Finding balance: Tree-ring isotopes differentiate between acclimation and stress-induced imbalance in a long-term irrigation experiment.

Scots pine (Pinus sylvestris L.) is a common European tree species, and understanding its acclimation to the rapidly changing climate through physiological, biochemical or structural adjustments is vital for predicting future growth. We investigated a long-term irrigation experiment at a naturally dry forest in Switzerland, comparing Scots pine trees that have been continuously irrigated for 17 years (irrigated) with those for which irrigation was interrupted after 10 years (stop) and non-irrigated trees (control), using tree growth, xylogenesis, wood anatomy, and carbon, oxygen and hydrogen stable isotope measurements in the water, sugars and cellulose of plant tissues. The dendrochronological analyses highlighted three distinct acclimation phases to the treatments: irrigated trees experienced (i) a significant growth increase in the first 4 years of treatment, (ii) high growth rates but with a declining trend in the following 8 years and finally (iii) a regression to pre-irrigation growth rates, suggesting the development of a new growth limitation (i.e. acclimation). The introduction of the stop treatment resulted in further growth reductions to below-control levels during the third phase. Irrigated trees showed longer growth periods and lower tree-ring δ13 C values, reflecting lower stomatal restrictions than control trees. Their strong tree-ring δ18 O and δ2 H (O-H) relationship reflected the hydrological signature similarly to the control. On the contrary, the stop trees had lower growth rates, conservative wood anatomical traits, and a weak O-H relationship, indicating a physiological imbalance. Tree vitality (identified by crown transparency) significantly modulated growth, wood anatomical traits and tree-ring δ13 C, with low-vitality trees of all treatments performing similarly regardless of water availability. We thus provide quantitative indicators for assessing physiological imbalance and tree acclimation after environmental stresses. We also show that tree vitality is crucial in shaping such responses. These findings are fundamental for the early assessment of ecosystem imbalances and decline under climate change.

Quantifying effects of cold acclimation and delayed springtime photosynthesis resumption in northern ecosystems.

Land carbon dynamics in temperate and boreal ecosystems are sensitive to environmental change. Accurately simulating gross primary productivity (GPP) and its seasonality is key for reliable carbon cycle projections. However, significant biases have been found in early spring GPP simulations of northern forests, where observations often suggest a later resumption of photosynthetic activity than predicted by models. Here, we used eddy covariance-based GPP estimates from 39 forest sites that differ by their climate and dominant plant functional types. We used a mechanistic and an empirical light use efficiency (LUE) model to investigate the magnitude and environmental controls of delayed springtime photosynthesis resumption (DSPR) across sites. We found DSPR reduced ecosystem LUE by 30-70% at many, but not all site-years during spring. A significant depression of LUE was found not only in coniferous but also at deciduous forests and was related to combined high radiation and low minimum temperatures. By embedding cold-acclimation effects on LUE that considers the delayed effects of minimum temperatures, initial model bias in simulated springtime GPP was effectively resolved. This provides an approach to improve GPP estimates by considering physiological acclimation and enables more reliable simulations of photosynthesis in northern forests and projections in a warming climate.

Absence of canopy temperature variation despite stomatal adjustment in Pinus sylvestris under multidecadal soil moisture manipulation.

Global warming and droughts push forests closer to their thermal limits, altering tree carbon uptake and growth. To prevent critical overheating, trees can adjust their thermotolerance (Tcrit ), temperature and photosynthetic optima (Topt and Aopt ), and canopy temperature (Tcan ) to stay below damaging thresholds. However, we lack an understanding of how soil droughts affect photosynthetic thermal plasticity and Tcan regulation. In this study, we measured the effect of soil moisture on the seasonal and diurnal dynamics of net photosynthesis (A), stomatal conductance (gs ), and Tcan , as well as the thermal plasticity of photosynthesis (Tcrit , Topt , and Aopt ), over the course of 1 yr using a long-term irrigation experiment in a drought-prone Pinus sylvestris forest in Switzerland. Irrigation resulted in higher needle-level A, gs , Topt , and Aopt compared with naturally drought-exposed trees. No daily or seasonal differences in Tcan were observed between treatments. Trees operated below their thermal thresholds (Tcrit ), independently of soil moisture content. Despite strong Tcan and Tair coupling, we provide evidence that drought reduces trees' temperature optimum due to a substantial reduction of gs during warm and dry periods of the year. These findings provide important insights regarding the effects of soil drought on the thermal tolerance of P. sylvestris.

Networking the forest infrastructure towards near real-time monitoring - A white paper.

