ALBERT

Notes: The need to assess the role of forests in the global cycling of carbon and how that role will change as the atmospheric concentration of CO2 increases has spawned many experiments over a range of scales. Experiments using open-top chambers have been established at many sites to test whether the short-term responses of tree seedlings described in controlled environments would be sustained over several growing seasons under field conditions. Here we review the results of those experiments, using the framework of the interacting cycles of carbon, water and nutrients, because that is the framework of the ecosystem models that are being used to address the decades-long response of forests.Our analysis suggests that most of what was learned in seedling studies was qualitatively correct. The evidence from field-grown trees suggests a continued and consistent stimulation of photosynthesis of about 60% for a 300 p.p.m. increase in [CO2], and there is little evidence of the long-term loss of sensitivity to CO2 that was suggested by earlier experiments with tree seedlings in pots. Despite the importance of respiration to a tree's carbon budget, no strong scientific consensus has yet emerged concerning the potential direct or acclimation response of woody plant respiration to CO2 enrichment. The relative effect of CO2 on above-ground dry mass was highly variable and greater than that indicated by most syntheses of seedling studies. Effects of CO2 concentration on static measures of response are confounded with the acceleration of ontogeny observed in elevated CO2. The trees in these open-top chamber experiments were in an exponential growth phase, and the large growth responses to elevated CO2 resulted from the compound interest associated with an increasing leaf area. This effect cannot be expected to persist in a closed-canopy forest where growth potential is constrained by a steady-state leaf area index. A more robust and informative measure of tree growth in these experiments is the annual increment in wood mass per unit leaf area, which increased 27% in elevated CO2. There is no support for the conclusion from many studies of seedlings that root-to-shoot ratio is increased by elevated CO2; the production of fine roots may be enhanced, but it is not clear that this response would persist in a forest. Foliar nitrogen concentrations were lower in CO2-enriched trees, but to a lesser extent than was indicated in seedling studies and only when expressed on a leaf mass basis. The prediction that leaf litter C/N ratio would increase was not supported in field experiments. Also contrasting with seedling studies, there is little evidence from the field studies that stomatal conductance is consistently affected by CO2; however, this is a topic that demands more study.Experiments with trees in open-top chambers under field conditions have provided data on longer-term, larger-scale responses of trees to elevated CO2 under field conditions, confirmed some of the conclusions from previous seedling studies, and challenged other conclusions. There remain important obstacles to using these experimental results to predict forest responses to rising CO2, but the studies are valuable nonetheless for guiding ecosystem model development and revealing the critical questions that must be addressed in new, larger-scale CO2 experiments.

Notes: A two-component model of growth and maintenance respiration is used to study the response of northern red oak (Quercus rubra L.) seedlings and 32-year-old trees to sub-ambient (10 μmol h; cumulative dose based on 7 h daily mean), ambient (43 μmol h), and twice-ambient (85 μmolh) ozone. The relative growth rates (RGR) of leaves sampled from seedlings and trees were similar across treatments, as were specific leaf respiration rates (SRR). Growth coefficients estimated from the SRR versus RGR relationship averaged 25-3 mol CO2 kg−1 leaf dry mass produced for seedlings and 21-5 mol kg−1 for trees. Maintenance coefficients ranged from 0-89 to 1-07 mol CO2 kg−1 leaf dry mass d−1 for seedlings and from 0-64 to 0-84 mol kg-1 d−1 for trees. Neither coefficient was affected by ozone. Leaves sampled throughout the growing season also showed little response of respiration to ozone. This occurred despite a 30% reduction in net photosynthesis for trees grown at twice-ambient ozone. These results suggest that growth and maintenance respiration in young northern red oak leaves are not affected by ozone and that in older leaves injury can occur without a parallel increase in so-called ‘maintenance’ respiration.

