ABSTRACT
Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ...ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO2 and/or O3. The plants were exposed either to ambient air (control), elevated CO2 (560 p.p.m.) elevated O3 (55 p.p.b.) or a mixture of elevated CO2 and O3 in a free air CO2 enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO2 and decreased by O3 at both control and elevated CO2. The relative stomatal limitation to photosynthesis (ls) was in both clones about 10% under control and elevated O3. Exposure to elevated CO2 + O3 in both clones and to elevated CO2 in clone 259, decreased ls even further – to about 5%. The corresponding changes in Rubisco content and the stability of Ci/Ca ratio suggest that the changes in photosynthesis in response to elevated CO2 and O3 were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O3 tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable Ci under the given treatment, that indicates close coupling between stomatal and mesophyll processes.
Ecosystem respiration (ER) was measured with the eddy covariance technique in 14 forest ecosystems in the Upper Great Lakes Region during the growing seasons of 2002 and 2003. The response of ER to ...soil temperature and moisture was analyzed using empirical models. On average, ER was higher in the intermediate and young than in the mature stands, and higher in hardwood than in conifer stands. The seasonal mean temperature-normalized respiration rate (
R
10) ranged from 1 to 3
μmol CO
2
m
−2
s
−1 and seasonal mean activation energy (
E
a) from 40 to 110
kJ
mol
−1. The variation in the residuals of temperature response function of ER was best explained by soil moisture content. ER showed higher temperature sensitivity (as indicated by lower
E
a) in the young than in the mature stands of coniferous forests, but not in the hardwood forests. The inclusion of soil moisture as an explicit driver of
R
10 explained an additional 8% (range 0–21%) of variability in ER. Significant moisture sensitivity of ER was detected in only 5 out of 20 site-years and it was associated with bimodal soil moisture distribution. Moisture sensitivity could partially be predicted from statistical moments kurtosis and interquartile range. The data implied greater moisture sensitivity with increasing stand age, possibly due to faster depletion of soil water supplies from a greater evaporative surface in the older stands. Additional limiting factors to ER were implicated.
Summary
1
The impacts of elevated atmospheric CO2 and/or O3 have been examined over 4 years using an open‐air exposure system in an aggrading northern temperate forest containing two different ...functional groups (the indeterminate, pioneer, O3‐sensitive species Trembling Aspen, Populus tremuloides and Paper Birch, Betula papyrifera, and the determinate, late successional, O3‐tolerant species Sugar Maple, Acer saccharum).
2
The responses to these interacting greenhouse gases have been remarkably consistent in pure Aspen stands and in mixed Aspen/Birch and Aspen/Maple stands, from leaf to ecosystem level, for O3‐tolerant as well as O3‐sensitive genotypes and across various trophic levels. These two gases act in opposing ways, and even at low concentrations (1·5 × ambient, with ambient averaging 34–36 nL L−1 during the summer daylight hours), O3 offsets or moderates the responses induced by elevated CO2.
3
After 3 years of exposure to 560 µmol mol−1 CO2, the above‐ground volume of Aspen stands was 40% above those grown at ambient CO2, and there was no indication of a diminishing growth trend. In contrast, O3 at 1·5 × ambient completely offset the growth enhancement by CO2, both for O3‐sensitive and O3‐tolerant clones. Implications of this finding for carbon sequestration, plantations to reduce excess CO2, and global models of forest productivity and climate change are presented.
Phenological events are integrative and sensitive indicators of ecosystem processes that respond to climate, water and nutrient availability, disturbance, and environmental change. The seasonality of ...ecosystem processes, including biogeochemical fluxes, can similarly be decomposed to identify key transition points and phase durations, which can be determined with high accuracy, and are specific to the processes of interest. As the seasonality of different processes differ, it can be argued that the interannual trends and responses to environmental forcings can be better described through the fluxes' own temporal characteristics than through correlation to traditional phenological events like bud break or leaf coloration. Here we present a global dataset of seasonality or phenological metrics calculated for gross primary productivity (GPP), ecosystem respiration (RE), latent heat (LE), and sensible heat (H) calculated for the FLUXNET2015 Dataset of about 200 sites and 1500 site years of data. The database includes metrics (i) on an absolute flux scale for comparisons with flux magnitudes and (ii) on a normalized scale for comparisons of change rates across different fluxes. Flux seasonality was characterized by fitting a single-pass double-logistic model to daily flux integrals, and the derivatives of the fitted time series were used to extract the phenological metrics marking key turning points, season lengths, and rates of change. Seasonal transition points could be determined with a 90 % confidence interval of 6-11 d for GPP, 8-14 d for RE, 10-15 d for LE, and 15-23 d for H. The phenology metrics derived from different partitioning methods diverged, at times significantly.
