Controls on the emission of plant volatiles through stomata: Differential sensitivity of emission rates to stomatal closure explained

[1] Volatile (VOC) flux from leaves may be expressed as G(S)DeltaP, where G(S) is stomatal conductance to specific compound and DeltaP partial pressure gradient between the atmosphere and substomatal cavities. It has been suggested that decreases in G(S) are balanced by increases in DeltaP such that stomata cannot control VOC emission. Yet, responses of emission rates of various volatiles to experimental manipulations of stomatal aperture are contrasting. To explain these controversies, a dynamic emission model was developed considering VOC distribution between gas and liquid phases using Henry's law constant (H, Pa m(3) mol(-1)). Our analysis demonstrates that highly volatile compounds such as isoprene and monoterpenes with H values on the order of 10(3) have gas and liquid pool half-times of a few seconds, and thus cannot be controlled by stomata. More soluble compounds such as alcohols and carboxylic acids with H values of 10(-2)-10(1) are controlled by stomata with the degree of stomatal sensitivity varying with H. Inability of compounds with high solubility to support a high partial pressure, and thus to balance DeltaP in response to a decrease in G(S) is the primary explanation for different stomatal sensitivities. For compounds with low H, the analysis predicts bursts of emission after stomatal opening that accord with experimental observations, but that cannot be currently explained. Large within-leaf VOC pool sizes in compounds with low H also increase the system inertia to environmental fluctuations. In conclusion, dynamic models are necessary to simulate diurnal variability of the emissions of compounds that preferably partition to aqueous phase

Nutrient limitation reduces land carbon uptake in simulations with a model of combined carbon, nitrogen and phosphorus cycling

Terrestrial carbon (C) cycle models applied for climate projections simulate a strong increase in net primary productivity (NPP) due to elevated atmospheric CO<sub>2</sub> concentration during the 21st century. These models usually neglect the limited availability of nitrogen (N) and phosphorus (P), nutrients that commonly limit plant growth and soil carbon turnover. To investigate how the projected C sequestration is altered when stoichiometric constraints on C cycling are considered, we incorporated a P cycle into the land surface model JSBACH (Jena Scheme for Biosphere–Atmosphere Coupling in Hamburg), which already includes representations of coupled C and N cycles. <br><br> The model reveals a distinct geographic pattern of P and N limitation. Under the SRES (Special Report on Emissions Scenarios) A1B scenario, the accumulated land C uptake between 1860 and 2100 is 13% (particularly at high latitudes) and 16% (particularly at low latitudes) lower in simulations with N and P cycling, respectively, than in simulations without nutrient cycles. The combined effect of both nutrients reduces land C uptake by 25% compared to simulations without N or P cycling. Nutrient limitation in general may be biased by the model simplicity, but the ranking of limitations is robust against the parameterization and the inflexibility of stoichiometry. After 2100, increased temperature and high CO<sub>2</sub> concentration cause a shift from N to P limitation at high latitudes, while nutrient limitation in the tropics declines. The increase in P limitation at high-latitudes is induced by a strong increase in NPP and the low P sorption capacity of soils, while a decline in tropical NPP due to high autotrophic respiration rates alleviates N and P limitations. The quantification of P limitation remains challenging. The poorly constrained processes of soil P sorption and biochemical mineralization are identified as the main uncertainties in the strength of P limitation. Even so, our findings indicate that global land C uptake in the 21st century is likely overestimated in models that neglect P and N limitations. In the long term, insufficient P availability might become an important constraint on C cycling at high latitudes. Accordingly, we argue that the P cycle must be included in global models used for C cycle projections

Does the law of diminishing returns in leaf scaling apply to vines? - Evidence from 12 species of climbing plants

Shapes, sizes and biomass investment per unit area (LMA) of vine leaves are characterized by high diversity that results in variation in leaf arrangement, light harvesting efficiency and photosynthetic activity. There exists a scaling relationship between leaf dry mass and surface area for many broad-leaved plants, and most estimates of the scaling exponent are greater than unity, implying that they follow the “law of diminishing returns”, i.e. that larger leaves require progressively greater investments of dry mass and accordingly have a greater LMA. Previous studies have primarily focused on trees and crops and there are few data available for vines. Yet, as vines have lower support investments in stems than self-supporting plants, they can have larger biomass investments in support within the leaves and stronger rise of biomass costs with increasing leaf area. In this study, we chose twelve species of vines (five woody vines and seven herbaceous vines) to investigate the following scientific questions: (i) whether there are significant differences in LMA between woody and herbaceous vines, (ii) whether leaf dry mass and surface area scaling relationships show evidence of diminishing returns in vines.We observed that LMA values of woody vines were significantly higher than those of the herbaceous vines. Leaf dry mass vs. surface area scaling relationship followed the law of diminishing returns in all 12 studied vine species. The existence of diminishing returns indicates that there is a trade-off between leaf surface area expansion and the energy investment for vines to support leaf physical structures

Estimations of isoprenoid emission capacity from enclosure studies: measurements, data processing, quality and standardized measurement protocols

The capacity for volatile isoprenoid production under standardized environmental conditions at a certain time (ES, the emission factor) is a key characteristic in constructing isoprenoid emission inventories. However, there is large variation in published ES estimates for any given species partly driven by dynamic modifications in ES due to acclimation and stress responses. Here we review additional sources of variation in ES estimates that are due to measurement and analytical techniques and calculation and averaging procedures, and demonstrate that estimations of ES critically depend on applied experimental protocols and on data processing and reporting. A great variety of experimental setups has been used in the past, contributing to study-to-study variations in ES estimates. We suggest that past experimental data should be distributed into broad quality classes depending on whether the data can or cannot be considered quantitative based on rigorous experimental standards. Apart from analytical issues, the accuracy of ES values is strongly driven by extrapolation and integration errors introduced during data processing. Additional sources of error, especially in meta-database construction, can further arise from inconsistent use of units and expression bases of ES. We propose a standardized experimental protocol for BVOC estimations and highlight basic meta-information that we strongly recommend to report with any ES measurement. We conclude that standardization of experimental and calculation protocols and critical examination of past reports is essential for development of accurate emission factor databases.JRC.H.7-Climate Risk Managemen

