Johan Uddling

Assessing foliar chlorophyll contents with the SPAD-502 chlorophyll meter: a calibration test with thirteen tree species of tropical rainforest in French Guiana

Abstract• Chlorophyll meters such as the SPAD-502 offer a simple, inexpensive and rapid method to estimate foliar chlorophyll content. However, values provided by SPAD-502 are unitless and require empirical calibrations between SPAD units and extracted chlorophyll values.• Leaves of 13 tree species from the tropical rain forest in French Guiana were sampled to select the most appropriate calibration model among the often-used linear, polynomial and exponential models, in addition to a novel homographic model that has a natural asymptote.• The homographic model best accurately predicted total chlorophyll content (μg cm−2) from SPAD units (R2 = 0.89). Interspecific differences in the homographic model parameters explain less than 7% of the variation in chlorophyll content in our data set.• The utility of the general homographic model for a variety of research and management applications clearly outweighs the slight loss of model accuracy due to the abandon of the species’ effect.

Leaf and canopy conductance in aspen and aspen-birch forests under free-air enrichment of carbon dioxide and ozone

Increasing concentrations of atmospheric carbon dioxide (CO2) and tropospheric ozone (O3) have the potential to affect tree physiology and structure, and hence forest feedbacks on climate. Here, we investigated how elevated concentrations of CO2 (+45%) and O3 (+35%), alone and in combination, affected conductance for mass transfer at the leaf and canopy levels in pure aspen (Populus tremuloides Michx.) and in mixed aspen and birch (Betula papyrifera Marsh.) forests in the free-air CO2-O3 enrichment experiment near Rhinelander, Wisconsin (Aspen FACE). The study was conducted during two growing seasons, when steady-state leaf area index (L) had been reached after > 6 years of exposure to CO2- and O3-enrichment treatments. Canopy conductance (g(c)) was estimated from stand sap flux, while leaf-level conductance of sun leaves in the upper canopy was derived by three different and independent methods: sap flux and L in combination with vertical canopy modelling, leaf 13C discrimination methodology in combination with photosynthesis modelling and leaf-level gas exchange. Regardless of the method used, the mean values of leaf-level conductance were higher in trees growing under elevated CO2 and/or O3 than in trees growing in control plots, causing a CO2 x O3 interaction that was statistically significant (P < or = 0.10) for sap flux- and (for birch) 13C-derived leaf conductance. Canopy conductance was significantly increased by elevated CO2 but not significantly affected by elevated O3. Investigation of a short-term gap in CO2 enrichment demonstrated a +10% effect of transient exposure of elevated CO2-grown trees to ambient CO2 on g(c). All treatment effects were similar in pure aspen and mixed aspen-birch communities. These results demonstrate that short-term primary stomatal closure responses to elevated CO2 and O3 were completely offset by long-term cumulative effects of these trace gases on tree and stand structure in determining canopy- and leaf-level conductance in pure aspen and mixed aspen-birch forests. Our results, together with the findings from other long-term FACE experiments with trees, suggest that model assumptions of large reductions in stomatal conductance under rising atmospheric CO2 are very uncertain for forests.

Constraints to nitrogen acquisition of terrestrial plants under elevated CO2.

A key part of the uncertainty in terrestrial feedbacks on climate change is related to how and to what extent nitrogen (N) availability constrains the stimulation of terrestrial productivity by elevated CO2 (eCO2 ), and whether or not this constraint will become stronger over time. We explored the ecosystem-scale relationship between responses of plant productivity and N acquisition to eCO2 in free-air CO2 enrichment (FACE) experiments in grassland, cropland and forest ecosystems and found that: (i) in all three ecosystem types, this relationship was positive, linear and strong (r(2) = 0.68), but exhibited a negative intercept such that plant N acquisition was decreased by 10% when eCO2 caused neutral or modest changes in productivity. As the ecosystems were markedly N limited, plants with minimal productivity responses to eCO2 likely acquired less N than ambient CO2 -grown counterparts because access was decreased, and not because demand was lower. (ii) Plant N concentration was lower under eCO2 , and this decrease was independent of the presence or magnitude of eCO2 -induced productivity enhancement, refuting the long-held hypothesis that this effect results from growth dilution. (iii) Effects of eCO2 on productivity and N acquisition did not diminish over time, while the typical eCO2 -induced decrease in plant N concentration did. Our results suggest that, at the decennial timescale covered by FACE studies, N limitation of eCO2 -induced terrestrial productivity enhancement is associated with negative effects of eCO2 on plant N acquisition rather than with growth dilution of plant N or processes leading to progressive N limitation.