Forests account for nearly 90 % of the world's terrestrial biomass in the form of carbon and they support 80 % of the global biodiversity. To understand the underlying forest dynamics, we need a long-term but also relatively high-frequency, networked monitoring system, as traditionally used in meteorology or hydrology. While there are numerous existing forest monitoring sites, particularly in temperate regions, the resulting data streams are rarely connected and do not provide information promptly, which hampers real-time assessments of forest responses to extreme climate events. The technology to build a better global forest monitoring network now exists. This white paper addresses the key structural components needed to achieve a novel meta-network. We propose to complement - rather than replace or unify - the existing heterogeneous infrastructure with standardized, quality-assured linking methods and interacting data processing centers to create an integrated forest monitoring network. These automated (research topic-dependent) linking methods in atmosphere, biosphere, and pedosphere play a key role in scaling site-specific results and processing them in a timely manner. To ensure broad participation from existing monitoring sites and to establish new sites, these linking methods must be as informative, reliable, affordable, and maintainable as possible, and should be supplemented by near real-time remote sensing data. The proposed novel meta-network will enable the detection of emergent patterns that would not be visible from isolated analyses of individual sites. In addition, the near real-time availability of data will facilitate predictions of current forest conditions (nowcasts), which are urgently needed for research and decision making in the face of rapid climate change. We call for international and interdisciplinary efforts in this direction.

Photosynthetic acclimation and sensitivity to short- and long-term environmental changes in a drought-prone forest.

Future climate will be characterized by an increase in frequency and duration of drought and warming that exacerbates atmospheric evaporative demand. How trees acclimate to long-term soil moisture changes and whether these long-term changes alter trees' sensitivity to short-term (day to months) variations of vapor pressure deficit (VPD) and soil moisture is largely unknown. Leaf gas exchange measurements were performed within a long-term (17 years) irrigation experiment in a drought-prone Scots pine-dominated forest in one of Switzerland's driest areas on trees in naturally dry (control), irrigated, and 'irrigation-stop' (after 11 years of irrigation) conditions. Seventeen years of irrigation increased photosynthesis (A) and stomatal conductance (gs) and reduced gs sensitivity to increasing VPD and soil drying. Following irrigation-stop, gas exchange decreased only after 3 years. After 5 years, maximum carboxylation (Vcmax) and electron transport (Jmax) rates in irrigation-stop recovered to similar levels as to before the irrigation-stop. These results suggest that long-term release from soil drought reduces the sensitivity to VPD and that atmospheric constraints may play an increasingly important role in combination with soil drought. Moreover, our study indicates that structural adjustments lead to an attenuation of initially strong leaf-level acclimation to strong multiple-year drought.

Drone-based physiological index reveals long-term acclimation and drought stress responses in trees.

Monitoring early tree physiological responses to drought is key to understanding progressive impacts of drought on forests and identifying resilient species. We combined drone-based multispectral remote sensing with measurements of tree physiology and environmental parameters over two growing seasons in a 100-y-old Pinus sylvestris forest subject to 17-y of precipitation manipulation. Our goal was to determine if drone-based photochemical reflectance index (PRI) captures tree drought stress responses and whether responses are affected by long-term acclimation. PRI detects changes in xanthophyll cycle pigment dynamics, which reflect increases in photoprotective non-photochemical quenching activity resulting from drought-induced photosynthesis downregulation. Here, PRI of never-irrigated trees was up to 10 times lower (higher stress) than PRI of irrigated trees. Long-term acclimation to experimental treatment, however, influenced the seasonal relationship between PRI and soil water availability. PRI also captured diurnal decreases in photochemical efficiency, driven by vapour pressure deficit. Interestingly, 5 years after irrigation was stopped for a subset of the irrigated trees, a positive legacy effect persisted, with lower stress responses (higher PRI) compared with never-irrigated trees. This study demonstrates the ability of remotely sensed PRI to scale tree physiological responses to an entire forest and the importance of long-term acclimation in determining current drought stress responses.

Canopy photosynthesis of six major arable crops is enhanced under diffuse light due to canopy architecture.