Notes: Light-saturated photosynthetic and stomatal responses to elevated CO2 were measured in upper and mid-canopy foliage of a sweetgum (Liquidambar styraciflua L) plantation exposed to free-air CO2 enrichment (FACE) for 3 years, to characterize environmental interactions with the sustained CO2 effects in an intact deciduous forest stand. Responses were evaluated in relation to one another, and to seasonal patterns and natural environmental stresses, including high  temperatures, vapour pressure deficits (VPD), and drought. Photosynthetic CO2 assimilation (A) averaged 46% higher in the +200 µmol mol−1 CO2 treatment, in mid- and upper canopy foliage. Stomatal conductance (gs) averaged 14% (mid-canopy) and 24% (upper canopy) lower under CO2 enrichment. Variations in the relative responses of A and gs were linked, such that greater relative stimulation of A was observed on dates when relative reductions in gs were slight. Dry soils and high VPD reduced gs and A in both treatments, and tended to diminish treatment differences. The absolute effects of CO2 on A and gs were minimized whenever gs was low (〈0·15 mol m−2 s−1), but relative effects, as the ratio of elevated to ambient rates, varied greatly under those conditions. Both stomatal and non-stomatal limitations of A were involved during late season droughts. Leaf temperature had a limited influence on A and gs, and there was no detectable relationship between prevailing temperature and CO2 effects on A or gs. The responsiveness of A and gs to elevated CO2, both absolute and relative, was maintained through time and within the canopy of this forest stand, subject to seasonal constraints and variability associated with limiting air and soil moisture.

Notes: Long-term exposure of plants to elevated [CO2] leads to a number of growth and physiological effects, many of which are interpreted in the context of ameliorating the negative impacts of drought. However, despite considerable study, a clear picture in terms of the influence of elevated [CO2] on plant water relations and the role that these effects play in determining the response of plants to elevated [CO2] under water-limited conditions has been slow to emerge. In this paper, four areas of research are examined that represent critical, yet uncertain, themes related to the response of plants to elevated [CO2] and drought. These include (1) fine-root proliferation and implications for whole-plant water uptake; (2) enhanced water-use efficiency and consequences for drought tolerance; (3) reductions in stomatal conductance and impacts on leaf water potential; and (4) solute accumulation, osmotic adjustment and dehydration tolerance of leaves. A survey of the literature indicates that the growth of plants at elevated [CO2] can lead to conditions whereby plants maintain higher (less negative) leaf water potentials. The mechanisms that contribute to this effect are not fully known, although CO2-induced reductions in stomatal conductance, increases in whole-plant hydraulic conductance and osmotic adjustment may be important. Less understood are the interactive effects of elevated [CO2] and drought on fine-root production and water-use efficiency, and the contribution of these processes to plant growth in water-limited environments. Increases in water-use efficiency and reductions in water use can contribute to enhanced soil water content under elevated [CO2]. Herbaceous crops and grasslands are most responsive in this regard. The conservation of soil water at elevated [CO2] in other systems has been less studied, but in terms of maintaining growth or carbon gain during drought, the benefits of CO2-induced improvements in soil water content appear relatively minor. Nonetheless, because even small effects of elevated [CO2] on plant and soil water relations can have important implications for ecosystems, we conclude that this area of research deserves continued investigation. Future studies that focus on cellular mechanisms of plant response to elevated [CO2] and drought are needed, as are whole-plant investigations that emphasize the integration of processes throughout the soil–plant–atmosphere continuum. We suggest that the hydraulic principles that govern water transport provide an integrating framework that would allow CO2-induced changes in stomatal conductance, leaf water potential, root growth and other processes to be uniquely evaluated within the context of whole-plant hydraulic conductance and water transport efficiency.

Notes: Responses of photosynthesis and stomatal conductance were monitored throughout a 3-year field exposure of Liriodendron tulipifera (yellow-poplar) and Quercus alba (white oak) to elevated concentrations of atmospheric CO2. Exposure to atmospheres enriched with +150 and +300 umol mol-1 CO2 increased net photosynthesis by 12–144% over the course of the study. Net photosynthesis was consistently higher at +300 than at +150 umol mol-1 CO2. The effect of CO2 enrichment on stomatal conductance was limited, but instantaneous leaf-level water use efficiency increased significantly. No decrease in the responsiveness of photosynthesis to CO2 enrichment over time was detected, and the responses were consistent throughout the canopy and across successive growth flushes and seasons. The relationships between internal CO2 concentration and photosynthesis (e.g. photosynthetic capacity and carboxylation efficiency) were not altered by growth at elevated concentrations of CO2. No alteration in the timing of leaf senescence or abscission was detected, suggesting that the seasonal duration of effective gas-exchange was unaffected by CO2 treatment. These results are consistent with data previously reported for these species in controlled-environment studies, and suggest that leaf-level photosynthesis does not down-regulate in these species as a result of acclimation to CO2 enrichment in the field. This sustained enhancement of photosynthesis provides the opportunity for increased growth and carbon storage by trees as the atmospheric concentration of CO2 rises, but many additional factors interact in determining whole-plant and forest responses to global change.

