Partitioning seasonal stem carbon dioxide efflux into stem respiration, bark photosynthesis, and transport-related flux in Scots pine.

Stem CO2 efflux is an important component of the carbon balance in forests. The efflux is considered to principally reflect the net result of two dominating and opposing processes: stem respiration and stem photosynthesis. In addition, transport of CO2 in xylem sap is thought to play an appreciable role in affecting the net flux. This work presents an approach to partition stem CO2 efflux among these processes using sap-flux data and CO2-exchange measurements from dark and transparent chambers placed on mature Scots pine (Pinus sylvestris) trees. Seasonal changes and monthly parameters describing the studied processes were determined. Respiration contributed most to stem net CO2 flux, reaching up to 79% (considering the sum of the absolute values of stem respiration, stem photosynthesis, and flux from CO2 transported in xylem sap to be 100%) in June, when stem growth was greatest. The contribution of photosynthesis accounted for up to 13% of the stem net CO2 flux, increasing over the monitoring period. CO2 transported axially with sap flow decreased towards the end of the growing season. At a reference temperature, respiration decreased starting around midsummer, while its temperature sensitivity increased during the summer. A decline was observed for photosynthetic quantum yield around midsummer together with a decrease in light-saturation point. The proposed approach facilitates modeling net stem CO2 flux at a range of time scales.

Does growing atmospheric CO2 explain increasing carbon sink in a boreal coniferous forest?

The terrestrial net ecosystem productivity (NEP) has increased during the past three decades, but the mechanisms responsible are still unclear. We analyzed 17 years (2001-2017) of eddy-covariance measurements of NEP, evapotranspiration (ET) and light and water use efficiency from a boreal coniferous forest in Southern Finland for trends and inter-annual variability (IAV). The forest was a mean annual carbon sink (252 [ ± 42] gC m-2a-1 ), and NEP increased at rate +6.4-7.0 gC m-2a-1 (or ca. +2.5% a-1 ) during the period. This was attributed to the increasing gross-primary productivity GPP and occurred without detectable change in ET. The start of annual carbon uptake period was advanced by 0.7 d a-1 , and increase in GPP and NEP outside the main growing season contributed ca. one-third and one-fourth of the annual trend, respectively. Meteorological factors were responsible for the IAV of fluxes but did not explain the long-term trends. The growing season GPP trend was strongest in ample light during the peak growing season. Using a multi-layer ecosystem model, we showed that direct CO2 fertilization effect diminishes when moving from leaf to ecosystem, and only 30-40% of the observed ecosystem GPP increase could be attributed to CO2 . The increasing trend in leaf-area index (LAI), stimulated by forest thinning in 2002, was the main driver of the enhanced GPP and NEP of the mid-rotation managed forest. It also compensated for the decrease of mean leaf stomatal conductance with increasing CO2 and LAI, explaining the apparent proportionality between observed GPP and CO2 trends. The results emphasize that attributing trends to their physical and physiological drivers is challenged by strong IAV, and uncertainty of LAI and species composition changes due to the dynamic flux footprint. The results enlighten the underlying mechanisms responsible for the increasing terrestrial carbon uptake in the boreal zone.

Influences of light and humidity on carbonyl sulfide-based estimates of photosynthesis.

Understanding climate controls on gross primary productivity (GPP) is crucial for accurate projections of the future land carbon cycle. Major uncertainties exist due to the challenge in separating GPP and respiration from observations of the carbon dioxide (CO2) flux. Carbonyl sulfide (COS) has a dominant vegetative sink, and plant COS uptake is used to infer GPP through the leaf relative uptake (LRU) ratio of COS to CO2 fluxes. However, little is known about variations of LRU under changing environmental conditions and in different phenological stages. We present COS and CO2 fluxes and LRU of Scots pine branches measured in a boreal forest in Finland during the spring recovery and summer. We find that the diurnal dynamics of COS uptake is mainly controlled by stomatal conductance, but the leaf internal conductance could significantly limit the COS uptake during the daytime and early in the season. LRU varies with light due to the differential light responses of COS and CO2 uptake, and with vapor pressure deficit (VPD) in the peak growing season, indicating a humidity-induced stomatal control. Our COS-based GPP estimates show that it is essential to incorporate the variability of LRU with environmental variables for accurate estimation of GPP on ecosystem, regional, and global scales.

Methane production and oxidation potentials along a fen-bog gradient from southern boreal to subarctic peatlands in Finland.

