Are chlorophyll concentrations and nitrogen across the vertical canopy profile affected by elevated CO2 in mature Quercus trees?

Key message: In mature Q. robur, chlorophyll varied with season and canopy height, whilst eCO2-driven changes were consistent with Marea, highlighting key factors for consideration when scaling photosynthetic processes and canopy N-use. Nitrogen-rich chlorophyll and carotenoid pigments are important in photosynthetic functioning. Photosynthetic pigments have been found to decrease with elevated CO2 (eCO2), but few such studies have been done in aged forest trees. This study aimed to assess the effects of eCO2 (150 μmol mol−1 above ambient) and canopy position on chlorophyll content in mature Quercus robur (Q. robur). Over 5000 in situ chlorophyll absorbance measurements, alongside laboratory chlorophyll extractions, were collected on canopy-dominant Q. robur in the 3rd and 4th season of CO2 fumigation of a free-air CO2 enrichment (FACE) study in central England. Mass-based chlorophyll concentration (Chlmass, mg g−1) was significantly higher in the lower canopy compared to upper canopy foliage (P < 0.05). In contrast, significantly higher chlorophyll content (Chlarea, mg m−2) was observed in the upper canopy. ECO2 did not affect Chlmass but Chlarea significantly increased, attributable to increased leaf mass per unit area (Marea, g m−2). We found no effect of eCO2 on mass-based or area-based nitrogen (Nmass, mg g−1 or Narea g m−2); however, Narea significantly increased with canopy height, again attributable to Marea. The parallel relationships between Marea, Narea and Chlarea suggest the allocation of N to light harvesting is maintained with eCO2 exposure as well as in the upper canopy, and that increased photosynthetic mass may help regulate the eCO2 variation. An understanding of changes in the light-harvesting machinery with eCO2 will be useful to assess canopy processes and, at larger scales, changes in biogeochemical cycles in future climate scenarios

A simple method for simulating drought effects on plants

Drought is expected to increase in frequency and severity in many regions in the future, so it is important to improve our understanding of how drought affects plant functional traits and ecological interactions. Imposing experimental water deficits is key to gaining this understanding, but has been hindered by logistic difficulties in maintaining consistently low water availability for plants. Here, we describe a simple method for applying soil water deficits to potted plants in glasshouse experiments. We modified an existing method (the “Snow and Tingey system”) in order to apply a gradual, moderate water deficit to 50 plant species of different life forms (grasses, vines, shrubs, trees). The method requires less maintenance and manual handling compared to other water deficit methods, so it can be used for extended periods of time and is relatively inexpensive to implement. With only a few modifications, it is possible to easily establish and maintain soil water deficits of differing intensity and duration, as well as to incorporate interacting stress factors. We tested this method by measuring physiological responses to an applied water deficit in a subset of 11 tree/shrub species with a wide range of drought tolerances and water-use strategies. For this subgroup of species, stomatal conductance was 2–17 times lower in droughted plants than controls, although only half of the species (5 out of 11) experienced midday leaf water potentials that exceeded their turgor loss (i.e., wilting) point. Leaf temperatures were up to 8°C higher in droughted plants than controls, indicating that droughted plants are at greater risk of thermal damage, relative to unstressed plants. The largest leaf temperature differences (between droughted and well-watered plants) were in species with high rates of water loss. Rapid osmotic adjustment was observed in leaves of five species when drought stress was combined with an experimental heatwave. These results highlight the potential value of further ecological and physiological experiments utilizing this simple water deficit method to study plant responses to drought stress

The influence of elevated CO2 and soil depth on rhizosphere activity and nutrient availability in a mature Eucalyptus woodland

Elevated carbon dioxide (eCO2) in the atmosphere increases forest biomass productivity but only where soil nutrients, particularly nitrogen (N) and phosphorus (P), are not limiting growth. eCO2, in turn, can impact rhizosphere nutrient availability. Our current understanding of nutrient cy cling under eCO2 is mainly derived from surface soil, leaving mechanisms of the impact of eCO2 on rhizosphere nutrient availability at deeper depths unexplored. To investigate the influence of eCO2 on nutrient availability in soil at depth, we studied various C, N, and P pools (extractable, microbial biomass, total soil C and N, and mineral-associated P) and nutrient cycling processes (enzyme activity and gross N mineralisation) associated with C, N, and P cycling in both bulk and rhizosphere soil at different depths at the Free Air CO2 enrichment facility in a native Australian mature Eucalyptus woodland (EucFACE) on a nutrient-poor soil. We found de creasing nutrient availability and gross N mineralisation with depth; however, this depth-associated decrease was reduced under eCO2, which we suggest is due to enhanced root influence. Increases in available PO3− 4 , adsorbed P, and the C : N and C : P ratio of enzyme activity with depth were observed. We conclude that the influences of roots and of eCO2 can affect available nutrient pools and processes well beyond the surface soil of a mature forest ecosystem. Our findings indicate a faster recycling of nutrients in the rhizosphere, rather than additional nutrients becoming available through soil organic matter (SOM) decomposition. If the plant growth response to eCO2 is reduced by the constraints of nutrient limitations, then the current results would call to question the potential for mature tree ecosystems to fix more C as biomass in response to eCO2. Future studies should address how accessible the available nutrients at depth are to deeply rooted plants and if fast recycling of nutrients is a meaningful contribution to biomass production and the accumulation of soil C in response to eCO2