Notes: Abstract. The construction and evaluation of a temperature-corrected in situ thermocouple psychrometer for measurement of leaf water potential (Ψ) is described. The instrument utilized two chromel-constantan thermocouples which allowed for detection of both the psychrometric zero offset and the temperature differential between the sample and the Peltier measuring junction. The psychrometer was subjected to stable temperature gradients while in contact with reference solutions of sodium chloride, and the effects of thermal gradients were quantified. Regression analysis indicated that temperature differentials were responsible for errors in water potential determinations of approximately –7.73 MPa°C−1. When installed on leaves of field-grown cotton (Gossypium hirsutum L.), corn (Zea mays L.) and soybean (Glycine max L. Merr) the instrument detected temperature differentials up to 0.1°C (–6.0 μV) which were associated with relatively small shifts in psychrometric zero offsets (–0.05––0.75 μV). Results indicated that substantial errors in apparent Ψ were caused by non-isothermal conditions between the leaf and the psychrometer measuring junction. The relative magnitude of these errors could be quantified and the corrected results showed good agreement with conventional psychrometric determination of Ψ on excised samples during a diurnal cycle.

Notes: Abstract. Cuticular resistance to water vapour diffusion is an important aspect of thermocouple psychrometry and may introduce significant error in the measurement of leaf water potential (Ψ). The effect of the citrus (Citrus mitis Blanco) leaf cuticle on water vapour movement was studied using the times required for vapour pressure equilibration during thermocouple psychrometric measurement of Ψ. Cuticular abrasion with various carborundum powders was used to reduce the diffusive resistance of both the adaxial and abaxial leaf surfaces, and the extent of the disruption to the leaf was investigated with light and electron microscopy.Cuticular abrasion resulted in reduced equilibration times due to decreased cuticular resistance and greater water vapour movement between the leaf and the psychrometer chamber. Equilibration times were reduced from over 5 h in the unabraded control leaves to 1 h with cuticle abrasion. This was associated with the decrease in diffusive resistance with cuticular abrasion from over 55 s cm−1 to less than 8 s cm−1 for both the adaxial and abaxial leaf surfaces.Scanning electron micrographs of the abraded leaf tissue revealed considerable disruption of the stomatal ledge and of the guard cells, surface smoothing and displacement of waxes into the stomatal aperture, and damage to veins. Observations with the transmission electron microscope revealed frequent disruption of epidermal cell walls, and damage to both the cytoplasmic and vacuolar membranes.

Notes: Abstract Two of the major uncertainties in forecasting future terrestrial sources and sinks of CO2 are the CO2-enhanced growth response of forests and soil warming effects on net CO2 efflux from forests. Carbon dioxide enrichment of tree seedlings over time periods less than 1 yr has generally resulted in enhanced rates of photosynthesis, decreased respiration, and increased growth, with minor increases in leaf area and small changes in C allocation. Exposure of woody species to elevated CO2 over several years has shown that high rates of photosynthesis may be sustained, but net C accumulation may not necessarily increase if CO2 release from soil respiration increases. The impact of the 25% rise in atmospheric CO2 with industrialization has been examined in tree ring chronologies from a range of species and locations. In contrast to the seedling tree results, there is no convincing evidence for CO2-enhanced stem growth of mature trees during the last several decades. However, if mature trees show a preferential root growth response to CO2 enrichment, the gain in root mass for an oak-hickory forest in eastern Tennessee is estimated to be only 9% over the last 40 years. Root data bases are inadequate for detecting such an effect. A very small shift in ecosystem nutrients from soil to vegetation could support CO2-enhanced growth. Climate warming and the accompanying increase in mean soil temperature could have a greater effect than CO2 enrichment on terrestrial sources and sinks of CO2. Soil respiration and N mineralization have been shown to increase with soil temperature. If plant growth increases with increased N availability, and more C is fixed in growth than is released by soil respiration, then a negative feedback on climate warming will occur. If warming results in a net increase in CO2 efflux from forests, then a positive feedback will follow. A 2 to 4°C increase in soil temperature could increase CO2 efflux from soil by 15 to 32% in eastern deciduous forests. Quantifying C budget responses of forests to future global change scenarios will be speculative until mature tree responses to CO2 enrichment and the effects of temperature on terrestrial sources and sinks of CO2 can be determined.