Methane (CH4 ) emissions from northern peatlands are projected to increase due to climate change, primarily because of projected increases in soil temperature. Yet, the rates and temperature responses of the two CH4 emission-related microbial processes (CH4 production by methanogens and oxidation by methanotrophs) are poorly known. Further, peatland sites within a fen-bog gradient are known to differ in the variables that regulate these two mechanisms, yet the interaction between peatland type and temperature lacks quantitative understanding. Here, we investigated potential CH4 production and oxidation rates for 14 peatlands in Finland located between c. 60 and 70°N latitude, representing bogs, poor fens, and rich fens. Potentials were measured at three different temperatures (5, 17.5, and 30℃) using the laboratory incubation method. We linked CH4 production and oxidation patterns to their methanogen and methanotroph abundance, peat properties, and plant functional types. We found that the rich fen-bog gradient-related nutrient availability and methanogen abundance increased the temperature response of CH4 production, with rich fens exhibiting the greatest production potentials. Oxidation potential showed a steeper temperature response than production, which was explained by aerenchymous plant cover, peat water holding capacity, peat nitrogen, and sulfate content. The steeper temperature response of oxidation suggests that, at higher temperatures, CH4 oxidation might balance increased CH4 production. Predicting net CH4  fluxes as an outcome of the two mechanisms is complicated due to their different controls and temperature responses. The lack of correlation between field CH4  fluxes and production/oxidation potentials, and the positive correlation with aerenchymous plants points toward the essential role of CH4 transport for emissions. The scenario of drying peatlands under climate change, which is likely to promote Sphagnum establishment over brown mosses in many places, will potentially reduce the predicted warming-related increase in CH4 emissions by shifting rich fens to Sphagnum-dominated systems.

Identifying dominant environmental predictors of freshwater wetland methane fluxes across diurnal to seasonal time scales.

While wetlands are the largest natural source of methane (CH4 ) to the atmosphere, they represent a large source of uncertainty in the global CH4 budget due to the complex biogeochemical controls on CH4 dynamics. Here we present, to our knowledge, the first multi-site synthesis of how predictors of CH4 fluxes (FCH4) in freshwater wetlands vary across wetland types at diel, multiday (synoptic), and seasonal time scales. We used several statistical approaches (correlation analysis, generalized additive modeling, mutual information, and random forests) in a wavelet-based multi-resolution framework to assess the importance of environmental predictors, nonlinearities and lags on FCH4 across 23 eddy covariance sites. Seasonally, soil and air temperature were dominant predictors of FCH4 at sites with smaller seasonal variation in water table depth (WTD). In contrast, WTD was the dominant predictor for wetlands with smaller variations in temperature (e.g., seasonal tropical/subtropical wetlands). Changes in seasonal FCH4 lagged fluctuations in WTD by ~17 ± 11 days, and lagged air and soil temperature by median values of 8 ± 16 and 5 ± 15 days, respectively. Temperature and WTD were also dominant predictors at the multiday scale. Atmospheric pressure (PA) was another important multiday scale predictor for peat-dominated sites, with drops in PA coinciding with synchronous releases of CH4 . At the diel scale, synchronous relationships with latent heat flux and vapor pressure deficit suggest that physical processes controlling evaporation and boundary layer mixing exert similar controls on CH4 volatilization, and suggest the influence of pressurized ventilation in aerenchymatous vegetation. In addition, 1- to 4-h lagged relationships with ecosystem photosynthesis indicate recent carbon substrates, such as root exudates, may also control FCH4. By addressing issues of scale, asynchrony, and nonlinearity, this work improves understanding of the predictors and timing of wetland FCH4 that can inform future studies and models, and help constrain wetland CH4 emissions.

Leaf carbon and water status control stomatal and nonstomatal limitations of photosynthesis in trees.

Photosynthetic rate is concurrently limited by stomatal limitations and nonstomatal limitations (NSLs). However, the controls on NSLs to photosynthesis and their coordination with stomatal control on different timescales remain poorly understood. According to a recent optimization hypothesis, NSLs depend on leaf osmotic or water status and are coordinated with stomatal control so as to maximize leaf photosynthesis. Drought and notching experiments were conducted on Pinus sylvestris, Picea abies, Betula Pendula and Populus tremula seedlings in glasshouse conditions to study the dependence of NSLs on leaf osmotic and water status, and their coordination with stomatal control, on timescales of minutes and weeks, to test the assumptions and predictions of the optimization hypothesis. Both NSLs and stomatal conductance followed power-law functions of leaf osmotic concentration and leaf water potential. Moreover, stomatal conductance was proportional to the square root of soil-to-leaf hydraulic conductance, as predicted by the optimization hypothesis. Though the detailed mechanisms underlying the dependence of NSLs on leaf osmotic or water status lie outside the scope of this study, our results support the hypothesis that NSLs and stomatal control are coordinated to maximize leaf photosynthesis and allow the effect of NSLs to be included in models of tree gas-exchange.

