Ecotrons: Powerful and versatile ecosystem analysers for ecology, agronomy and environmental science.

Ecosystems integrity and services are threatened by anthropogenic global changes. Mitigating and adapting to these changes require knowledge of ecosystem functioning in the expected novel environments, informed in large part through experimentation and modelling. This paper describes 13 advanced controlled environment facilities for experimental ecosystem studies, herein termed ecotrons, open to the international community. Ecotrons enable simulation of a wide range of natural environmental conditions in replicated and independent experimental units while measuring various ecosystem processes. This capacity to realistically control ecosystem environments is used to emulate a variety of climatic scenarios and soil conditions, in natural sunlight or through broad-spectrum lighting. The use of large ecosystem samples, intact or reconstructed, minimizes border effects and increases biological and physical complexity. Measurements of concentrations of greenhouse trace gases as well as their net exchange between the ecosystem and the atmosphere are performed in most ecotrons, often quasi continuously. The flow of matter is often tracked with the use of stable isotope tracers of carbon and other elements. Equipment is available for measurements of soil water status as well as root and canopy growth. The experiments ran so far emphasize the diversity of the hosted research. Half of them concern global changes, often with a manipulation of more than one driver. About a quarter deal with the impact of biodiversity loss on ecosystem functioning and one quarter with ecosystem or plant physiology. We discuss how the methodology for environmental simulation and process measurements, especially in soil, can be improved and stress the need to establish stronger links with modelling in future projects. These developments will enable further improvements in mechanistic understanding and predictive capacity of ecotron research which will play, in complementarity with field experimentation and monitoring, a crucial role in exploring the ecosystem consequences of environmental changes.

Integration of ecosystem science into radioecology: A consensus perspective.

In the Fall of 2016 a workshop was held which brought together over 50 scientists from the ecological and radiological fields to discuss feasibility and challenges of reintegrating ecosystem science into radioecology. There is a growing desire to incorporate attributes of ecosystem science into radiological risk assessment and radioecological research more generally, fueled by recent advances in quantification of emergent ecosystem attributes and the desire to accurately reflect impacts of radiological stressors upon ecosystem function. This paper is a synthesis of the discussions and consensus of the workshop participant's responses to three primary questions, which were: 1) How can ecosystem science support radiological risk assessment? 2) What ecosystem level endpoints potentially could be used for radiological risk assessment? and 3) What inference strategies and associated methods would be most appropriate to assess the effects of radionuclides on ecosystem structure and function? The consensus of the participants was that ecosystem science can and should support radiological risk assessment through the incorporation of quantitative metrics that reflect ecosystem functions which are sensitive to radiological contaminants. The participants also agreed that many such endpoints exit or are thought to exit and while many are used in ecological risk assessment currently, additional data need to be collected that link the causal mechanisms of radiological exposure to these endpoints. Finally, the participants agreed that radiological risk assessments must be designed and informed by rigorous statistical frameworks capable of revealing the causal inference tying radiological exposure to the endpoints selected for measurement.

Resource Concentration Modulates the Fate of Dissimilated Nitrogen in a Dual-Pathway Actinobacterium.

Respiratory ammonification and denitrification are two evolutionarily unrelated dissimilatory nitrogen (N) processes central to the global N cycle, the activity of which is thought to be controlled by carbon (C) to nitrate (NO3 -) ratio. Here we find that Intrasporangium calvum C5, a novel dual-pathway denitrifier/respiratory ammonifier, disproportionately utilizes ammonification rather than denitrification when grown under low C concentrations, even at low C:NO3 - ratios. This finding is in conflict with the paradigm that high C:NO3 - ratios promote ammonification and low C:NO3 - ratios promote denitrification. We find that the protein atomic composition for denitrification modules (NirK) are significantly cost minimized for C and N compared to ammonification modules (NrfA), indicating that limitation for C and N is a major evolutionary selective pressure imprinted in the architecture of these proteins. The evolutionary precedent for these findings suggests ecological importance for microbial activity as evidenced by higher growth rates when I. calvum grows predominantly using its ammonification pathway and by assimilating its end-product (ammonium) for growth under ammonium-free conditions. Genomic analysis of I. calvum further reveals a versatile ecophysiology to cope with nutrient stress and redox conditions. Metabolite and transcriptional profiles during growth indicate that enzyme modules, NrfAH and NirK, are not constitutively expressed but rather induced by nitrite production via NarG. Mechanistically, our results suggest that pathway selection is driven by intracellular redox potential (redox poise), which may be lowered when resource concentrations are low, thereby decreasing catalytic activity of upstream electron transport steps (i.e., the bc1 complex) needed for denitrification enzymes. Our work advances our understanding of the biogeochemical flexibility of N-cycling organisms, pathway evolution, and ecological food-webs.