Diffuse radiation generally increases photosynthetic rates if total radiation is kept constant. Different hypotheses have been proposed to explain this enhancement of photosynthesis, but conclusive results over a wide range of diffuse conditions or about the effect of canopy architecture are lacking. Here, we show the response of canopy photosynthesis to different fractions of diffuse light conditions for five major arable crops (pea, potato, wheat, barley, rapeseed) and cover crops characterized by different canopy architecture. We used 13 years of flux and microclimate measurements over a field with a typical 4 year crop rotation scheme in Switzerland. We investigated the effect of diffuse light on photosynthesis over a gradient of diffuse light fractions ranging from 100% diffuse (overcast sky) to 11% diffuse light (clear-sky conditions). Gross primary productivity (GPP) increased with diffuse fraction and thus was greater under diffuse than direct light conditions if the absolute photon flux density per unit surface area was kept constant. Mean leaf tilt angle (MTA) and canopy height were found to be the best predictors of the diffuse versus direct radiation effect on photosynthesis. Climatic factors, such as the drought index and growing degree days (GDD), had a significant influence on initial quantum yield under direct but not diffuse light conditions, which depended primarily on MTA. The maximum photosynthetic rate at 2,000 µmol m-2  s-1 photosynthetically active radiation under direct conditions strongly depended on GDD, MTA, leaf area index (LAI) and the interaction between MTA and LAI, while under diffuse conditions, this parameter depended mostly on MTA and only to a minor extent on canopy height and their interaction. The strongest photosynthesis enhancement under diffuse light was found for wheat, barley and rapeseed, whereas the lowest was for pea. Thus, we suggest that measuring canopy architecture and diffuse radiation will greatly improve GPP estimates of global cropping systems.

High-throughput drone-based remote sensing reliably tracks phenology in thousands of conifer seedlings.

Phenology is an important indicator of environmental variation and climate change impacts on tree responses. In conifers, monitoring phenology of photosynthesis through remote sensing has been unreliable, because needle foliage varies little throughout the year. This is challenging for modelling ecosystem carbon uptake and monitoring phenology for enhanced breeding (genomic selection) and forest health. Here, we demonstrate that drone-based carotenoid-sensitive spectral indices, such as the Chl/carotenoid index (CCI), can be used to track phenology in conifers by taking advantage of the close relationship between seasonally changing carotenoid levels and the variation of photosynthetic activity. Physiological ground measurements, including photosynthetic pigments and maximum quantum yield of Chl fluorescence, indicated that CCI tracked the variation of photosynthetic activity better than other vegetation indices for 30 white spruce seedlings measured over 1 yr. A machine-learning approach, using CCI derived from drone-based multispectral imagery, was used to model phenology of photosynthesis for the entire pedigree population (6000 seedlings). This high-throughput drone-based phenotyping approach is suitable for studying climate change impacts and environmental variation on the physiological status of thousands of field-grown conifers at unprecedented speed and scale.

Tracking the phenology of photosynthesis using carotenoid-sensitive and near-infrared reflectance vegetation indices in a temperate evergreen and mixed deciduous forest.

Photosynthetic phenology is an important indicator of annual gross primary productivity (GPP). Assessing photosynthetic phenology remotely is difficult for evergreen conifers as they remain green year-round. Carotenoid-based vegetation indices such as the photochemical reflectance index (PRI) and chlorophyll/carotenoid index (CCI) are promising tools to remotely track the invisible phenology of photosynthesis by assessing carotenoid pigment dynamics. PRI, CCI and the near-infrared reflectance of vegetation (NIRV ) index may act as proxies of photosynthetic efficiency (ɛ), an important parameter in light-use efficiency models, or direct proxies of photosynthesis. To understand the physiological mechanisms reflected by PRI and CCI and the ability of vegetation indices to act as proxies of photosynthetic activity for estimating GPP, we measured leaf pigment composition, PRI, CCI, NIRV and photosynthetic activity at the leaf and canopy scales over 2 years in an evergreen and mixed deciduous forest. PRI and CCI captured the large seasonal carotenoid/chlorophyll ratio changes and good relationships were observed between PRI-ɛ and CCI-photosynthesis and NIRV -photosynthesis. PRI-, CCI- and NIRV -based models effectively tracked observed seasonal GPP. We propose that carotenoid-based and near-infrared reflectance vegetation indices may provide useful proxies of photosynthetic activity and can improve remote sensing-based models of GPP in evergreen and deciduous forests.

Light and VPD gradients drive foliar nitrogen partitioning and photosynthesis in the canopy of European beech and silver fir.

