Forests exchange large amounts of CO2 with the atmosphere and can influence and be influenced by atmospheric CO2. There has been a recent proliferation of literature on the effects of atmospheric CO2 on forest trees. More than 300 studies of trees on five different continents have been published in the last five years. These include an increasing number of field studies with a long-term focus and involving CO2×stress or environment interactions. The recent data on long-term effects of elevated atmospheric CO2 on trees indicate a potential for a persistent enhancement of tree growth for several years, although the only relevant long-term datasets currently available are for juvenile trees.

The current literature indicates a significantly larger average long-term biomass increment under elevated CO2 for conifers (130%) than for deciduous trees (49%) in studies not involving stress components. However, stimulation of photosynthesis by elevated CO2 in long-term studies was similar for conifers (62%) and deciduous trees (53%). Recent studies indicate that elevated CO2 causes a more persistent stimulation of biomass increment and photosynthesis than previously expected. Results of seedling studies, however, might not be applicable to other stages of tree development because of complications of age-dependent and size-dependent shifts in physiology and carbon allocation, which are accelerated by elevated CO2. In addition, there are many possible avenues to down-regulation, making the predicted canopy CO2 exchange and growth of mature trees and forests in a CO2-rich atmosphere uncertain. Although, physiological down-regulation of photosynthetic rates has been documented in field situations, it is rarely large enough to offset entirely photosynthetic gains in elevated CO2. A persistent growth stimulation of individual mature trees has been demonstrated although this effect is more uncertain in trees in natural stands.

Resource interactions can both constrain tree responses to elevated CO2 and be altered by them. Although drought can reduce gas-exchange rates and offset the benefits of elevated CO2, even in well watered trees, stomatal conductance is remarkably less responsive to elevated CO2 than in herbaceous species. Stomata of a number of tree species have been demonstrated to be unresponsive to elevated CO2. We conclude that positive effects of CO2 on leaf area can be at least as important in determining canopy transpiration as negative, direct effects of CO2 on stomatal aperture. With respect to nutrition, elevated CO2 has the potential to alter tree–soil interactions that might influence future changes in ecosystem productivity. There is continued evidence that in most cases nutrient limitations diminish growth and photosynthetic responses to elevated CO2 at least to some degree, and that elevated CO2 can accelerate the appearance of nutrient limitations with increasing time of treatment. In many studies, tree biomass responses to CO2 are artefacts in the sense that they are merely responses to CO2-induced changes in internal nutritional status of the tree.

There are numerous interactions between CO2 and factors of the biotic and abiotic environment. The importance of increasing atmospheric CO2 concentrations for productivity is likely to be overestimated if these are not taken into account. Many interactions, however, are simply additive rather than synergistic or antagonistic. This appears to hold true for many parameters under elevated CO2 in combination with temperature, elevated O3, and other atmospheric pollutants. However, there is currently little evidence that elevated CO2 will counteract O3 damage. When the foliage content of C, mineral nutrients and secondary metabolites is altered by elevated CO2, tree×insect interactions are modified. In most trees, mycorrhizal interactions might be less important for direct effects of CO2 than for alleviating general nutrient deficiencies.

Since many responses to elevated CO2 and their interactions with stress show considerable variability among species/genotypes, one principal research need is for comparative studies of a large variety of woody species and ecosystems under realistic conditions. We still need more long-term experiments on mature trees and stands to address critical scaling issues likely to advance our understanding of responses to elevated CO2 at different stages of forest development and their interactions with climate and environment. The only tools available at present for coping with the consequences of rising CO2 are management of resources and selection of genotypes suitable for the future climate and environment.

Tree and forest functioning in an enriched CO2 atmosphere

A free-air CO2 enrichment (FACE) system was designed to permit the experimental exposure of tall vegetation such as stands of forest trees to elevated atmospheric CO2 concentrations ([CO2]a) without enclosures that alter tree microenvironment. We describe a prototype FACE system currently in operation in forest plots in a maturing loblolly pine (Pinus taeda L.) stand in North Carolina, USA. The system uses feedback control technology to control [CO2] in a 26 m diameter forest plot that is over 10 m tall, while monitoring the 3D plot volume to characterize the whole-stand CO2 regime achieved during enrichment. In the second summer season of operation of the FACE system, atmospheric CO2 enrichment was conducted in the forest during all daylight hours for 96.7% of the scheduled running time from 23 May to 14 October with a preset target [CO2] of 550 μmol mol–1, ≈ 200 μmol mol–1 above ambient [CO2]. The system provided spatial and temporal control of [CO2] similar to that reported for open-top chambers over trees, but without enclosing the vegetation. The daily average daytime [CO2] within the upper forest canopy at the centre of the FACE plot was 552 ± 9 μmol mol–1 (mean ± SD). The FACE system maintained 1-minute average [CO2] to within ± 110 μmol mol–1 of the target [CO2] for 92% of the operating time. Deviations of [CO2] outside of this range were short-lived (most lasting < 60 s) and rare, with fewer than 4 excursion events of a minute or longer per day. Acceptable spatial control of [CO2] by the system was achieved, with over 90% of the entire canopy volume within ± 10% of the target [CO2] over the exposure season. CO2 consumption by the FACE system was much higher than for open-top chambers on an absolute basis, but similar to that of open-top chambers and branch bag chambers on a per unit volume basis. CO2 consumption by the FACE system was strongly related to windspeed, averaging 50 g CO2 m–3 h–1 for the stand for an average windspeed of 1.5 m s–1 during summer. The [CO2] control results show that the free-air approach is a tractable way to study long-term and short-term alterations in trace gases, even within entire tall forest ecosystems. The FACE approach permits the study of a wide range of forest stand and ecosystem processes under manipulated [CO2]a that were previously impossible or intractable to study in true forest ecosystems.