Biotic degradation at night, abiotic degradation at day: positive feedbacks on litter decomposition in drylands.

The arid and semi-arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO2 ) uptake, which depends largely on plant litter decomposition and the subsequent release of CO2 back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO2 efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO2 efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long-term trend of global carbon sequestration.

Plant species effects on soil nutrients and chemistry in arid ecological zones.

The presence of vegetation strongly influences ecosystem function by controlling the distribution and transformation of nutrients across the landscape. The magnitude of vegetation effects on soil chemistry is largely dependent on the plant species and the background soil chemical properties of the site, but has not been well quantified along vegetation transects in the Great Basin. We studied the effects of plant canopy cover on soil chemistry within five different ecological zones, subalpine, montane, pinyon-juniper, sage/Mojave transition, and desert shrub, in the Great Basin of Nevada all with similar underlying geology. Although plant species differed in their effects on soil chemistry, the desert shrubs Sarcobatus vermiculatus, Atriplex spp., Coleogyne ramosissima, and Larrea tridentata typically exerted the most influence on soil chemistry, especially amounts of K(+) and total nitrogen, beneath their canopies. However, the extent to which vegetation affected soil nutrient status in any given location was not only highly dependent on the species present, and presumably the nutrient requirements and cycling patterns of the plant species, but also on the background soil characteristics (e.g., parent material, weathering rates, leaching) where plant species occurred. The results of this study indicate that the presence or absence of a plant species, especially desert shrubs, could significantly alter soil chemistry and subsequently ecosystem biogeochemistry and function.

Earthworm effects on native grassland root system dynamics under natural and increased rainfall.

Earthworms (EWs) can modify soil structure and nutrient availability, and hence alter conditions for plant growth through their burrowing and casting activities. However, few studies have specifically quantified EW effects by experimentally manipulating earthworm densities (EWDs). In an earlier field study in native grassland ecosystems exposed to ambient and experimentally elevated rainfall (+280 mm year(-1), projected under some climate change scenarios), we found no effects of EWDs (37, 114, 169 EW m(-2)) and corresponding EW activity on aboveground net primary productivity (ANPP), even though soil nutrient availability likely increased with increasing EWDs. The lack of effects of EWDs on ANPP suggested that EWs may have adversely affected root systems in that study in some way. The objective of the present study was to quantify responses of root length density (RLD), using data collected from the same grassland plots during the earlier study. RLDs were highest in plots with low EWDs and decreased in plots with higher EWDs. Elevated rainfall primarily increased RLDs in the low EWD treatment (by almost +40%). Reductions in RLDs resulting from increased EWDs did not affect ANPP. Our results indicate that elevating EWDs above ambient levels may limit root growth through large increases in soil bioturbation, but concurrent increases in cast production and nutrient availability may compensate for the suppression of root nutrient absorbing surface area leaving ANPP unchanged, but with shifts in growth (biomass) allocation toward shoots. Similarly, reductions in EWDs appeared to promote higher RLDs that increased soil nutrient foraging in soil with lower amounts of nutrients because of reduced casting activity. Amplified responses observed when rainfall during the growing season was increased suggest that EWDs may mainly affect RLDs and above- vs. belowground growth (biomass) allocation under climate changes that include more frequent wetter-than-average growing seasons.

