Elevated CO2 did not affect the hydrological balance of a mature native Eucalyptus woodland

Elevated atmospheric CO₂ concentration (eC_a) might reduce forest water‐use, due to decreased transpiration, following partial stomatal closure, thus enhancing water‐use efficiency and productivity at low water availability. If evapotranspiration (E_t) is reduced, it may subsequently increase soil water storage (ΔS) or surface runoff (R) and drainage (D_g), although these could be offset or even reversed by changes in vegetation structure, mainly increased leaf area index (L). To understand the effect of eC_a in a water‐limited ecosystem, we tested whether 2 years of eC_a (~40% increase) affected the hydrological partitioning in a mature water‐limited Eucalyptus woodland exposed to Free‐Air CO₂ Enrichment (FACE). This timeframe allowed us to evaluate whether physiological effects of eC_a reduced stand water‐use irrespective of L, which was unaffected by eC_a in this timeframe. We hypothesized that eC_a would reduce tree‐canopy transpiration (E_tree), but excess water from reduced E_tree would be lost via increased soil evaporation and understory transpiration (E_floor) with no increase in ΔS, R or D_g. We computed E_t, ΔS, R and D_g from measurements of sapflow velocity, L, soil water content (θ), understory micrometeorology, throughfall and stemflow. We found that eC_a did not affect E_tree, E_floor, ΔS or θ at any depth (to 4.5 m) over the experimental period. We closed the water balance for dry seasons with no differences in the partitioning to R and D_g between C_a levels. Soil temperature and θ were the main drivers of E_floor while vapour pressure deficit‐controlled E_tree, though eC_a did not significantly affect any of these relationships. Our results suggest that in the short‐term, eC_a does not significantly affect ecosystem water‐use at this site. We conclude that water‐savings under eC_a mediated by either direct effects on plant transpiration or by indirect effects via changes in L or soil moisture availability are unlikely in water‐limited mature eucalypt woodlands.     

Canopy leaf area of a mature evergreen Eucalyptus woodland does not respond to elevated atmospheric [CO2] but tracks water availability

Canopy leaf area, quantified by the leaf area index (L), is a crucial driver of forest productivity, water use and energy balance. Because L responds to environmental drivers, it can represent an important feedback to climate change, but its responses to rising atmospheric [CO₂] and water availability of forests have been poorly quantified. We studied canopy leaf area dynamics for 28 months in a native evergreen Eucalyptus woodland exposed to free‐air CO₂ enrichment (the EucFACE experiment), in a subtropical climate where water limitation is common. We hypothesized that, because of expected stimulation of productivity and water‐use efficiency, L should increase with elevated [CO₂]. We estimated L from diffuse canopy transmittance, and measured monthly leaf litter production. Contrary to expectation, L did not respond to elevated [CO₂]. We found that L varied between 1.10 and 2.20 across the study period. The dynamics of L showed a quick increase after heavy rainfall and a steady decrease during periods of low rainfall. Leaf litter production was correlated to changes in L, both during periods of decreasing L (when no leaf growth occurred) and during periods of increasing L (active shedding of old foliage when new leaf growth occurred). Leaf lifespan, estimated from mean L and total annual litter production, was up to 2 months longer under elevated [CO₂] (1.18 vs. 1.01 years; P = 0.05). Our main finding that L was not responsive to elevated CO₂ is consistent with other forest FACE studies, but contrasts with the positive response of L commonly predicted by many ecosystem models.     

Atmospheric CO2 concentration may directly affect leaf respiration measurement in tobacco,but not respiration itself*

When atmospheric CO₂ concentration increases, various consequences for plant metabolism have been suggested, such as changes in photosynthesis, photorespiration or respiration which can affect growth and carbon sequestration. In addition to long‐term (indirect) effects on respiration, short‐term (direct) effects of CO₂ concentration on the respiration of leaves, shoots and roots are described in the literature. In most cases, respiration is reported to be inhibited by increased CO₂ concentration, but the mechanism(s) are not yet understood. It has been shown previously that, when the respective technical problems and properties of a gas exchange system are fully considered, a short‐term increase in CO₂ (up to 4200 µmol mol^?1) had no effect on respiration of Phaseolus or Populus leaves (Jahnke, Plant, Cell and Environment 24, 1139–1151, 2001). However, in the present study, large (apparent) CO₂ effects were found with mature Nicotiana leaves whereas, in young leaves, the effect was absent. The experimental results clearly show that the observed direct CO₂ effect on dark CO₂ efflux in the mature tobacco leaves was caused by leakage of CO₂ inside the leaves (and the magnitude of the effect was dependent on the size of the leakage). Nicotiana leaves are, in contrast to Phaseolus and Populus leaves (which are heterobaric), characterized by a homobaric anatomy in which intercellular air spaces are not compartmented and provide a continuous system of open pores in the lateral (paradermal) direction of the leaves. Mesophyll porosity increases with leaf development, which explains the differences between young and mature tobacco leaves. When internal leakage was experimentally restricted, the CO₂ inhibition on CO₂ efflux was no longer observed. It is concluded that the measured direct CO₂ effect(s) on leaf CO₂ efflux in the dark are artefactual, and that a true direct CO₂ effect on leaf respiration does not exist.     