While foliar photosynthetic relationships with light, nitrogen, and water availability have been well described, environmental factors driving vertical gradients of foliar traits within forest canopies are still not well understood. We, therefore, examined how light availability and vapour pressure deficit (VPD) co-determine vertical gradients (between 12 and 42 m and in the understorey) of foliar photosynthetic capacity (Amax), 13C fractionation (∆), specific leaf area (SLA), chlorophyll (Chl), and nitrogen (N) concentrations in canopies of Fagus sylvatica and Abies alba growing in a mixed forest in Switzerland in spring and summer 2017. Both species showed lower Chl/N and lower SLA with higher light availability and VPD at the top canopy. Despite these biochemical and morphological acclimations, Amax during summer remained relatively constant and the photosynthetic N-use efficiency (PNUE) decreased with higher light availability for both species, suggesting suboptimal N allocation within the canopy. ∆ of both species were lower at the canopy top compared to the bottom, indicating high water-use efficiency (WUE). VPD gradients strongly co-determined the vertical distribution of Chl, N, and PNUE in F. sylvatica, suggesting stomatal limitation of photosynthesis in the top canopy, whereas these traits were only related to light availability in A. alba. Lower PNUE in F. sylvatica with higher WUE clearly indicated a trade-off in water vs. N use, limiting foliar acclimation to high light and VPD at the top canopy. Species-specific trade-offs in foliar acclimation to environmental canopy gradients may thus be considered for scaling photosynthesis from leaf to canopy to landscape levels.

Vertical patterns of photosynthesis and related leaf traits in two contrasting agricultural crops.

To include within-canopy leaf acclimation responses to light and other resource gradients in photosynthesis modelling, it is imperative to understand the variation of leaf structural, biochemical and physiological traits from canopy top to bottom. In the present study, leaf photosynthetic traits for top and bottom canopy leaves, canopy structure and light profiles, were measured over one growing season for two contrasting crop types, winter barley (Hordeum vulgare L.) and rape seed (Brassica napus L.). With the exception of quantum yield, other traits such as maximum photosynthetic capacity (Amax), dark respiration, leaf nitrogen and chlorophyll contents, and leaf mass per area, showed consistently higher (P<0.05) values for top leaves throughout the growing season and for both crop types. Even though Amax was higher for top leaves, the bottom half of the canopy intercepted more light and thus contributed the most to total canopy photosynthesis up until senescence set in. Incorporating this knowledge into a simple top/bottom-leaf upscaling scheme, separating top and bottom leaves, resulted in a better match between estimated and measured total canopy photosynthesis, compared with a one-leaf upscaling scheme. Moreover, aggregating to daily and weekly temporal resolutions progressively increased the linearity of the leaf photosynthetic responses to light for top leaves.

Changes in vegetation phenology are not reflected in atmospheric CO2 and 13 C/12 C seasonality.

Northern terrestrial ecosystems have shown global warming-induced advances in start, delays in end, and thus increased lengths of growing season and gross photosynthesis in recent decades. The tradeoffs between seasonal dynamics of two opposing fluxes, CO2 uptake through photosynthesis and release through respiration, determine the influence of the terrestrial ecosystem on the atmospheric CO2 and 13 C/12 C seasonality. Here, we use four CO2 observation stations in the Northern Hemisphere, namely Alert, La Jolla, Point Barrow, and Mauna Loa Observatory, to determine how changes in vegetation productivity and phenology, respiration, and air temperature affect both the atmospheric CO2 and 13 C/12 C seasonality. Since the 1960s, the only significant long-term trend of CO2 and 13 C/12 C seasonality was observed at the northern most station, Alert, where the spring CO2 drawdown dates advanced by 0.65 ± 0.55 days yr-1 , contributing to a nonsignificant increase in length of the CO2 uptake period (0.74 ± 0.67 days yr-1 ). For Point Barrow station, vegetation phenology changes in well-watered ecosystems such as the Canadian and western Siberian wetlands contributed the most to 13 C/12 C seasonality while the CO2 seasonality was primarily linked to nontree vegetation. Our results indicate significant increase in the Northern Hemisphere soil respiration. This means, increased respiration of 13 C depleted plant materials cancels out the 12 C gain from enhanced vegetation activities during the start and end of growing season. These findings suggest therefore that parallel warming-induced increases both in photosynthesis and respiration contribute to the long-term stability of CO2 and 13 C/12 C seasonality under changing climate and vegetation activity. The summer photosynthesis and the soil respiration in the dormant seasons have become more vigorous which lead to increased peak-to-through CO2 amplitude. As the relative magnitude of the increased photosynthesis in summer months is more than the increased respiration in dormant months, we have the increased overall carbon uptake rates in the northern ecosystems.

Citizen science: best practices to remove observer bias in trend analysis.