To determine whether elevated CO2 reduces or exacerbates the detrimental effects of O3 on aspen (Populus tremuloides Michx.). aspen clones 216 and 271 (O3 tolerant), and 259 (O3 sensitive) were ...exposed to ambient levels of CO2 and O3 or elevated levels of CO2, O3, or CO2 + O3 in the FACTS II (Aspen FACE) experiment, and physiological and molecular responses were measured and compared. Clone 259. the most O3-sensitive clone, showed the greatest amount of visible foliar symptoms as well as significant decreases in chlorophyll, carotenoid, starch, and ribulose-1, 5-bisphosphate carboxylase/oxygenase (Rubisco) concentrations and transcription levels for the Rubisco small subunit. Generally, the constitutive (basic) transcript levels for phenylalanine ammonialyase (PAL) and chalcone synthase (CHS) and the average antioxidant activities were lower for the ozone sensitive clone 259 as compared to the more tolerant 216 and 271 clones. A significant decrease in chlorophyll a, b and total (a + b) concentrations in CO2, O3, and CO2 + O3 plants was observed for all clones. Carotenoid concentrations were also significantly lower in all clones; however. CHS transcript levels were not significantly affected, suggesting a possible degradation of carotenoid pigments in O3-stressed plants. Antioxidant activities and PAL and 1-aminocyclopropane-l-carboxylic acid (ACC)-oxidase transcript levels showed a general increase in all O3 treated clones, while remaining low in CO2 and CO2 + O3 plants (although not all differences were significant). Our results suggest that the ascorbate-glutathione and phenylpropanoid pathways were activated under ozone stress and suppressed during exposure to elevated CO2. Although CO2 + O2 treatment resulted in a slight reduction of O3-induced leaf injury, it did not appear to ameliorate all of the harmful affects of O3 and, in fact. may have contributed to an increase in chloroplast damage in all three aspen clones.
In this article dedicated to Olevi Kull (June 22, 1955-January 31, 2007), we draw on his writings (in English and translated) to outline his thoughts on the relationship between scientists and ...science. We provide a brief synthesis of his most important work, give a short account of his career and, to bring the man into focus, share some personal stories of interactions with him. Kull considered that for a personal understanding to become scientific knowledge it must be explained convincingly based on theory and empirical support, and then taught to others in both spoken and written words. He saw the last step as the main distinction between learning and science. Olevi Kull's approach to science relied on two principles: first, linking theory and experiments in challenging settings, e.g., to test the generality of his ideas he often challenged them in multi-layered, mixed-species canopies. Second, he insisted on setting experiments to test assumptions used in quantitative analyses or in explaining an observed outcome; this, at times, led to falsification of commonly held ideas, thus enhancing ecophysiological understanding. After describing Kull's application of these principles, we give a brief synthesis of his most important work, in which he demonstrated through experimentation and modeling how the vertical distribution of leaves in canopies is consistent with the acclimation of the photosynthetic apparatus. We also review some of his findings on the interactive effects of carbon dioxide and ozone on canopy photosynthesis.
As a primary flux in the global water cycle, evapotranspiration (ET) connects hydrologic and biological processes and is directly affected by water and land management, land use change and climate ...variability. Satellite remote sensing provides an effective means for diagnosing ET patterns over heterogeneous landscapes; however, limitations on the spatial and temporal resolution of satellite data, combined with the effects of cloud contamination, constrain the amount of detail that a single satellite can provide. In this study, we describe an application of a multi-sensor ET data fusion system over a mixed forested/agricultural landscape in North Carolina, USA, during the growing season of 2013. The fusion system ingests ET estimates from the Two-Source Energy Balance Model (TSEB) applied to thermal infrared remote sensing retrievals of land surface temperature from multiple satellite platforms: hourly geostationary satellite data at 4 km resolution, daily 1 km imagery from the Moderate Resolution Imaging Spectroradiometer (MODIS) and biweekly Landsat thermal data sharpened to 30 m. These multiple ET data streams are combined using the Spatial and Temporal Adaptive Reflectance Fusion Model (STARFM) to estimate daily ET at 30 m resolution to investigate seasonal water use behavior at the level of individual forest stands and land cover patches. A new method, also exploiting the STARFM algorithm, is used to fill gaps in the Landsat ET retrievals due to cloud cover and/or the scan-line corrector (SLC) failure on Landsat 7. The retrieved daily ET time series agree well with observations at two AmeriFlux eddy covariance flux tower sites in a managed pine plantation within the modeling domain: US-NC2 located in a mid-rotation (20-year-old) loblolly pine stand and US-NC3 located in a recently clear-cut and replanted field site. Root mean square errors (RMSEs) for NC2 and NC3 were 0.99 and 1.02 mm day−1, respectively, with mean absolute errors of approximately 29 % at the daily time step, 12 % at the monthly time step and 0.7 % over the full study period at the two flux tower sites. Analyses of water use patterns over the plantation indicate increasing seasonal ET with stand age for young to mid-rotation stands up to 20 years, but little dependence on age for older stands. An accounting of consumptive water use by major land cover classes representative of the modeling domain is presented, as well as relative partitioning of ET between evaporation (E) and transpiration (T) components obtained with the TSEB. The study provides new insights about the effects of management and land use change on water yield over forested landscapes.
