Limits to photosynthesis: seasonal shifts in supply and demand for CO2 in Scots pine

Boreal forests undergo a strong seasonal photosynthetic cycle; however, the underlying processes remain incompletely characterized. Here, we present a novel analysis of the seasonal diffusional and biochemical limits to photosynthesis (A(net)) relative to temperature and light limitations in high-latitude mature Pinus sylvestris, including a high-resolution analysis of the seasonality of mesophyll conductance (g(m)) and its effect on the estimation of carboxylation capacity (VCmax). We used a custom-built gas-exchange system coupled to a carbon isotope analyser to obtain continuous measurements for the estimation of the relevant shoot gas-exchange parameters and quantified the biochemical and diffusional controls alongside the environmental controls over A(net). The seasonality of A(net) was strongly dependent on VCmax and the diffusional limitations. Stomatal limitation was low in spring and autumn but increased to 31% in June. By contrast, mesophyll limitation was nearly constant (19%). We found that VCmax limited A(net) in the spring, whereas daily temperatures and the gradual reduction of light availability limited A(net) in the autumn, despite relatively high VCmax. We describe for the first time the role of mesophyll conductance in connection with seasonal trends in net photosynthesis of P. sylvestris, revealing a strong coordination between g(m) and A(net), but not between g(m) and stomatal conductance

Accumulation of antimony and lead in leaves and needles of trees: The role of traffic emissions

Antimony (Sb) is a toxic metalloid, which has been increasingly used in the brake lining of ve-hicles, and increased concentrations have been found in soils near abundant traffic. However, since very few investigations of Sb accumulation in urban vegetation have been undertaken there exists a knowledge gap. We studied the concentrations of Sb in leaves and needles of trees in the Gothenburg City area, Sweden. In addition, lead (Pb), also associated with traffic, was investi-gated. Sb and Pb concentrations of Quercus palustris leaves at seven sites with contrasting traffic intensity varied substantially, correlated with the traffic-related PAH (polycyclic aromatic hy-drocarbon) air pollution at the sites and increased during the growing season. Sb but not Pb concentrations were significantly higher in needles of Picea abies and Pinus sylvestris near major roads compared to sites at larger distances. In Pinus nigra needles at two urban streets both Sb and Pb were higher compared to an urban nature park environment, emphasising the role of traffic emissions for these elements. A continued accumulation of Sb and Pb in three years old needles of Pinus nigra, two years old needles of Pinus sylvestris and eleven years old needles of Picea abies was observed. Our data suggest a pronounced link between traffic pollution and Sb accumulation in leaves and needles, where the particles carrying Sb seem not to be transported very far from the source. We also conclude that there exists a strong potential for Sb and Pb bioaccumulation over time in leaves and needles. Implications of these findings are that increased concentrations of toxic Sb and Pb are likely to prevail in environments with high traffic intensity and that Sb can enter the ecological food chain by accumulation in leaves and needles, which is important for the biogeochemical cycling

Components explain, but do eddy fluxes constrain? Carbon budget of a nitrogen-fertilized boreal Scots pine forest

Nitrogen (N) fertilization increases biomass and soil organic carbon (SOC) accumulation in boreal pine forests, but the underlying mechanisms remain uncertain. At two Scots pine sites, one undergoing annual N fertilization and the other a reference, we sought to explain these responses.We measured component fluxes, including biomass production, SOC accumulation, and respiration, and summed them into carbon budgets. We compared the resulting summations to ecosystem fluxes measured by eddy covariance.N fertilization increased most component fluxes (P −2 yr−1. Stemwood production increases were ascribed to this partitioning shift, gross primary production (GPP), and carbon-use efficiency, in that order. The methods agreed in their estimates of GPP in both stands (P > 0.05), but the components detected an increase in net ecosystem production (NEP) (190 (54) g C m−2 yr−1; P −2 yr−1; ns).The pairing of plots, the simplicity of the sites, and the strength of response provide a compelling description of N effects on the C budget. However, the disagreement between methods calls for further paired tests of N fertilization effects in simple forest ecosystems

Limited vertical CO2 transport in stems of mature boreal Pinus sylvestris trees

Several studies have suggested that CO2 transport in the transpiration stream can considerably bias estimates of root and stem respiration in ring-porous and diffuse-porous tree species. Whether this also happens in species with tracheid xylem anatomy and lower sap flow rates, such as conifers, is currently unclear. We infused C-13-labelled solution into the xylem near the base of two 90-year-old Pinus sylvestris L. trees. A custom-built gas exchange system and an online isotopic analyser were used to sample the CO2 efflux and its isotopic composition continuously from four positions along the bole and one upper canopy shoot in each tree. Phloem and needle tissue C-13 enrichment was also evaluated at these positions. Most of the C-13 label was lost by diffusion within a few metres of the infusion point indicating rapid CO2 loss during vertical xylem transport. No C-13 enrichment was detected in the upper bole needle tissues. Furthermore, mass balance calculations showed that c. 97% of the locally respired CO2 diffused radially to the atmosphere. Our results support the notion that xylem CO2 transport is of limited magnitude in conifers. This implies that the concerns that stem transport of CO2 derived from root respiration biases chamber-based estimates of forest carbon cycling may be unwarranted for mature conifer stands.Peer reviewe