Early snowmelt significantly enhances boreal springtime carbon uptake.

We determine the annual timing of spring recovery from space-borne microwave radiometer observations across northern hemisphere boreal evergreen forests for 1979-2014. We find a trend of advanced spring recovery of carbon uptake for this period, with a total average shift of 8.1 d (2.3 d/decade). We use this trend to estimate the corresponding changes in gross primary production (GPP) by applying in situ carbon flux observations. Micrometeorological CO2 measurements at four sites in northern Europe and North America indicate that such an advance in spring recovery would have increased the January-June GPP sum by 29 g⋅C⋅m-2 [8.4 g⋅C⋅m-2 (3.7%)/decade]. We find this sensitivity of the measured springtime GPP to the spring recovery to be in accordance with the corresponding sensitivity derived from simulations with a land ecosystem model coupled to a global circulation model. The model-predicted increase in springtime cumulative GPP was 0.035 Pg/decade [15.5 g⋅C⋅m-2 (6.8%)/decade] for Eurasian forests and 0.017 Pg/decade for forests in North America [9.8 g⋅C⋅m-2 (4.4%)/decade]. This change in the springtime sum of GPP related to the timing of spring snowmelt is quantified here for boreal evergreen forests.

New insights into the covariation of stomatal, mesophyll and hydraulic conductances from optimization models incorporating nonstomatal limitations to photosynthesis.

Optimization models of stomatal conductance (gs ) attempt to explain observed stomatal behaviour in terms of cost--benefit tradeoffs. While the benefit of stomatal opening through increased CO2 uptake is clear, currently the nature of the associated cost(s) remains unclear. We explored the hypothesis that gs maximizes leaf photosynthesis, where the cost of stomatal opening arises from nonstomatal reductions in photosynthesis induced by leaf water stress. We analytically solved two cases, CAP and MES, in which reduced leaf water potential leads to reductions in carboxylation capacity (CAP) and mesophyll conductance (gm ) (MES). Both CAP and MES predict the same one-parameter relationship between the intercellular : atmospheric CO2 concentration ratio (ci /ca ) and vapour pressure deficit (VPD, D), viz. ci /ca  ≈ ξ/(ξ + âˆšD), as that obtained from previous optimization models, with the novel feature that the parameter ξ is determined unambiguously as a function of a small number of photosynthetic and hydraulic variables. These include soil-to-leaf hydraulic conductance, implying a stomatal closure response to drought. MES also predicts that gs /gm is closely related to ci /ca and is similarly conservative. These results are consistent with observations, give rise to new testable predictions, and offer new insights into the covariation of stomatal, mesophyll and hydraulic conductances.

Joint control of terrestrial gross primary productivity by plant phenology and physiology.

Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate-carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy-covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000-2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r(2) = 0.90) and GPP recovery after a fire disturbance in South Dakota (r(2) = 0.88). Additional analysis of the eddy-covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

The uncertain climate footprint of wetlands under human pressure.

Significant climate risks are associated with a positive carbon-temperature feedback in northern latitude carbon-rich ecosystems, making an accurate analysis of human impacts on the net greenhouse gas balance of wetlands a priority. Here, we provide a coherent assessment of the climate footprint of a network of wetland sites based on simultaneous and quasi-continuous ecosystem observations of CO2 and CH4 fluxes. Experimental areas are located both in natural and in managed wetlands and cover a wide range of climatic regions, ecosystem types, and management practices. Based on direct observations we predict that sustained CH4 emissions in natural ecosystems are in the long term (i.e., several centuries) typically offset by CO2 uptake, although with large spatiotemporal variability. Using a space-for-time analogy across ecological and climatic gradients, we represent the chronosequence from natural to managed conditions to quantify the "cost" of CH4 emissions for the benefit of net carbon sequestration. With a sustained pulse-response radiative forcing model, we found a significant increase in atmospheric forcing due to land management, in particular for wetland converted to cropland. Our results quantify the role of human activities on the climate footprint of northern wetlands and call for development of active mitigation strategies for managed wetlands and new guidelines of the Intergovernmental Panel on Climate Change (IPCC) accounting for both sustained CH4 emissions and cumulative CO2 exchange.