Does the leaf economic spectrum hold within plant functional types? A Bayesian multivariate trait meta-analysis

The leaf economic spectrum is a widely studied axis of plant trait variability that defines a trade-off between leaf longevity and productivity. While this has been investigated at the global scale, where it is robust, and at local scales, where deviations from it are common, it has received less attention at the intermediate scale of plant functional types (PFTs). We investigated whether global leaf economic relationships are also present within the scale of plant functional types (PFTs) commonly used by Earth System models, and the extent to which this global-PFT hierarchy can be used to constrain trait estimates. We developed a hierarchical multivariate Bayesian model that assumes separate means and covariance structures within and across PFTs and fit this model to seven leaf traits from the TRY database related to leaf longevity, morphology, biochemistry, and photosynthetic metabolism. Although patterns of trait covariation were generally consistent with the leaf economic spectrum, we found three approximate tiers to this consistency. Relationships among morphological and biochemical traits (specific leaf area [SLA], N, P) were the most robust within and across PFTs, suggesting that covariation in these traits is driven by universal leaf construction trade-offs and stoichiometry. Relationships among metabolic traits (dark respiration [R-d], maximum RuBisCo carboxylation rate [V-c,V-max], maximum electron transport rate [J(max)]) were slightly less consistent, reflecting in part their much sparser sampling (especially for high-latitude PFTs), but also pointing to more flexible plasticity in plant metabolistm. Finally, relationships involving leaf lifespan were the least consistent, indicating that leaf economic relationships related to leaf lifespan are dominated by across-PFT differences and that within-PFT variation in leaf lifespan is more complex and idiosyncratic. Across all traits, this covariance was an important source of information, as evidenced by the improved imputation accuracy and reduced predictive uncertainty in multivariate models compared to univariate models. Ultimately, our study reaffirms the value of studying not just individual traits but the multivariate trait space and the utility of hierarchical modeling for studying the scale dependence of trait relationships.Environmental Biolog

The Coordination of Leaf Photosynthesis Links C and N Fluxes in C3 Plant Species

Photosynthetic capacity is one of the most sensitive parameters in vegetation models and its relationship to leaf nitrogen content links the carbon and nitrogen cycles. Process understanding for reliably predicting photosynthetic capacity is still missing. To advance this understanding we have tested across C3 plant species the coordination hypothesis, which assumes nitrogen allocation to photosynthetic processes such that photosynthesis tends to be co-limited by ribulose-1,5-bisphosphate (RuBP) carboxylation and regeneration. The coordination hypothesis yields an analytical solution to predict photosynthetic capacity and calculate area-based leaf nitrogen content (Na). The resulting model linking leaf photosynthesis, stomata conductance and nitrogen investment provides testable hypotheses about the physiological regulation of these processes. Based on a dataset of 293 observations for 31 species grown under a range of environmental conditions, we confirm the coordination hypothesis: under mean environmental conditions experienced by leaves during the preceding month, RuBP carboxylation equals RuBP regeneration. We identify three key parameters for photosynthetic coordination: specific leaf area and two photosynthetic traits (k3, which modulates N investment and is the ratio of RuBP carboxylation/oxygenation capacity () to leaf photosynthetic N content (Npa); and Jfac, which modulates photosynthesis for a given k3 and is the ratio of RuBP regeneration capacity (Jmax) to). With species-specific parameter values of SLA, k3 and Jfac, our leaf photosynthesis coordination model accounts for 93% of the total variance in Na across species and environmental conditions. A calibration by plant functional type of k3 and Jfac still leads to accurate model prediction of Na, while SLA calibration is essentially required at species level. Observed variations in k3 and Jfac are partly explained by environmental and phylogenetic constraints, while SLA variation is partly explained by phylogeny. These results open a new avenue for predicting photosynthetic capacity and leaf nitrogen content in vegetation models

Contrasting effects of long term versus short-term nitrogen addition on photosynthesis and respiration in the Arctic

We examined the effects of short (<1–4 years) and long-term (22 years) nitrogen (N) and/or phosphorus (P) addition on the foliar CO2 exchange parameters of the Arctic species Betula nana and Eriophorum vaginatum in northern Alaska. Measured variables included: the carboxylation efficiency of Rubisco (Vcmax), electron transport capacity (Jmax), dark respiration (Rd), chlorophyll a and b content (Chl), and total foliar N (N). For both B. nana and E. vaginatum, foliar N increased by 20–50 % as a consequence of 1–22 years of fertilisation, respectively, and for B. nana foliar N increase was consistent throughout the whole canopy. However, despite this large increase in foliar N, no significant changes in Vcmax and Jmax were observed. In contrast, Rd was significantly higher (>25 %) in both species after 22 years of N addition, but not in the shorter-term treatments. Surprisingly, Chl only increased in both species the first year of fertilisation (i.e. the first season of nutrients applied), but not in the longer-term treatments. These results imply that: (1) under current (low) N availability, these Arctic species either already optimize their photosynthetic capacity per leaf area, or are limited by other nutrients; (2) observed increases in Arctic NEE and GPP with increased nutrient availability are caused by structural changes like increased leaf area index, rather than increased foliar photosynthetic capacity and (3) short-term effects (1–4 years) of nutrient addition cannot always be extrapolated to a larger time scale, which emphasizes the importance of long-term ecological experiments