Differentiation in stem and leaf traits among sympatric lianas, scandent shrubs and trees in a subalpine cold temperate forest

The scandent shrub plant form is a variant of liana that has upright and self-supporting stems when young but later becomes a climber. We aimed to explore the associations of stem and leaf traits among sympatric lianas, scandent shrubs and trees, and the effects of growth form and leaf habit on variation in stem or leaf traits. We measured 16 functional traits related to stem xylem anatomy, leaf morphology and nutrient stoichiometry in eight liana, eight scandent shrub and 21 tree species co-occurring in a subalpine cold temperate forest at an elevation of 2600–3200 m in Southwest China. Overall, lianas, scandent shrubs and trees were ordered along a fast-slow continuum of stem and leaf functional traits, with some traits overlapping. We found a consistent pattern of lianas > scandent shrubs > trees for hydraulically weighted vessel diameter, maximum vessel diameter and theoretical hydraulic conductivity. Vessel density and sapwood density showed a pattern of lianas = scandent shrubs < trees, and lianas < scandent shrubs = trees, respectively. Lianas had significantly higher specific leaf area and lower carbon concentration than co-occurring trees, with scandent shrubs showing intermediate values that overlapped with lianas and trees. The differentiation among lianas, scandent shrubs and trees was mainly explained by variation in stem traits. Additionally, deciduous lianas were positioned at the fast end of the trait spectrum, and evergreen trees at the slow end of the spectrum. Our results showed for the first time clear differentiation in stem and leaf traits among sympatric liana, scandent shrub and tree species in a subalpine cold temperate forest. This work will contribute to understanding the mechanisms responsible for variation in ecological strategies of different growth forms of woody plants

Extreme heat increases stomatal conductance and drought-induced mortality risk in vulnerable plant species

Tree mortality during global-change-type drought is usually attributed to xylem dysfunction, but as climate change increases the frequency of extreme heat events, it is necessary to better understand the interactive role of heat stress. We hypothesized that some drought-stressed plants paradoxically open stomata in heatwaves to prevent leaves from critically overheating. We experimentally imposed heat (&gt;40°C) and drought stress onto 20 broadleaf evergreen tree/shrub species in a glasshouse study. Most well-watered plants avoided lethal overheating, but drought exacerbated thermal damage during heatwaves. Thermal safety margins (TSM) quantifying the difference between leaf surface temperature and leaf critical temperature, where photosynthesis is disrupted, identified species vulnerability to heatwaves. Several mechanisms contributed to high heat tolerance and avoidance of damaging leaf temperatures—small leaf size, low leaf osmotic potential, high leaf mass per area (i.e., thick, dense leaves), high transpirational capacity, and access to water. Water-stressed plants had smaller TSM, greater crown dieback, and a fundamentally different stomatal heatwave response relative to well-watered plants. On average, well-watered plants closed stomata and decreased stomatal conductance (gs) during the heatwave, but droughted plants did not. Plant species with low gs, either due to isohydric stomatal behavior under water deficit or inherently low transpirational capacity, opened stomata and increased gs under high temperatures. The current paradigm maintains that stomata close before hydraulic thresholds are surpassed, but our results suggest that isohydric species may dramatically increase gs (over sixfold increases) even past their leaf turgor loss point. By actively increasing water loss at high temperatures, plants can be driven toward mortality thresholds more rapidly than has been previously recognized. The inclusion of TSM and responses to heat stress could improve our ability to predict the vulnerability of different tree species to future droughts

Stomatal responses of terrestrial plants to global change

Quantifying the stomatal responses of plants to global change factors is crucial for modeling terrestrial carbon and water cycles. Here we synthesize worldwide experimental data to show that stomatal conductance (gs) decreases with elevated carbon dioxide (CO2), warming, decreased precipitation, and tropospheric ozone pollution, but increases with increased precipitation and nitrogen (N) deposition. These responses vary with treatment magnitude, plant attributes (ambient gs, vegetation biomes, and plant functional types), and climate. All two-factor combinations (except warming + N deposition) significantly reduce gs, and their individual effects are commonly additive but tend to be antagonistic as the effect sizes increased. We further show that rising CO2 and warming would dominate the future change of plant gs across biomes. The results of our meta-analysis provide a foundation for understanding and predicting plant gs across biomes and guiding manipulative experiment designs in a real world where global change factors do not occur in isolation