Do the energy fluxes and surface conductance of boreal coniferous forests in Europe scale with leaf area?

Earth observing systems are now routinely used to infer leaf area index (LAI) given its significance in spatial aggregation of land surface fluxes. Whether LAI is an appropriate scaling parameter for daytime growing season energy budget, surface conductance (Gs ), water- and light-use efficiency and surface-atmosphere coupling of European boreal coniferous forests was explored using eddy-covariance (EC) energy and CO2 fluxes. The observed scaling relations were then explained using a biophysical multilayer soil-vegetation-atmosphere transfer model as well as by a bulk Gs representation. The LAI variations significantly alter radiation regime, within-canopy microclimate, sink/source distributions of CO2 , H2 O and heat, and forest floor fluxes. The contribution of forest floor to ecosystem-scale energy exchange is shown to decrease asymptotically with increased LAI, as expected. Compared with other energy budget components, dry-canopy evapotranspiration (ET) was reasonably 'conservative' over the studied LAI range 0.5-7.0 m2 m-2 . Both ET and Gs experienced a minimum in the LAI range 1-2 m2 m-2 caused by opposing nonproportional response of stomatally controlled transpiration and 'free' forest floor evaporation to changes in canopy density. The young forests had strongest coupling with the atmosphere while stomatal control of energy partitioning was strongest in relatively sparse (LAI ~2 m2 m-2 ) pine stands growing on mineral soils. The data analysis and model results suggest that LAI may be an effective scaling parameter for net radiation and its partitioning but only in sparse stands (LAI <3 m2 m-2 ). This finding emphasizes the significance of stand-replacing disturbances on the controls of surface energy exchange. In denser forests, any LAI dependency varies with physiological traits such as light-saturated water-use efficiency. The results suggest that incorporating species traits and site conditions are necessary when LAI is used in upscaling energy exchanges of boreal coniferous forests.

Do small spores disperse further than large spores?

In species that disperse by airborne propagules an inverse relationship is often assumed between propagule size and dispersal distance. However, for microscopic spores the evidence for the relationship remains ambiguous. Lagrangian stochastic dispersion models that have been successful in predicting seed dispersal appear to predict similar dispersal for all spore sizes up to -40 microm diameter. However, these models have assumed that spore size affects only the downwards drift of particles due to gravitation and have largely omitted the highly size-sensitive deposition process to surfaces such as forest canopy. On the other hand, they have assumed that spores are certain to deposit when the air parcel carrying them touches the ground. Here, we supplement a Lagrangian stochastic dispersion model with a mechanistic deposition model parameterized by empirical deposition data for 1-10 microm spores. The inclusion of realistic deposition improved the ability of the model to predict empirical data on the dispersal of a wood-decay fungus (aerodynamic spore size 3.8 microm). Our model predicts that the dispersal of 1-10 microm spores is in fact highly sensitive to spore size, with 97-98% of 1 microm spores but only 12-58% of 10-microm spores dispersing beyond 2 km in the simulated range of wind and canopy conditions. Further, excluding the assumption of certain deposition at the ground greatly increased the expected dispersal distances throughout the studied spore size range. Our results suggest that by evolutionary adjustment of spore size, release height and timing of release, fungi and other organisms with microscopic spores can change the expected distribution of dispersal locations markedly. The complex interplay of wind and canopy conditions in determining deposition resulted in some counterintuitive predictions, such as that spores disperse furthest under intermediate wind, providing intriguing hypotheses to be tested empirically in future studies.

Latent heat exchange in the boreal and arctic biomes.

In this study latent heat flux (λE) measurements made at 65 boreal and arctic eddy-covariance (EC) sites were analyses by using the Penman-Monteith equation. Sites were stratified into nine different ecosystem types: harvested and burnt forest areas, pine forests, spruce or fir forests, Douglas-fir forests, broadleaf deciduous forests, larch forests, wetlands, tundra and natural grasslands. The Penman-Monteith equation was calibrated with variable surface resistances against half-hourly eddy-covariance data and clear differences between ecosystem types were observed. Based on the modeled behavior of surface and aerodynamic resistances, surface resistance tightly control λE in most mature forests, while it had less importance in ecosystems having shorter vegetation like young or recently harvested forests, grasslands, wetlands and tundra. The parameters of the Penman-Monteith equation were clearly different for winter and summer conditions, indicating that phenological effects on surface resistance are important. We also compared the simulated λE of different ecosystem types under meteorological conditions at one site. Values of λE varied between 15% and 38% of the net radiation in the simulations with mean ecosystem parameters. In general, the simulations suggest that λE is higher from forested ecosystems than from grasslands, wetlands or tundra-type ecosystems. Forests showed usually a tighter stomatal control of λE as indicated by a pronounced sensitivity of surface resistance to atmospheric vapor pressure deficit. Nevertheless, the surface resistance of forests was lower than for open vegetation types including wetlands. Tundra and wetlands had higher surface resistances, which were less sensitive to vapor pressure deficits. The results indicate that the variation in surface resistance within and between different vegetation types might play a significant role in energy exchange between terrestrial ecosystems and atmosphere. These results suggest the need to take into account vegetation type and phenology in energy exchange modeling.