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2-year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha(-1)  yr(-1) . Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.

Loss of plant biodiversity eliminates stimulatory effect of elevated CO2 on earthworm activity in grasslands.

Earthworms are among the world's most important ecosystem engineers because of their effects on soil fertility and plant productivity. Their dependence on plants for carbon, however, means that any changes in plant community structure or function caused by rising atmospheric CO2 or loss of plant species diversity could affect earthworm activity, which may feed back on plant communities. Production of surface casts measured during three consecutive years in field experimental plots (n = 24, 1.2 m(2)) planted with local calcareous grassland species that varied in plant species richness (diversity levels: high, 31 species; medium, 12; low, 5) and were exposed to ambient (356 µl CO2 l(-1)) or elevated (600 µl CO2 l(-1)) CO2 was only consistently stimulated in high diversity plots exposed to elevated CO2 (+120 %, 31 spp: 603 ± 52 under ambient CO2 vs. 1,325 ± 204 g cast dwt. m(-2) year(-1) under elevated CO2 in 1996; +77 %, 940 ± 44 vs. 1,663 ± 204 g cast dwt. m(-2) year(-1) in 1998). Reductions in plant diversity had little effect on cast production in ecosystems maintained at ambient CO2, but the stimulatory effect of elevated CO2 on cast production disappeared when plant species diversity was decreased to 12 and 5 species. High diversity plots were also the only communities that included plant species that an earlier field study showed to be among the most responsive to elevated CO2 and to be most preferred by earthworms to deposit casts near. Further, the +87 % CO2-induced increase in cast production measured over the 3 years corresponded to a parallel increase in cumulative total nitrogen of 5.7 g N m(-2) and would help explain the large stimulation of aboveground plant biomass production observed in high-diversity communities under elevated CO2. The results of this study demonstrate how the loss of plant species from communities can alter responses of major soil heterotrophs and consequently ecosystem biogeochemistry.

Concurrent and lagged impacts of an anomalously warm year on autotrophic and heterotrophic components of soil respiration: a deconvolution analysis.

*Partitioning soil respiration into autotrophic (R(A)) and heterotrophic (R(H)) components is critical for understanding their differential responses to climate warming. *Here, we used a deconvolution analysis to partition soil respiration in a pulse warming experiment. We first conducted a sensitivity analysis to determine which parameters can be identified by soil respiration data. A Markov chain Monte Carlo technique was then used to optimize those identifiable parameters in a terrestrial ecosystem model. Finally, the optimized parameters were employed to quantify R(A) and R(H) in a forward analysis. *Our results displayed that more than one-half of parameters were constrained by daily soil respiration data. The optimized model simulation showed that warming stimulated R(H) and had little effect on R(A) in the first 2 months, but decreased both R(H) and R(A) during the remainder of the treatment and post-treatment years. Clipping of above-ground biomass stimulated the warming effect on R(H) but not on R(A). Overall, warming decreased R(A) and R(H) significantly, by 28.9% and 24.9%, respectively, during the treatment year and by 27.3% and 33.3%, respectively, during the post-treatment year, largely as a result of decreased canopy greenness and biomass. *Lagged effects of climate anomalies on soil respiration and its components are important in assessing terrestrial carbon cycle feedbacks to climate warming.

On the consequences of the energy imbalance for calculating surface conductance to water vapour.

The Penman-Monteith combination equation, which is most frequently used to derive the surface conductance to water vapour (Gs), implicitly assumes the energy balance to be closed. Any energy imbalance (positive or negative) will thus affect the calculated Gs. Using eddy covariance energy flux data from a temperate grassland and a desert shrub ecosystem we explored five possible approaches of closing the energy imbalance and show that calculated Gs may differ considerably between these five approaches depending on the relative magnitudes of sensible and latent heat fluxes, and the magnitude and sign of the energy imbalance. Based on our limited understanding of the nature of the energy imbalance, we tend to favour an approach which preserves the Bowen-ratio and closes the energy balance on a larger time scale.