Stomatal and non-stomatal fluxes of ozone to a northern mixed hardwood forest

Measurements of ozone, sensible heat, and latent heat fluxes and plant physiological parameters were made at a northern mixed hardwood forest located at the University of Michigan Biological Station in northern Michigan from June 27 to September 28, 2002. These measurements were used to calculate total ozone flux and partitioning between stomatal and non-stomatal sinks. Total ozone flux varied diurnally with maximum values reaching 100 μmol m-2 h-1 at midday and minimums at or near zero at night. Mean daytime canopy conductance was 0.5 mol m-2 s-1. During daytime, non-stomatal ozone conductance accounted for as much as 66% of canopy conductance, with the non-stomatal sink representing 63% of the ozone flux. Stomatal conductance showed expected patterns of behaviour with respect to photosynthetic photon flux density (PPFD) and vapour pressure defecit (VPD). Non-stomatal conductance for ozone increased monotonically with increasing PPFD, increased with temperature (T) before falling off again at high T, and behaved similarly for VPD. Day-time non-stomatal ozone sinks are large and vary with time and environmental drivers, particularly PPFD and T. This information is crucial to deriving mechanistic models that can simulate ozone uptake by different vegetation types

Photosynthetic temperature responses of tree species in Rwanda: evidence of pronounced negative effects of high temperature in montane rainforest climax species

The sensitivity of photosynthetic metabolism to temperature has been identified as a key uncertainty for projecting the magnitude of the terrestrial feedback on future climate change. While temperature responses of photosynthetic capacities have been comparatively well investigated in temperate species, the responses of tropical tree species remain unexplored. We compared the responses of seedlings of native cold-adapted tropical montane rainforest tree species with those of exotic warm-adapted plantation species, all growing in an intermediate temperature common garden in Rwanda. Leaf gas exchange responses to carbon dioxide (CO2 ) at different temperatures (20-40°C) were used to assess the temperature responses of biochemical photosynthetic capacities. Analyses revealed a lower optimum temperature for photosynthetic electron transport rates than for Rubisco carboxylation rates, along with lower electron transport optima in the native cold-adapted than in the exotic warm-adapted species. The photosynthetic optimum temperatures were generally exceeded by daytime peak leaf temperatures, in particular in the native montane rainforest climax species. This study thus provides evidence of pronounced negative effects of high temperature in tropical trees and indicates high susceptibility of montane rainforest climax species to future global warming.

Ozone risk for vegetation in the future climate of Europe based on stomatal ozone uptake calculations

The negative impacts of surface ozone (O3) on vegetation are determined by external exposure, leaf gas exchange and plant antioxidant defence capacity, all dependent on climate and CO2 concentrations. In this study the influence of climate change on simulated stomatal O3 uptake of a generic crop and a generic deciduous tree at ten European sites was investigated, using the LRTAP Mapping Manual stomatal flux model. O3 concentrations are calculated by a chemistry transport model (MATCH) for three 30-yr time-windows (1961–1990, 2021–2050, 2071–2100), with constant precursor emissions and meteorology from a regional climate model (RCA3). Despite substantially increased modelled future O3 concentrations in central and southern Europe, the flux-based risk for O3 damage to vegetation is predicted to remain unchanged or decrease at most sites, mainly as a result of projected reductions in stomatal conductance under rising CO2 concentrations. Drier conditions in southern Europe are also important for this result. At northern latitudes, the current parameterisation of the stomatal conductance model suggest O3 uptake to be mainly limited by temperature. This study demonstrates the importance of accounting for the influences by climate and CO2 on stomatal O3 uptake, and of developing their representation in models, for risk assessment involving climate change.