Net carbon dioxide losses of northern ecosystems in response to autumn warming.

The carbon balance of terrestrial ecosystems is particularly sensitive to climatic changes in autumn and spring, with spring and autumn temperatures over northern latitudes having risen by about 1.1 degrees C and 0.8 degrees C, respectively, over the past two decades. A simultaneous greening trend has also been observed, characterized by a longer growing season and greater photosynthetic activity. These observations have led to speculation that spring and autumn warming could enhance carbon sequestration and extend the period of net carbon uptake in the future. Here we analyse interannual variations in atmospheric carbon dioxide concentration data and ecosystem carbon dioxide fluxes. We find that atmospheric records from the past 20 years show a trend towards an earlier autumn-to-winter carbon dioxide build-up, suggesting a shorter net carbon uptake period. This trend cannot be explained by changes in atmospheric transport alone and, together with the ecosystem flux data, suggest increasing carbon losses in autumn. We use a process-based terrestrial biosphere model and satellite vegetation greenness index observations to investigate further the observed seasonal response of northern ecosystems to autumnal warming. We find that both photosynthesis and respiration increase during autumn warming, but the increase in respiration is greater. In contrast, warming increases photosynthesis more than respiration in spring. Our simulations and observations indicate that northern terrestrial ecosystems may currently lose carbon dioxide in response to autumn warming, with a sensitivity of about 0.2 PgC degrees C(-1), offsetting 90% of the increased carbon dioxide uptake during spring. If future autumn warming occurs at a faster rate than in spring, the ability of northern ecosystems to sequester carbon may be diminished earlier than previously suggested.

Ruptures of mixed lipid monolayers under tension and supercooling: implications for nanobubbles in plants.

Mixed phospholipid and glycolipid monolayers likely coat the surfaces of pressurised gas nanobubbles within the hydraulic systems of plants. The lipid coatings bond to water under negative pressure and are thus stretched out of equilibrium. In this work, we have used molecular dynamics simulations to produce trajectories of a biologically relevant mixed monolayer, pulled at mild negative pressures (-1.5 to -4.5 MPa). Pore formation within the monolayer is observed at both 270 and 310 K, and proceeds as an activated process once the lipid tails fully transition from the two dimensional liquid condensed to liquid expanded phase. Pressure:area isotherms showed reduced surface pressure under slight supercooling (T = 270 K) at all observed areas per lipid. Finally, Rayleigh-Plesset simulations were used to predict evolving nanobubble size using the calculated pressure:area isotherms as dynamic surface tensions. We confirm the existence of a second critical radius with respect to runaway growth, above the homogeneous cavitation radius.

Assimilate transport in phloem sets conditions for leaf gas exchange.

Carbon uptake and transpiration in plant leaves occurs through stomata that open and close. Stomatal action is usually considered a response to environmental driving factors. Here we show that leaf gas exchange is more strongly related to whole tree level transport of assimilates than previously thought, and that transport of assimilates is a restriction of stomatal opening comparable with hydraulic limitation. Assimilate transport in the phloem requires that osmotic pressure at phloem loading sites in leaves exceeds the drop in hydrostatic pressure that is due to transpiration. Assimilate transport thus competes with transpiration for water. Excess sugar loading, however, may block the assimilate transport because of viscosity build-up in phloem sap. Therefore, for given conditions, there is a stomatal opening that maximizes phloem transport if we assume that sugar loading is proportional to photosynthetic rate. Here we show that such opening produces the observed behaviour of leaf gas exchange. Our approach connects stomatal regulation directly with sink activity, plant structure and soil water availability as they all influence assimilate transport. It produces similar behaviour as the optimal stomatal control approach, but does not require determination of marginal cost of water parameter.