Prolonged suppression of ecosystem carbon dioxide uptake after an anomalously warm year.

Terrestrial ecosystems control carbon dioxide fluxes to and from the atmosphere through photosynthesis and respiration, a balance between net primary productivity and heterotrophic respiration, that determines whether an ecosystem is sequestering carbon or releasing it to the atmosphere. Global and site-specific data sets have demonstrated that climate and climate variability influence biogeochemical processes that determine net ecosystem carbon dioxide exchange (NEE) at multiple timescales. Experimental data necessary to quantify impacts of a single climate variable, such as temperature anomalies, on NEE and carbon sequestration of ecosystems at interannual timescales have been lacking. This derives from an inability of field studies to avoid the confounding effects of natural intra-annual and interannual variability in temperature and precipitation. Here we present results from a four-year study using replicate 12,000-kg intact tallgrass prairie monoliths located in four 184-m(3) enclosed lysimeters. We exposed 6 of 12 monoliths to an anomalously warm year in the second year of the study and continuously quantified rates of ecosystem processes, including NEE. We find that warming decreases NEE in both the extreme year and the following year by inducing drought that suppresses net primary productivity in the extreme year and by stimulating heterotrophic respiration of soil biota in the subsequent year. Our data indicate that two years are required for NEE in the previously warmed experimental ecosystems to recover to levels measured in the control ecosystems. This time lag caused net ecosystem carbon sequestration in previously warmed ecosystems to be decreased threefold over the study period, compared with control ecosystems. Our findings suggest that more frequent anomalously warm years, a possible consequence of increasing anthropogenic carbon dioxide levels, may lead to a sustained decrease in carbon dioxide uptake by terrestrial ecosystems.

Atmospheric mercury exchange with a tallgrass prairie ecosystem housed in mesocosms.

This study focused on characterizing air-surface mercury Hg exchange for individual surfaces (soil, litter-covered soil and plant shoots) and ecosystem-level flux associated with tallgrass prairie ecosystems housed inside large mesocosms over three years. The major objectives of this project were to determine if individual surface fluxes could be combined to predict ecosystem-level exchange and if this low-Hg containing ecosystem was a net source or sink for atmospheric Hg. Data collected in the field were used to validate fluxes obtained in the mesocosm setting. Because of the controlled experimental design and ease of access to the mesocosms, data collected allowed for assessment of factors controlling flux and comparison of models developed for soil Hg flux versus environmental conditions at different temporal resolution (hourly, daily and monthly). Evaluation of hourly data showed that relationships between soil Hg flux and environmental conditions changed over time, and that there were interactions between parameters controlling exchange. Data analyses demonstrated that to estimate soil flux over broad temporal scales (e.g. annual flux) coarse-resolution data (monthly averages) are needed. Plant foliage was a sink for atmospheric Hg with uptake influenced by plant functional type and age. Individual system component fluxes (bare soil and plant) could not be directly combined to predict the measured whole system flux (soil, litter and plant). Emissions of Hg from vegetated and litter-covered soil were lower than fluxes from adjacent bare soil and the difference between the two was seasonally dependent and greatest when canopy coverage was greatest. Thus, an index of plant canopy development (canopy greenness) was used to model Hg flux from vegetated soil. Accounting for ecosystem Hg inputs (precipitation, direct plant uptake of atmospheric Hg) and modeled net exchange between litter-and-plant covered soils, the tallgrass prairie was found to be a net annual sink of atmospheric Hg.

Divergence of reproductive phenology under climate warming.