Polycyclic aromatic hydrocarbon (PAH) accumulation in Quercus palustris and Pinus nigra in the urban landscape of Gothenburg, Sweden

Trees have the potential to improve urban air quality as leaves and needles capture air pollutants from the air, but further empirical data has been requested to quantify these effects. We measured the concentration of 32 poly-cyclic aromatic hydrocarbons (PAHs) in leaves of pin oak (Quercus palustris) and needles of black pine (Pinus nigra) in the City of Gothenburg, Sweden, during the summer of 2018. Oak leaves were collected twice (June, Sep-tember), while one-year-old (C + 1) and three-year-old (C + 3) pine needles were sampled in June to study the temporal development of leaf/needle PAH concentrations. Specific leaf area (SLA) was estimated, which permit-ted calculation of leaf/needle area-based PAH content that were compared with the mass-based concentration. In addition, the air concentration of PAHs and NO2 was measured using passive samplers. There was a strong correlation between air concentrations of PAH and NO2, indicating that the pollutants to a large degree originate from the same sources. In the oak leaves there was a significant decrease in low molecular mass PAHs (L-PAH, mainly gaseous) between June and September, but a significant increase in high molecular mass PAHs (H-PAH, mainly particle-bound). There was a strong correlation between L-PAH concentration in leaves and in air indicating an influence of equilibrium processes between ambient air and leaf. In the pine needles, there was a significant increase of both L-PAH and H-PAH in three-year-old needles compared to one-year-old needles. Pine was superior to oak in accumulating PAHs from the air, especially for L-PAHs when comparing area-based content. However, H-PAH concentrations were higher in oak leaves compared to pine needles on a leaf mass basis, emphasizing the importance of how concentrations are expressed. The results from this study can contribute to the development of urban planning strategies regarding the effect of vegetation on air quality. (C) 2021 The Authors. Published by Elsevier B.V

Variation in Polycyclic Aromatic Compound (PAC) Concentrations in a Norway Spruce Stand Close to a Major Traffic Route-Influence of Distance and Season

Air concentrations of polycyclic aromatic compounds (PACs) were measured in a Norway spruce (Picea abies) stand at four different distances from a major traffic route near Gothenburg, Sweden, during summer and winter 2019, using passive samplers. In total, the concentrations of 32 polycyclic aromatic hydrocarbons (PAHs) and six dibenzothiophenes (DBTs) and NO2 were analysed. PAC air concentrations were on average 2.5 times higher during the winter compared to the summer. All investigated PAH categories and compounds as well as dibenzothiophene declined linearly with the logarithmic distance from the traffic route. This indicates that the length of pollutant pathways through vegetation is a valid proxy to assess health risks of traffic-related PAC pollutants. Concentrations of heavier PAHs declined faster (concentration at site furthest away from the road was 6% and 33% of concentration at site closest to road, for summer and winter respectively) with increasing distance from traffic compared to lighter PAHs (63% and 68% for summer and winter respectively). As a result, the mix of PAHs changed with the depth into the forest stand (i.e. distance from the road) with potential effects on the toxicity as well as on diagnostic ratios used for source attribution. This is likely caused by different deposition rates among the PAH compounds to the vegetation surfaces, although the exact mechanism needs to be further investigated. This study provides new information that can improve air quality risk assessment and limit the exposure of the population to toxic air pollutants such as PACs

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (Na) is equal to the light gradient. However, it is widely observed that Na declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (ma). Using a simple model of the carbon-nitrogen (C-N) balance of trees with a steady-state canopy, we implement this constraint within the framework of the MAXX optimization hypothesis that maximizes net canopy C export. Virtually all canopy traits predicted by MAXX (leaf N gradient, leaf N concentration, leaf photosynthetic capacity, canopy N content, leaf-area index) are in close agreement with the values observed in a mature stand of Norway spruce trees (Picea abies L. Karst.). An alternative upper-bound constraint on leaf photosynthetic capacity (Asat) does not reproduce the canopy traits of this stand. MAXX subject to a lower bound on ma is also qualitatively consistent with co-variations in leaf N gradient, ma and Asat observed across a range of temperate and tropical tree species. Our study highlights the key role of constraints in optimization models of plant function

Variation in Polycyclic Aromatic Compound (PAC) Concentrations in a Norway Spruce Stand Close to a Major Traffic Route-Influence of Distance and Season