A single-point modeling approach for the intercomparison and evaluation of ozone dry deposition across chemical transport models (Activity 2 of AQMEII4).

A primary sink of air pollutants and their precursors is dry deposition. Dry deposition estimates differ across chemical transport models, yet an understanding of the model spread is incomplete. Here, we introduce Activity 2 of the Air Quality Model Evaluation International Initiative Phase 4 (AQMEII4). We examine 18 dry deposition schemes from regional and global chemical transport models as well as standalone models used for impact assessments or process understanding. We configure the schemes as single-point models at eight Northern Hemisphere locations with observed ozone fluxes. Single-point models are driven by a common set of site-specific meteorological and environmental conditions. Five of eight sites have at least 3 years and up to 12 years of ozone fluxes. The interquartile range across models in multiyear mean ozone deposition velocities ranges from a factor of 1.2 to 1.9 annually across sites and tends to be highest during winter compared with summer. No model is within 50 % of observed multiyear averages across all sites and seasons, but some models perform well for some sites and seasons. For the first time, we demonstrate how contributions from depositional pathways vary across models. Models can disagree with respect to relative contributions from the pathways, even when they predict similar deposition velocities, or agree with respect to the relative contributions but predict different deposition velocities. Both stomatal and nonstomatal uptake contribute to the large model spread across sites. Our findings are the beginning of results from AQMEII4 Activity 2, which brings scientists who model air quality and dry deposition together with scientists who measure ozone fluxes to evaluate and improve dry deposition schemes in the chemical transport models used for research, planning, and regulatory purposes.

Intercomparison of methods to estimate gross primary production based on CO2 and COS flux measurements.

Separating the components of ecosystem-scale carbon exchange is crucial in order to develop better models and future predictions of the terrestrial carbon cycle. However, there are several uncertainties and unknowns related to current photosynthesis estimates. In this study, we evaluate four different methods for estimating photosynthesis at a boreal forest at the ecosystem scale, of which two are based on carbon dioxide (CO2) flux measurements and two on carbonyl sulfide (COS) flux measurements. The CO2-based methods use traditional flux partitioning and artificial neural networks to separate the net CO2 flux into respiration and photosynthesis. The COS-based methods make use of a unique 5-year COS flux data set and involve two different approaches to determine the leaf-scale relative uptake ratio of COS and CO2 (LRU), of which one (LRUCAP) was developed in this study. LRUCAP was based on a previously tested stomatal optimization theory (CAP), while LRUPAR was based on an empirical relation to measured radiation. For the measurement period 2013-2017, the artificial neural network method gave a GPP estimate very close to that of traditional flux partitioning at all timescales. On average, the COS-based methods gave higher GPP estimates than the CO2-based estimates on daily (23% and 7% higher, using LRUPAR and LRUCAP, respectively) and monthly scales (20% and 3% higher), as well as a higher cumulative sum over 3 months in all years (on average 25% and 3% higher). LRUCAP was higher than LRU estimated from chamber measurements at high radiation, leading to underestimation of midday GPP relative to other GPP methods. In general, however, use of LRUCAP gave closer agreement with CO2-based estimates of GPP than use of LRUPAR. When extended to other sites, LRUCAP may be more robust than LRUPAR because it is based on a physiological model whose parameters can be estimated from simple measurements or obtained from the literature. In contrast, the empirical radiation relation in LRUPAR may be more site-specific. However, this requires further testing at other measurement sites.

Dynamic Surface Tension Enhances the Stability of Nanobubbles in Xylem Sap.

Air seeded nanobubbles have recently been observed within tree sap under negative pressure. They are stabilized by an as yet unidentified process, although some embolize their vessels in extreme circumstances. Current literature suggests that a varying surface tension helps bubbles survive, but few direct measurements of this quantity have been made. Here, we present calculations of dynamic surface tension for two biologically relevant lipids using molecular dynamics simulations. We find that glycolipid monolayers resist expansion proportionally to the rate of expansion. Their surface tension increases with the tension applied, in a similar way to the viscosity of a non-Newtonian fluid. In contrast, a prototypical phospholipid was equally resistant to all applied tensions, suggesting that the fate of a given nanobubble is dependent on its surface composition. By incorporating our results into a Classical Nucleation Theory (CNT) framework, we predict nanobubble stability with respect to embolism. We find that the metastable radius of glycolipid coated nanobubbles is approximately 35 nm, and that embolism is in this case unlikely when the external pressure is less negative than -1.5 MPa.