When things get MESI: The Manipulation Experiments Synthesis Initiative-A coordinated effort to synthesize terrestrial global change experiments.

Responses of the terrestrial biosphere to rapidly changing environmental conditions are a major source of uncertainty in climate projections. In an effort to reduce this uncertainty, a wide range of global change experiments have been conducted that mimic future conditions in terrestrial ecosystems, manipulating CO2 , temperature, and nutrient and water availability. Syntheses of results across experiments provide a more general sense of ecosystem responses to global change, and help to discern the influence of background conditions such as climate and vegetation type in determining global change responses. Several independent syntheses of published data have yielded distinct databases for specific objectives. Such parallel, uncoordinated initiatives carry the risk of producing redundant data collection efforts and have led to contrasting outcomes without clarifying the underlying reason for divergence. These problems could be avoided by creating a publicly available, updatable, curated database. Here, we report on a global effort to collect and curate 57,089 treatment responses across 3644 manipulation experiments at 1145 sites, simulating elevated CO2 , warming, nutrient addition, and precipitation changes. In the resulting Manipulation Experiments Synthesis Initiative (MESI) database, effects of experimental global change drivers on carbon and nutrient cycles are included, as well as ancillary data such as background climate, vegetation type, treatment magnitude, duration, and, unique to our database, measured soil properties. Our analysis of the database indicates that most experiments are short term (one or few growing seasons), conducted in the USA, Europe, or China, and that the most abundantly reported variable is aboveground biomass. We provide the most comprehensive multifactor global change database to date, enabling the research community to tackle open research questions, vital to global policymaking. The MESI database, freely accessible at doi.org/10.5281/zenodo.7153253, opens new avenues for model evaluation and synthesis-based understanding of how global change affects terrestrial biomes. We welcome contributions to the database on GitHub.

Accelerated sea-level rise is suppressing CO2 stimulation of tidal marsh productivity: A 33-year study.

Accelerating relative sea-level rise (RSLR) is threatening coastal wetlands. However, rising CO2 concentrations may also stimulate carbon sequestration and vertical accretion, counterbalancing RSLR. A coastal wetland dominated by a C3 plant species was exposed to ambient and elevated levels of CO2 in situ from 1987 to 2019 during which time ambient CO2 concentration increased 18% and sea level rose 23 cm. Plant production did not increase in response to gradually rising ambient CO2 concentration during this period. Elevated CO2 increased shoot production relative to ambient CO2 for the first two decades, but from 2005 to 2019, elevated CO2 stimulation of production was diminished. The decline coincided with increases in relative sea level above a threshold that hindered root productivity. While elevated CO2 stimulation of elevation gain has the potential to moderate the negative impacts of RSLR on tidal wetland productivity, benefits for coastal wetland resilience will diminish in the long term as rates of RSLR accelerate.

Do trade-offs govern plant species' responses to different global change treatments?

Plants are subject to trade-offs among growth strategies such that adaptations for optimal growth in one condition can preclude optimal growth in another. Thus, we predicted that a plant species that responds positively to one global change treatment would be less likely than average to respond positively to another treatment, particularly for pairs of treatments that favor distinct traits. We examined plant species' abundances in 39 global change experiments manipulating two or more of the following: CO2 , nitrogen, phosphorus, water, temperature, or disturbance. Overall, the directional response of a species to one treatment was 13% more likely than expected to oppose its response to a another single-factor treatment. This tendency was detectable across the global data set, but held little predictive power for individual treatment combinations or within individual experiments. Although trade-offs in the ability to respond to different global change treatments exert discernible global effects, other forces obscure their influence in local communities.

Determinants of community compositional change are equally affected by global change.

Global change is impacting plant community composition, but the mechanisms underlying these changes are unclear. Using a dataset of 58 global change experiments, we tested the five fundamental mechanisms of community change: changes in evenness and richness, reordering, species gains and losses. We found 71% of communities were impacted by global change treatments, and 88% of communities that were exposed to two or more global change drivers were impacted. Further, all mechanisms of change were equally likely to be affected by global change treatments-species losses and changes in richness were just as common as species gains and reordering. We also found no evidence of a progression of community changes, for example, reordering and changes in evenness did not precede species gains and losses. We demonstrate that all processes underlying plant community composition changes are equally affected by treatments and often occur simultaneously, necessitating a wholistic approach to quantifying community changes.