Citizen science, time series records over long periods of time, and wide geographic areas offer many opportunities for scientists to answer questions that would otherwise be impractical to investigate. Citizen scientists currently play active roles in a wide range of ecological projects; however, observer biases such as varying perception of events or objects being observed and quality of observations present challenges to successfully derive interannual variability and trend statistics from time series records. It is recommended that citizen science records, particularly those involving events such as plant phenology, should not be directly averaged across sites. The interannual variability expressed as an anomaly and trend expressed as a regression slope should be calculated for each site. Only the site level anomaly and regression slopes should be averaged to suppress observer biases.

Performance assessment of onboard and scene-based methods for Airborne Prism Experiment spectral characterization.

Accurate spectral calibration of airborne and spaceborne imaging spectrometers is essential for proper preprocessing and scientific exploitation of high spectral resolution measurements of the land and atmosphere. A systematic performance assessment of onboard and scene-based methods for in-flight monitoring of instrument spectral calibration is presented for the first time in this paper. Onboard and ground imaging data were collected at several flight altitudes using the Airborne Prism Experiment (APEX) imaging spectrometer. APEX is equipped with an in-flight characterization (IFC) facility allowing the evaluation of radiometric, spectral, and geometric system properties, both in-flight and on-ground for the full field of view. Atmospheric and onboard filter spectral features present in at-sensor radiances are compared with the same features in reference transmittances convolved to varying instrument spectral configurations. A spectrum-matching algorithm, taking advantage of the high sensitivity of measurements around sharp spectral features toward spectrometer spectral performance, is used to retrieve channel center wavelength and bandwidth parameters. Results showed good agreement between spectral parameters estimated using onboard IFC and ground imaging data. The average difference between estimates obtained using the O(2) and H(2)O features and those obtained using the corresponding filter features amounted to about 0.3 nm (0.05 of a spectral pixel). A deviation from the nominal laboratory instrument spectral calibration and an altitude-dependent performance was additionally identified. The relatively good agreement between estimates obtained by the two approaches in similar spectral windows suggests they can be used in a complementary fashion: while the method relying on atmospheric features can be applied without the need for dedicated calibration acquisitions, the IFC allows assessment at user-selectable wavelength positions by custom filters as well as for the system on-ground.

In-flight spectral performance monitoring of the Airborne Prism Experiment.

Spectral performance of an airborne dispersive pushbroom imaging spectrometer cannot be assumed to be stable over a whole flight season given the environmental stresses present during flight. Spectral performance monitoring during flight is commonly accomplished by looking at selected absorption features present in the Sun, atmosphere, or ground, and their stability. The assessment of instrument performance in two different environments, e.g., laboratory and airborne, using precisely the same calibration reference, has not been possible so far. The Airborne Prism Experiment (APEX), an airborne dispersive pushbroom imaging spectrometer, uses an onboard in-flight characterization (IFC) facility, which makes it possible to monitor the sensor's performance in terms of spectral, radiometric, and geometric stability in flight and in the laboratory. We discuss in detail a new method for the monitoring of spectral instrument performance. The method relies on the monitoring of spectral shifts by comparing instrument-induced movements of absorption features on ground and in flight. Absorption lines originate from spectral filters, which intercept the full field of view (FOV) illuminated using an internal light source. A feature-fitting algorithm is used for the shift estimation based on Pearson's correlation coefficient. Environmental parameter monitoring, coregistered on board with the image and calibration data, revealed that differential pressure and temperature in the baffle compartment are the main driving parameters explaining the trend in spectral performance deviations in the time and the space (across-track) domains, respectively. The results presented in this paper show that the system in its current setup needs further improvements to reach a stable performance. Findings provided useful guidelines for the instrument revision currently under way. The main aim of the revision is the stabilization of the instrument for a range of temperature and pressure conditions to be encountered during operation.

APEX - the Hyperspectral ESA Airborne Prism Experiment.

The airborne ESA-APEX (Airborne Prism Experiment) hyperspectral mission simulator is described with its distinct specifications to provide high quality remote sensing data. The concept of an automatic calibration, performed in the Calibration Home Base (CHB) by using the Control Test Master (CTM), the In-Flight Calibration facility (IFC), quality flagging (QF) and specific processing in a dedicated Processing and Archiving Facility (PAF), and vicarious calibration experiments are presented. A preview on major applications and the corresponding development efforts to provide scientific data products up to level 2/3 to the user is presented for limnology, vegetation, aerosols, general classification routines and rapid mapping tasks. BRDF (Bidirectional Reflectance Distribution Function) issues are discussed and the spectral database SPECCHIO (Spectral Input/Output) introduced. The optical performance as well as the dedicated software utilities make APEX a state-of-the-art hyperspectral sensor, capable of (a) satisfying the needs of several research communities and (b) helping the understanding of the Earth's complex mechanisms.