Because the flowering and fruiting phenology of plants is sensitive to environmental cues such as temperature and moisture, climate change is likely to alter community-level patterns of reproductive phenology. Here we report a previously unreported phenomenon: experimental warming advanced flowering and fruiting phenology for species that began to flower before the peak of summer heat but delayed reproduction in species that started flowering after the peak temperature in a tallgrass prairie in North America. The warming-induced divergence of flowering and fruiting toward the two ends of the growing season resulted in a gap in the staggered progression of flowering and fruiting in the community during the middle of the season. A double precipitation treatment did not significantly affect flowering and fruiting phenology. Variation among species in the direction and magnitude of their response to warming caused compression and expansion of the reproductive periods of different species, changed the amount of overlap between the reproductive phases, and created possibilities for an altered selective environment to reshape communities in a future warmed world.

Net carbon exchange and evapotranspiration in postfire and intact sagebrush communities in the Great Basin.

Invasion of non-native annuals across the Intermountain West is causing a widespread transition from perennial sagebrush communities to fire-prone annual herbaceous communities and grasslands. To determine how this invasion affects ecosystem function, carbon and water fluxes were quantified in three, paired sagebrush and adjacent postfire communities in the northern Great Basin using a 1-m3 gas exchange chamber. Most of the plant cover in the postfire communities was invasive species including Bromus tectorum L., Agropyron cristatum (L.) Gaertn and Sisymbrium altissimum L. Instantaneous morning net carbon exchange (NCE) and evapotranspiration (ET) in native shrub plots were greater than either intershrub or postfire plots. Native sagebrush communities were net carbon sinks (mean NCE 0.2-4.3 micromol m-2 s-1) throughout the growing season. The magnitude and seasonal variation of NCE in the postfire communities were controlled by the dominant species and availability of soil moisture. Net C exchange in postfire communities dominated by perennial bunchgrasses was similar to sagebrush. However, communities dominated by annuals (cheatgrass and mustard) had significantly lower NCE than sagebrush and became net sources of carbon to the atmosphere (NCE declined to -0.5 micromol m-2 s-1) with increased severity of the summer drought. Differences in the patterns of ET led to lower surface soil moisture content and increased soil temperatures during summer in the cheatgrass-dominated community compared to the adjacent sagebrush community. Intensive measurements at one site revealed that temporal and spatial patterns of NCE and ET were correlated most closely with changes in leaf area in each community. By altering the patterns of carbon and water exchange, conversion of native sagebrush to postfire invasive communities may disrupt surface-atmosphere exchange and degrade the carbon storage capacity of these systems.

Influence of elevated CO2 on canopy development and red:far-red ratios in two-storied stands ofRicinus communis.

Vertical structure of plant stands and canopies may change under conditions of elevated CO2 due to differential responses of overstory and understory plants or plant parts. In the long term, seedling recruitment, competition, and thus population or community structure may be affected. Aside from the possible differential direct effects of elevated CO2 on photosynthesis and growth, both the quantity and quality of the light below the overstory canopy could be indirectly affected by CO2-induced changes in overstory leaf area index (LAI) and/or changes in overstory leaf quality. In order to explore such possible interactions, we compared canopy leaf area development, canopy light extinction and the quality of light beneath overstory leaves of two-storied monospecific stands ofRicinus communis exposed to ambient (340 µl l-1) and elevated (610 µl l-1) CO2. Plants in each stand were grown in a common soil as closed "artificial ecosystems" with a ground area of 6.7 m2. LAI of overstory plants in all ecosystems more than doubled during the experiment but was not different between CO2 treatments at the end. As a consequence, extinction of photosynthetically active radiation (PAR) was also not altered. However, under elevated CO2 the red to far-red ratio (R:FR) measured beneath overstory leaves was 10% lower than in ecosystems treated with ambient CO2. This reduction was associated with increased thickness of palisade layers of overstory leaves and appears to be a plausible explanation for the specific enhancement of stem elongation of understory plants (without a corresponding biomass response) under elevated CO2. CO2 enrichment led to increased biomass of overstory plants (mainly stem biomass) but had no effect on understory biomass. The results of this study raise the possibility of an important indirect effect of elevated CO2 at the stand-level. We suggest that, under elevated CO2, reductions in the R:FR ratio beneath overstory canopies may affect understory plant development independently of the effects of PAR extinction.