Mangrove growth response to experimental warming is greatest near the range limit in northeast Florida.

Shrubs are invading into grasslands around the world, but we don't yet know how these shrubs will fare in a warmer future. In ecotonal coastal wetland ecosystems, woody mangroves are encroaching into herbaceous salt marshes owing to changes in temperature, precipitation, and sediment dynamics. Increasing mangrove biomass in wetlands often increases carbon storage, which is high in these productive ecosystems, but little is known about how mangrove growth will change in response to warming. To address this knowledge gap, we deployed warming experiments at three coastal wetland sites along a latitudinal gradient in northeast Florida where Avicennia germinans, black mangroves, are encroaching into salt marshes. We achieved air temperature warming (+1.6°C during the day) at all three sites and measured stem elongation, canopy height and area changes, and leaf and node number. After 2 yr of warming, we found that mangrove growth rate in height increased due to warming. Warming increased stem elongation by 130% over unwarmed control plots after 1 yr at the northern site. Mangrove growth in canopy area did not respond to warming. Site differences in growth rate were pronounced, and mangrove growth in both height and area were lowest at the northern site, despite greater impacts of warming at that site. We also found that area-based relative growth rate was five times higher across all treatments than height-based relative growth rate, indicating that mangroves are growing wider rather than taller in these ecotonal environments. Our findings indicate that the growth effect of experimental warming depends on site characteristics and growth parameter measured. We also propose that differential mangrove growth across the three sites may be driven by biotic factors such as the identity of the salt marsh species into which mangroves are encroaching. Our results suggest that, as seen in other ecosystems, wetland plants may respond most strongly to warming at their poleward range edge.

Plant species determine tidal wetland methane response to sea level rise.

Blue carbon (C) ecosystems are among the most effective C sinks of the biosphere, but methane (CH4) emissions can offset their climate cooling effect. Drivers of CH4 emissions from blue C ecosystems and effects of global change are poorly understood. Here we test for the effects of sea level rise (SLR) and its interactions with elevated atmospheric CO2, eutrophication, and plant community composition on CH4 emissions from an estuarine tidal wetland. Changes in CH4 emissions with SLR are primarily mediated by shifts in plant community composition and associated plant traits that determine both the direction and magnitude of SLR effects on CH4 emissions. We furthermore show strong stimulation of CH4 emissions by elevated atmospheric CO2, whereas effects of eutrophication are not significant. Overall, our findings demonstrate a high sensitivity of CH4 emissions to global change with important implications for modeling greenhouse-gas dynamics of blue C ecosystems.

Representing the function and sensitivity of coastal interfaces in Earth system models.

Between the land and ocean, diverse coastal ecosystems transform, store, and transport material. Across these interfaces, the dynamic exchange of energy and matter is driven by hydrological and hydrodynamic processes such as river and groundwater discharge, tides, waves, and storms. These dynamics regulate ecosystem functions and Earth's climate, yet global models lack representation of coastal processes and related feedbacks, impeding their predictions of coastal and global responses to change. Here, we assess existing coastal monitoring networks and regional models, existing challenges in these efforts, and recommend a path towards development of global models that more robustly reflect the coastal interface.

Exploring the oxygen sensitivity of wetland soil carbon mineralization.