Air concentrations of polycyclic aromatic compounds (PACs) were measured in a Norway spruce (Picea abies) stand at four different distances from a major traffic route near Gothenburg, Sweden, during summer and winter 2019, using passive samplers. In total, the concentrations of 32 polycyclic aromatic hydrocarbons (PAHs) and six dibenzothiophenes (DBTs) and NO2 were analysed. PAC air concentrations were on average 2.5 times higher during the winter compared to the summer. All investigated PAH categories and compounds as well as dibenzothiophene declined linearly with the logarithmic distance from the traffic route. This indicates that the length of pollutant pathways through vegetation is a valid proxy to assess health risks of traffic-related PAC pollutants. Concentrations of heavier PAHs declined faster (concentration at site furthest away from the road was 6% and 33% of concentration at site closest to road, for summer and winter respectively) with increasing distance from traffic compared to lighter PAHs (63% and 68% for summer and winter respectively). As a result, the mix of PAHs changed with the depth into the forest stand (i.e. distance from the road) with potential effects on the toxicity as well as on diagnostic ratios used for source attribution. This is likely caused by different deposition rates among the PAH compounds to the vegetation surfaces, although the exact mechanism needs to be further investigated. This study provides new information that can improve air quality risk assessment and limit the exposure of the population to toxic air pollutants such as PACs

Acclimation of light and dark respiration to experimental and seasonal warming are mediated by changes in leaf nitrogen in Eucalyptus globulus

Quantifying the adjustments of leaf respiration in response to seasonal temperature variation and climate warming is crucial because carbon loss from vegetation is a large but uncertain part of the global carbon cycle. We grew fast-growing Eucalyptus globulus Labill. trees exposed to +3 °C warming and elevated CO2 in 10-m tall whole-tree chambers and measured the temperature responses of leaf mitochondrial respiration, both in light (RLight) and in darkness (RDark), over a 20–40 °C temperature range and during two different seasons. RLight was assessed using the Laisk method. Respiration rates measured at a standard temperature (25 °C – R25) were higher in warm-grown trees and in the warm season, related to higher total leaf nitrogen (N) investment with higher temperatures (both experimental and seasonal), indicating that leaf N concentrations modulated the respiratory capacity to changes in temperature. Once differences in leaf N were accounted for, there were no differences in R25 but the Q10 (i.e., short-term temperature sensitivity) was higher in late summer compared with early spring. The variation in RLight between experimental treatments and seasons was positively correlated with carboxylation capacity and photorespiration. RLight was less responsive to short-term changes in temperature than RDark, as shown by a lower Q10 in RLight compared with RDark. The overall light inhibition of R was ∼40%. Our results highlight the dynamic nature of leaf respiration to temperature variation and that the responses of RLight do not simply mirror those of RDark. Therefore, it is important not to assume that RLight is the same as RDark in ecosystem models, as doing so may lead to large errors in predicting plant CO2 release and productivity

Spring photosynthetic recovery of boreal Norway spruce under conditions of elevated [CO2] and air temperature

Accumulated carbon uptake, apparent quantum yield (AQY) and light-saturated net CO2 assimilation (A(sat)) were used to assess the responses of photosynthesis to environmental conditions during spring for three consecutive years. Whole-tree chambers were used to expose 40-year-old field-grown Norway spruce trees in northern Sweden to an elevated atmospheric CO2 concentration, [CO2], of 700 mu mol CO2 mol(-1) (C-E) and an air temperature (T) between 2.8 and 5.6 degrees C above ambient T (T-E), during summer and winter. Net shoot CO2 exchange (A(net)) was measured continuously on 1-year-old shoots and was used to calculate the accumulated carbon uptake and daily A(sat) and AQY. The accumulated carbon uptake, from 1 March to 30 June, was stimulated by 33, 44 and 61% when trees were exposed to C-E, T-E, and C-E and T-E combined, respectively. Air temperature strongly influenced the timing and extent of photosynthetic recovery expressed as AQY and A(sat) during the spring. Under elevated T (T-E), the recovery of AQY and A(sat) commenced similar to 10 days earlier and the activity of these parameters was significantly higher throughout the recovery period. In the absence of frost events, the photosynthetic recovery period was less than a week. However, frost events during spring slowed recovery so that full recovery could take up to 60 days to complete. Elevated [CO2] stimulated AQY and A(sat) on average by similar to 10 and similar to 50%, respectively, throughout the recovery period, but had minimal or no effect on the onset and length of the photosynthetic recovery period during the spring. However, AQY, A(sat) and A(net) all recovered at significantly higher T (average +2.2 degrees C) in T-E than in T-A, possibly caused by acclimation or by shorter days and lower light levels during the early part of the recovery in T-E compared with T-A. The results suggest that predicted future climate changes will cause prominent stimulation of photosynthetic CO2 uptake in boreal Norway spruce forest during spring, mainly caused by elevated T, but also elevated [CO2]. However, the effects of elevated T may not be linearly extrapolated to future warmer climates