The evapotranspiration / potential evapotranspiration (AET / PET) ratio is traditionally termed as the crop coefficient (Kc) and has been generally used as ecosystem evaporative stress index. In the ...current hydrology literature, Kc has been widely used as a parameter to estimate crop water demand by water managers but has not been well examined for other types of ecosystems such as forests and other perennial vegetation. Understanding the seasonal dynamics of this variable for all ecosystems is important for projecting the ecohydrological responses to climate change and accurately quantifying water use at watershed to global scales. This study aimed at deriving monthly Kc for multiple vegetation cover types and understanding its environmental controls by analyzing the accumulated global eddy flux (FLUXNET) data. We examined monthly Kc data for seven vegetation covers, including open shrubland (OS), cropland (CRO), grassland (GRA), deciduous broad leaf forest (DBF), evergreen needle leaf forest (ENF), evergreen broad leaf forest (EBF), and mixed forest (MF), across 81 sites. We found that, except for evergreen forests (EBF and ENF), Kc values had large seasonal variation across all land covers. The spatial variability of Kc was well explained by latitude, suggesting site factors are a major control on Kc. Seasonally, Kc increased significantly with precipitation in the summer months, except in EBF. Moreover, leaf area index (LAI) significantly influenced monthly Kc in all land covers, except in EBF. During the peak growing season, forests had the highest Kc values, while croplands (CRO) had the lowest. We developed a series of multivariate linear monthly regression models for Kc by land cover type and season using LAI, site latitude, and monthly precipitation as independent variables. The Kc models are useful for understanding water stress in different ecosystems under climate change and variability as well as for estimating seasonal ET for large areas with mixed land covers.
A major concern for coastal freshwater wetland function and health is the effects of saltwater intrusion on greenhouse gas production from peat soils. Coastal freshwater forested wetlands are likely ...to experience increased hydroperiod with rising sea level, as well as saltwater intrusion. These potential changes to wetland hydrology may also alter forested wetland structure and lead to a transition from forest to shrub/marsh wetland ecosystems. Loss of forested wetlands is already evident by dying trees and dead standing trees (“ghost” forests) along the Atlantic coast of the US, which will result in significant alterations to plant carbon (C) inputs, particularly that of coarse woody debris, to soils. We investigated the effects of salinity and wood C inputs on soils collected from a coastal freshwater forested wetland in North Carolina, USA, and incubated in the laboratory with either freshwater or saltwater (2.5 or 5.0 ppt) and with or without the additions of wood. Saltwater additions at 2.5 and 5.0 ppt reduced CO2 production by 41 % and 37 %, respectively, compared to freshwater. Methane production was reduced by 98 % (wood-free incubations) and by 75 %–87 % (wood-amended incubations) in saltwater treatments compared to the freshwater plus wood treatment. Additions of wood also resulted in lower CH4 production from the freshwater treatment and higher CH4 production from saltwater treatments compared to wood-free incubations. The δ13CH4-C isotopic signature suggested that, in wood-free incubations, CH4 produced from the freshwater treatment originated primarily from the acetoclastic pathway, while CH4 produced from the saltwater treatments originated primarily from the hydrogenotrophic pathway. These results suggest that saltwater intrusion into coastal freshwater forested wetlands will reduce CH4 production, but long-term changes in C dynamics will likely depend on how changes in wetland vegetation and microbial function influence C cycling in peat soils.
We review 15 techniques for estimating missing values of net ecosystem CO
2 exchange (NEE) in eddy covariance time series and evaluate their performance for different artificial gap scenarios based ...on a set of 10 benchmark datasets from six forested sites in Europe.
The goal of gap filling is the reproduction of the NEE time series and hence this present work focuses on estimating missing NEE values, not on editing or the removal of suspect values in these time series due to systematic errors in the measurements (e.g., nighttime flux, advection). The gap filling was examined by generating 50 secondary datasets with artificial gaps (ranging in length from single half-hours to 12 consecutive days) for each benchmark dataset and evaluating the performance with a variety of statistical metrics. The performance of the gap filling varied among sites and depended on the level of aggregation (native half-hourly time step versus daily), long gaps were more difficult to fill than short gaps, and differences among the techniques were more pronounced during the day than at night.
The non-linear regression techniques (NLRs), the look-up table (LUT), marginal distribution sampling (MDS), and the semi-parametric model (SPM) generally showed good overall performance. The artificial neural network based techniques (ANNs) were generally, if only slightly, superior to the other techniques. The simple interpolation technique of mean diurnal variation (MDV) showed a moderate but consistent performance. Several sophisticated techniques, the dual unscented Kalman filter (UKF), the multiple imputation method (MIM), the terrestrial biosphere model (BETHY), but also one of the ANNs and one of the NLRs showed high biases which resulted in a low reliability of the annual sums, indicating that additional development might be needed. An uncertainty analysis comparing the estimated random error in the 10 benchmark datasets with the artificial gap residuals suggested that the techniques are already at or very close to the noise limit of the measurements. Based on the techniques and site data examined here, the effect of gap filling on the annual sums of NEE is modest, with most techniques falling within a range of ±25
g
C
m
−2
year
−1.
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