Soil oxygen availability may influence blue carbon, which is carbon stored in coastal wetlands, by controlling the decomposition of soil organic matter. We are beginning to quantify soil oxygen availability in wetlands, but we lack a precise understanding of how oxygen controls soil carbon dynamics. In this paper, we synthesize existing data from oxic and anoxic wetland soil incubations to determine how oxygen controls carbon mineralization. We define the oxygen sensitivity of carbon mineralization as the ratio of carbon mineralization rate in oxic soil to this rate in anoxic soil, such that higher values of this ratio indicate greater sensitivity of carbon mineralization to oxygen. The estimates of oxygen sensitivity we derived from existing literature show a wide range of ratios, from 0.8 to 33, across wetlands. We then report oxygen sensitivities from an experimental mesocosm we developed to manipulate soil oxygen status in realistic soils. The variation in oxygen sensitivity we uncover from this systematic review and experiment indicates that Earth system models may misrepresent the oxygen sensitivity of carbon mineralization, and how it varies with context, in wetland soils. We suggest that altered soil oxygen availability could be an important driver of future blue carbon storage in coastal wetlands.

Globally consistent influences of seasonal precipitation limit grassland biomass response to elevated CO2.

Rising atmospheric carbon dioxide concentration should stimulate biomass production directly via biochemical stimulation of carbon assimilation, and indirectly via water savings caused by increased plant water-use efficiency. Because of these water savings, the CO2 fertilization effect (CFE) should be stronger at drier sites, yet large differences among experiments in grassland biomass response to elevated CO2 appear to be unrelated to annual precipitation, preventing useful generalizations. Here, we show that, as predicted, the impact of elevated CO2 on biomass production in 19 globally distributed temperate grassland experiments reduces as mean precipitation in seasons other than spring increases, but that it rises unexpectedly as mean spring precipitation increases. Moreover, because sites with high spring precipitation also tend to have high precipitation at other times, these effects of spring and non-spring precipitation on the CO2 response offset each other, constraining the response of ecosystem productivity to rising CO2. This explains why previous analyses were unable to discern a reliable trend between site dryness and the CFE. Thus, the CFE in temperate grasslands worldwide will be constrained by their natural rainfall seasonality such that the stimulation of biomass by rising CO2 could be substantially less than anticipated.

Ambient changes exceed treatment effects on plant species abundance in global change experiments.

The responses of species to environmental changes will determine future community composition and ecosystem function. Many syntheses of global change experiments examine the magnitude of treatment effect sizes, but we lack an understanding of how plant responses to treatments compare to ongoing changes in the unmanipulated (ambient or background) system. We used a database of long-term global change studies manipulating CO2 , nutrients, water, and temperature to answer three questions: (a) How do changes in plant species abundance in ambient plots relate to those in treated plots? (b) How does the magnitude of ambient change in species-level abundance over time relate to responsiveness to global change treatments? (c) Does the direction of species-level responses to global change treatments differ from the direction of ambient change? We estimated temporal trends in plant abundance for 791 plant species in ambient and treated plots across 16 long-term global change experiments yielding 2,116 experiment-species-treatment combinations. Surprisingly, for most species (57%) the magnitude of ambient change was greater than the magnitude of treatment effects. However, the direction of ambient change, whether a species was increasing or decreasing in abundance under ambient conditions, had no bearing on the direction of treatment effects. Although ambient communities are inherently dynamic, there is now widespread evidence that anthropogenic drivers are directionally altering plant communities in many ecosystems. Thus, global change treatment effects must be interpreted in the context of plant species trajectories that are likely driven by ongoing environmental changes.

Partitioning direct and indirect effects reveals the response of water-limited ecosystems to elevated CO2.

Increasing concentrations of atmospheric carbon dioxide are expected to affect carbon assimilation and evapotranspiration (ET), ultimately driving changes in plant growth, hydrology, and the global carbon balance. Direct leaf biochemical effects have been widely investigated, whereas indirect effects, although documented, elude explicit quantification in experiments. Here, we used a mechanistic model to investigate the relative contributions of direct (through carbon assimilation) and indirect (via soil moisture savings due to stomatal closure, and changes in leaf area index) effects of elevated CO2 across a variety of ecosystems. We specifically determined which ecosystems and climatic conditions maximize the indirect effects of elevated CO2 The simulations suggest that the indirect effects of elevated CO2 on net primary productivity are large and variable, ranging from less than 10% to more than 100% of the size of direct effects. For ET, indirect effects were, on average, 65% of the size of direct effects. Indirect effects tended to be considerably larger in water-limited ecosystems. As a consequence, the total CO2 effect had a significant, inverse relationship with the wetness index and was directly related to vapor pressure deficit. These results have major implications for our understanding of the CO2 response of ecosystems and for global projections of CO2 fertilization, because, although direct effects are typically understood and easily reproducible in models, simulations of indirect effects are far more challenging and difficult to constrain. Our findings also provide an explanation for the discrepancies between experiments in the total CO2 effect on net primary productivity.