Konvensyen Myprospec tumpu revolusi industri 4.0

Rising atmospheric concentrations of CO 2 (C a) can reduce stomatal conductance and transpiration rate in trees, but the magnitude of this effect varies considerably among experiments. The theory of optimal stomatal behaviour predicts that the ratio of photosynthesis to transpiration (instantaneous transpiration efficiency, ITE) should increase in proportion to C a. We hypothesized that plants regulate stomatal conductance optimally in response to rising C a. We tested this hypothesis with data from young Eucalyptus saligna Sm. trees grown in 12 climate-controlled whole-tree chambers for 2 years at ambient and elevated C a. Elevated C a was ambient + 240 ppm, 60% higher than ambient C a. Leaf-scale gas exchange was measured throughout the second year of the study and leaf-scale ITE increased by 60% under elevated C a, as predicted. Values of leaf-scale ITE depended strongly on vapour pressure deficit (D) in both CO 2 treatments. Whole-canopy CO 2 and H 2O fluxes were also monitored continuously for each chamber throughout the second year. There were small differences in D between C a treatments, which had important effects on values of canopy-scale ITE. However, when C a treatments were compared at the same D, canopy-scale ITE was consistently increased by 60%, again as predicted. Importantly, leaf and canopy-scale ITE were not significantly different, indicating that ITE was not scale-dependent. Observed changes in transpiration rate could be explained on the basis that ITE increased in proportion to C a. The effect of elevated C a on photosynthesis increased with rising D. At high D, C a had a large effect on photosynthesis and a small effect on transpiration rate. At low D, in contrast, there was a small effect of C a on photosynthesis, but a much larger effect on transpiration rate. If shown to be a general response, the proportionality of ITE with C a will allow us to predict the effects of C a on transpiration rate

Probing the inner sanctum of leaf phosphorus : measuring the fractions of leaf P

Background: In spite of the importance of phosphorus (P) to plant physiological function and growth, relatively few studies have quantified foliar P fractions in native plants in natural environments. Understanding how these P fractions vary with P availability, soil type and parent material should provide information on the importance of P storage versus its partitioning to cell ultrastructure versus active biochemical compounds. In the latest study evaluating foliar P fractions, McQuillan et al. (2020), this issue, have enlisted a novel technique to estimate these foliar P fractions for major groups of functional and structural compounds in native species of different taxa across sites west of the Great Dividing Range of Australia. Scope: Combined with recent studies of diverse tropical species, there is a conservative amount of lipid-membrane P and nucleic acid P across a threefold range of leaf P concentrations, from very low leaf P concentration to what could arguably be considered moderately low leaf P concentrations (0.3 to 1.0 mg g−1 leaf P concentration). Conclusions: The findings provide insight into how overall leaf P concentrations are partitioned, including that P investment in structural components of the leaf like membrane phospholipids is remarkably conservatively regulated. Further insights await a quantification of organelle-specific P fractions on well-preserved samples, so importance of the storage versus biochemical functions of orthophosphate can be elucidated. These insights will be important for incorporating functional components of P and P biogeochemistry into models of ecosystem function, for understanding how P may regulate global change responses

Leaf inclination angle and foliage clumping in an evergreen broadleaf Eucalyptus forest under elevated atmospheric CO2

How leaves are presented affects interaction of atmospheric CO2, energy (light), and plant physiology. Plant productivity is primarily determined by the amount of leaf area, leaf orientation and distribution in space. Not much attention has been paid to possible changes in leaf orientation and distribution with elevated CO2, but its effect on plant growth could alter the proportions of sunlit and shaded leaf areas and feedback on carbohydrate available for further growth. We report on first measurements of leaf inclination angle distribution and foliage clumping in a native evergreen Eucalyptus woodland in ambient CO2 and under +150 ppm elevated CO2. We found that a spherical leaf angle distribution was not an appropriate supposition for present species (Eucalyptus tereticornis Sm.) at this site. Our measurements of leaf inclination angles from imagery indicated an erectophile, highly skewed unimodal leaf inclination angle distribution function. We conclude that despite the measured steeper angles under elevated CO2 concentrations, the leaf angle change is not significant and falls within the expected natural variability and uncertainties connected with the measurement method. The lack of a clear response of leaf orientation and foliage clumping to elevated CO2 concentration indicates that the previously produced datasets of leaf inclination angles and foliage clumping maps with Earth observation data may be suitable while modelling carbon and water cycles under climate change

Is phosphorus limiting in a mature Eucalyptus woodland? : phosphorus fertilisation stimulates stem growth

Aims: Few direct tests of phosphorus (P) limitation on highly-weathered soils have been conducted, especially in mature, native Eucalyptus stands. We tested whether growth in a mature >80-year old stand of Eucalyptus tereticornis in Cumberland Plain Woodland was limited by P, and whether this P-limitation affected leaf photosynthetic capacity. Methods: P was added to trees at the native woodland site at 50 kg ha-1 year-1 in each of 3 years, and stem and leaf responses were measured. Results: Leaf P concentrations before fertilisation were 2], because photosynthesis in elevated [CO2] may become further constrained by required phosphate pools within the photosynthetic apparatus