Stomatal uptake of O3 in aspen and aspen-birch forests under free-air CO2 and O3 enrichment

Rising atmospheric carbon dioxide (CO2) may alleviate the toxicological impacts of concurrently rising tropospheric ozone (O3) during the present century if higher CO2 is accompanied by lower stomatal conductance (gs), as assumed by many models. We investigated how elevated concentrations of CO2 and O3, alone and in combination, affected the accumulated stomatal flux of O3 (AFst) by canopies and sun leaves in closed aspen and aspen-birch forests in the free-air CO2-O3 enrichment experiment near Rhinelander, Wisconsin. Stomatal conductance for O3 was derived from sap flux data and AFst was estimated either neglecting or accounting for the potential influence of non-stomatal leaf surface O3 deposition. Leaf-level AFst (AFst(l)) was not reduced by elevated CO2. Instead, there was a significant CO2 x O(3) interaction on AFst(l), as a consequence of lower values of gs in control plots and the combination treatment than in the two single-gas treatments. In addition, aspen leaves had higher AFst(l) than birch leaves, and estimates of AFst(l) were not very sensitive to non-stomatal leaf surface O3 deposition. Our results suggest that model projections of large CO2-induced reductions in gs alleviating the adverse effect of rising tropospheric O3 may not be reasonable for northern hardwood forests.

Exposure to moderate concentrations of tropospheric ozone impairs tree stomatal response to carbon dioxide.

With rising concentrations of both atmospheric carbon dioxide (CO(2)) and tropospheric ozone (O(3)), it is important to better understand the interacting effects of these two trace gases on plant physiology affecting land-atmosphere gas exchange. We investigated the effect of growth under elevated CO(2) and O(3), singly and in combination, on the primary short-term stomatal response to CO(2) concentration in paper birch at the Aspen FACE experiment. Leaves from trees grown in elevated CO(2) and/or O(3) exhibited weaker short-term responses of stomatal conductance to both an increase and a decrease in CO(2) concentration from current ambient level. The impairement of the stomatal CO(2) response by O(3) most likely developed progressively over the growing season as assessed by sap flux measurements. Our results suggest that expectations of plant water-savings and reduced stomatal air pollution uptake under rising atmospheric CO(2) may not hold for northern hardwood forests under concurrently rising tropospheric O(3).

Mycorrhiza Symbiosis Increases the Surface for Sunlight Capture in Medicago truncatula for Better Photosynthetic Production

Arbuscular mycorrhizal (AM) fungi play a prominent role in plant nutrition by supplying mineral nutrients, particularly inorganic phosphate (Pi), and also constitute an important carbon sink. AM stimulates plant growth and development, but the underlying mechanisms are not well understood. In this study, Medicago truncatula plants were grown with Rhizophagus irregularis BEG141 inoculum (AM), mock inoculum (control) or with Pi fertilization. We hypothesized that AM stimulates plant growth through either modifications of leaf anatomy or photosynthetic activity per leaf area. We investigated whether these effects are shared with Pi fertilization, and also assessed the relationship between levels of AM colonization and these effects. We found that increased Pi supply by either mycorrhization or fertilization led to improved shoot growth associated with increased nitrogen uptake and carbon assimilation. Both mycorrhized and Pi-fertilized plants had more and longer branches with larger and thicker leaves than the control plants, resulting in an increased photosynthetically active area. AM-specific effects were earlier appearance of the first growth axes and increased number of chloroplasts per cell section, since they were not induced by Pi fertilization. Photosynthetic activity per leaf area remained the same regardless of type of treatment. In conclusion, the increase in growth of mycorrhized and Pi-fertilized Medicago truncatula plants is linked to an increase in the surface for sunlight capture, hence increasing their photosynthetic production, rather than to an increase in the photosynthetic activity per leaf area.