A free‐air enrichment system for exposing tall forest vegetation to elevated atmospheric CO2

A review of greenhouse engineering developments during the 1990s

A closed CO2 and temperature-controlled, long-term chamber system has been developed and set up in a typical boreal forest of Scots pine (Pinus sylvestris L.) near the Mekrijarvi Research Station (62°47′N, 30°58′E, 145 m above sea level) belonging to the University of Joensuu, Finland. The main objectives of the experiment were to provide a means of assessing the medium to long-term effects of elevated atmospheric CO2 concentration (EC) and temperature (ET) on photosynthesis, respiration, growth, and biomass at the whole-tree level and to measure instantaneous whole-system CO2 exchange. The system consists of 16 chambers with individual facilities for controlling CO2 concentration, temperature, and the combination of the two. The chambers can provide a wide variety of climatic conditions that are similar to natural regimes. In this experiment the target CO2 concentration in the EC chambers was set at a fixed constant of 700 µmol mol−1 and the target air temperature in the ET chambers to track the ambient temperature but with a specified addition. Chamber performance was assessed on the base of recordings covering three consecutive years. The CO2 and temperature control in these closed chambers was in general accurate and reliable. CO2 concentration in the EC chambers was within 600–725 µmol mol−1 for 90 % of the exposure time during the "growing-season" (15 April – 15 September) and 625–725 µmol mol−1 for 88 % of the time in the "off-season" (16 September – 14 April), while temperatures in the chambers were within ±2.0 °C of the ambient or target temperature in the "growing season" and within ±3.0 °C in the "off season". There were still some significant chamber effects. Solar radiation in the chambers was reduced by 50–60 % for 82 % of the time in the "growing season" and 55–65 % for 78 % of the time in the "off season", and the relative humidity of the air was increased by 5–10 % for 72 % of the time in the "growing season" and 2–12 % for 91 % of the time in the "off season". The crown architecture and main phenophase of the trees were not modified significantly by enclosure in the chambers, but some physiological parameters changed significantly, e.g., the radiant energy-saturated photosynthesis rate, transpiration rate, maximum photochemical efficiency of photosystem 2, and chlorophyll content.

Controlled Environment Chambers for Investigating Tree Response to Elevated CO2 and Temperature Under Boreal Conditions

A free-air CO2 enrichment (FACE) system was designed to permit the experimental exposure of tall vegetation such as stands of forest trees to elevated atmospheric CO2 concentrations ([CO2]a) without enclosures that alter tree microenvironment. We describe a prototype FACE system currently in operation in forest plots in a maturing loblolly pine (Pinus taeda L.) stand in North Carolina, USA. The system uses feedback control technology to control [CO2] in a 26 m diameter forest plot that is over 10 m tall, while monitoring the 3D plot volume to characterize the whole-stand CO2 regime achieved during enrichment. In the second summer season of operation of the FACE system, atmospheric CO2 enrichment was conducted in the forest during all daylight hours for 96.7% of the scheduled running time from 23 May to 14 October with a preset target [CO2 ]o f 550 μmol mol ‐1 , µ 200 μmol mol ‐1 above ambient [CO2]. The system provided spatial and temporal control of [CO2] similar to that reported for open-top chambers over trees, but without enclosing the vegetation. The daily average daytime [CO2] within the upper forest canopy at the centre of the FACE plot was 552 6 9 μmol mol ‐1 (mean 6 SD). The FACE system maintained 1-minute average [CO2] to within 6 110 μmol mol ‐1 of the target [CO2] for 92% of the operating time. Deviations of [CO2] outside of this range were short-lived (most lasting , 60 s) and rare, with fewer than 4 excursion events of a minute or longer per day. Acceptable spatial control of [CO2] by the system was achieved, with over 90% of the entire canopy volume within 6 10% of the target [CO2] over the exposure season. CO2 consumption by the FACE system was much higher than for open-top chambers on an absolute basis, but similar to that of open-top chambers and branch bag chambers on a per unit volume basis. CO2 consumption by the FACE system was strongly related to windspeed, averaging 50 g CO2 m ‐3 h ‐1 for the stand for an average windspeed of 1.5 m s ‐1 during summer. The [CO2] control results show that the free-air approach is a tractable way to study long-term and short-term alterations in trace gases, even within entire tall forest ecosystems. The FACE approach permits the study of a wide range of forest stand and ecosystem processes under manipulated [CO2]a that were previously impossible or intractable to study in true forest ecosystems.

A. Lockwood

Papers

Performance of a controlled-ventilation open-top chamber for climate change research

Detecting surface condensation on tomato plants in greenhouses