome of the meta analysis, especially worthy of further investigation in the context of carbon sink strategies for greenhouse gas mitigation, is that broadleaf tree plantations placed onto prior native forest or pastures did not affect soil C stocks whereas pine plantations reduced soil C stocks by 12–15%.

Soil carbon stocks and land use change: a meta analysis

Plant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 000 out of the world's 300 000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation – but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models.

https://onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2486.2011.02451.x

TRY - a global database of plant traits

Stomata, the small pores on the surfaces of leaves and stalks, regulate the flow of gases in and out of leaves and thus plants as a whole. They adapt to local and global changes on all timescales from minutes to millennia. Recent data from diverse fields are establishing their central importance to plant physiology, evolution and global ecology. Stomatal morphology, distribution and behaviour respond to a spectrum of signals, from intracellular signalling to global climatic change. Such concerted adaptation results from a web of control systems, reminiscent of a 'scale-free' network, whose untangling requires integrated approaches beyond those currently used.

http://www.bio.bristol.ac.uk/research/HetheringtonLab/pdfs/nature01843.pdf

The role of stomata in sensing and driving environmental change.

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2007.01641.x

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions.

• Data from 13 long-term (> 1 yr), field-based studies of the effects of elevated CO2 concentration ([CO2]) on European forest tree species were analysed using meta-analysis and modelling. Meta-analysis was used to determine mean responses across the data sets, and data were fitted to two commonly used models of stomatal conductance in order to explore response to environmental conditions and the relationship with assimilation.
• Meta-analysis indicated a significant decrease (21%) in stomatal conductance in response to growth in elevated [CO2] across all studies. The response to [CO2] was significantly stronger in young trees than old trees, in deciduous compared to coniferous trees, and in water stressed compared to nutrient stressed trees. No evidence of acclimation of stomatal conductance to elevated [CO2] was found.
• Fits of data to the first model showed that growth in elevated [CO2] did not alter the response of stomatal conductance to vapour pressure deficit, soil water content or atmospheric [CO2]. Fits of data to the second model indicated that conductance and assimilation responded in parallel to elevated [CO2] except when water was limiting.
• Data were compared to a previous meta-analysis and it was found that the response of gs to elevated [CO2] was much more consistent in long-term (> 1 yr) studies, emphasising the need for long-term elevated [CO2] studies. By interpreting data in terms of models, the synthesis will aid future modelling studies of responses of forest trees to elevated [CO2].

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1046/j.1469-8137.2001.00028.x

Stomatal conductance of forest species after long‐term exposure to elevated CO2 concentration: a synthesis

The effects of elevated atmospheric CO2 concentration on growth of forest tree species are difficult to predict because practical limitations restrict experiments to much shorter than the average life-span of a tree. Long-term, processbased computer models must be used to extrapolate from shorter-term experiments. A key problem is to ensure a strong flow of information between experiments and models. In this study, meta-analysis techniques were used to summarize a suite of photosynthetic model parameters obtained from 15 field-based elevated [CO2] experiments on European forest tree species. The parameters studied are commonly used in modelling photosynthesis, and include observed light-saturated photosynthetic rates (Amax), the potential electron transport rate (Jmax), the maximum Rubisco activity (Vcmax) and leaf nitrogen concentration on mass (Nm) and area (Na) bases. Across all experiments, light-saturated photosynthesis was strongly stimulated by growth in elevated [CO2]. However, significant down-regulation of photosynthesis was also observed; when measured at the same CO2 concentration, photosynthesis was reduced by 10‐20%. The underlying biochemistry of photosynthesis was affected, as shown by a down-regulation of the parameters Jmax and Vcmax of the order of 10%. This reduction in Jmax and Vcmax was linked to the effects of elevated [CO2] on leaf nitrogen concentration. It was concluded that the current model is adequate to model photosynthesis in elevated [CO2]. Tables of model parameter values for different European forest species are given.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.1999.00523.x

P. Roberntz

Papers

Stomatal conductance of forest species after long‐term exposure to elevated CO2 concentration: a synthesis

Effects of elevated (CO2) on photosynthesis in European forest species: a meta-analysis of model parameters

Tradition as asset or burden for transitions from forests as cropping systems to multifunctional forest landscapes: Sweden as a case study