Understorey productivity in temperate grassy woodland responds to soil water availability but not to elevated [CO2]

Rising atmospheric [CO₂] and associated climate change are expected to modify primary productivity across a range of ecosystems globally. Increasing aridity is predicted to reduce grassland productivity, although rising [CO₂] and associated increases in plant water use efficiency may partially offset the effect of drying on growth. Difficulties arise in predicting the direction and magnitude of future changes in ecosystem productivity, due to limited field experimentation investigating climate and CO₂ interactions. We use repeat near‐surface digital photography to quantify the effects of water availability and experimentally manipulated elevated [CO₂] (eCO₂) on understorey live foliage cover and biomass over three growing seasons in a temperate grassy woodland in south‐eastern Australia. We hypothesised that (i) understorey herbaceous productivity is dependent upon soil water availability, and (ii) that eCO₂ will increase productivity, with greatest stimulation occurring under conditions of low water availability. Soil volumetric water content (VWC) determined foliage cover and growth rates over the length of the growing season (August to March), with low VWC (<0.1 m³ m^?3) reducing productivity. However, eCO₂ did not increase herbaceous cover and biomass over the duration of the experiment, or mitigate the effects of low water availability on understorey growth rates and cover. Our findings suggest that projected increases in aridity in temperate woodlands are likely to lead to reduced understorey productivity, with little scope for eCO₂ to offset these changes.     

Water availability affects seasonal CO2‐induced photosynthetic enhancement in herbaceous species in a periodically dry woodland

Elevated atmospheric CO₂ (eCO₂) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g_s) and increased soil water content (V_SWC) and second, through increased leaf internal CO₂ (C_i) and decreased stomatal limitations (S_lim). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C₃ herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO₂‐induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%–50% with a 150 μmol mol^?1 increase in atmospheric CO₂ across seasons. This eCO₂‐induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V_SWC. The highest photosynthetic enhancement by eCO₂ (>30%) was observed during the most water‐limited period, for example, with V_SWC <0.07 in this sandy surface soil. Under eCO₂ there was neither a significant decrease in g_s in the three herbaceous species, nor increases in V_SWC, indicating no “water‐savings effect” of eCO₂. Periods of low V_SWC showed lower g_s (less than ≈ 0.12 mol m^?2 s^?1), higher relative S_lim (>30%) and decreased C_i under the ambient CO₂ concentration (aCO₂), with leaf photosynthesis strongly carboxylation‐limited. The alleviation of S_lim by eCO₂ was facilitated by increasing C_i, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO₂, controls g_s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO₂ has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the “water‐savings effect” usually invoked in grasslands.     

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time‐to‐mortality in plants exposed to prolonged drought, while elevated [CO₂] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO₂] and temperature on the inter‐dependent carbon and hydraulic characteristics associated with drought‐induced mortality in Eucalyptus radiata seedlings grown in two [CO₂] (400 and 640 μL L^?1) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO₂] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO₂], may be the primary contributors to drought‐induced seedling mortality under future climates.     

Elevated CO2 does not stimulate carbon sink in a semi‐arid grassland

Elevated CO₂ is widely accepted to enhance terrestrial carbon sink, especially in arid and semi‐arid regions. However, great uncertainties exist for the CO₂ fertilisation effects, particularly when its interactions with other global change factors are considered. A four‐factor (CO₂, temperature, precipitation and nitrogen) experiment revealed that elevated CO₂ did not affect either gross ecosystem productivity or ecosystem respiration, and consequently resulted in no changes of net ecosystem productivity in a semi‐arid grassland despite whether temperature, precipitation and nitrogen were elevated or not. The observations could be primarily attributable to the offset of ecosystem carbon uptake by enhanced soil carbon release under CO₂ enrichment. Our findings indicate that arid and semi‐arid ecosystems may not be sensitive to CO₂ enrichment as previously expected and highlight the urgent need to incorporate this mechanism into most IPCC carbon‐cycle models for convincing projection of terrestrial carbon sink and its feedback to climate change.