Above vs. belowground plant biomass along a barrier island: Implications for dune stabilization.

Coastal regions are inherently and increasingly vulnerable and geomorphologically unstable, yet are invaluable economic and residential hubs. Dunes are dynamic buffers to erosion and the most natural, economical, and effective defense for coastal communities. Vegetation is integral to dune structure as it facilitates accretion and stabilization. Differences in the vegetation and root density likely translate to variability in coastal erosion prevention, but this notion has been largely unconsidered. We directly compared stabilizing factors, depth and density, of the root systems of two dominant mid-Atlantic dune plant species, native American beach grass (Ammophila breviligulata) and invasive Asiatic sand sedge (Carex kobomugi). Despite high plant density, C. kobomugi is targeted for removal in restoration efforts as its roots are assumed to provide less effective stabilization than A. breviligulata. We collected 30 cores and hand dug 14 A. breviligulata ramets at Island Beach State Park, New Jersey to examine biomass, root:shoot ratios, and root density. C. kobomugi had a more extensive root system with a root:shoot ratio of 11.36:1 compared to 1.62:1 for A. breviligulata. Similarly, cores 60 cm deep and 7.6 cm wide were sufficient to attain fully intact A. breviligulata roots, which did not extend deeper than 40 cm, but insufficient for C. kobomugi roots which extended beyond the sampling system vertically and horizontally. Scaling these findings to m(-2), aboveground biomass is relatively equal, but C. kobomugi had over 700% more root mass m(-2) than A. breviligulata. These results have strong implications for dune management. The root system of C. kobomugi may be better adapted to stabilize dunes and thus protect coastal areas during small and large-scale perturbations than previously supposed. This is a unique situation whereby the creation of monocultures will hyperstabilize dunes and make them more resistant to erosion at the cost of reduced biodiversity within the framework of resiliency.

Elevated CO2 promotes long-term nitrogen accumulation only in combination with nitrogen addition.

Biogeochemical models that incorporate nitrogen (N) limitation indicate that N availability will control the magnitude of ecosystem carbon uptake in response to rising CO2 . Some models, however, suggest that elevated CO2 may promote ecosystem N accumulation, a feedback that in the long term could circumvent N limitation of the CO2 response while mitigating N pollution. We tested this prediction using a nine-year CO2 xN experiment in a tidal marsh. Although the effects of CO2 are similar between uplands and wetlands in many respects, this experiment offers a greater likelihood of detecting CO2 effects on N retention on a decadal timescale because tidal marshes have a relatively open N cycle and can accrue soil organic matter rapidly. To determine how elevated CO2 affects N dynamics, we assessed the three primary fates of N in a tidal marsh: (1) retention in plants and soil, (2) denitrification to the atmosphere, and (3) tidal export. We assessed changes in N pools and tracked the fate of a (15) N tracer added to each plot in 2006 to quantify the fraction of added N retained in vegetation and soil, and to estimate lateral N movement. Elevated CO2 alone did not increase plant N mass, soil N mass, or (15) N label retention. Unexpectedly, CO2 and N interacted such that the combined N+CO2 treatment increased ecosystem N accumulation despite the stimulation in N losses indicated by reduced (15) N label retention. These findings suggest that in N-limited ecosystems, elevated CO2 is unlikely to increase long-term N accumulation and circumvent progressive N limitation without additional N inputs, which may relieve plant-microbe competition and allow for increased plant N uptake.

Evolutionary history and novel biotic interactions determine plant responses to elevated CO2 and nitrogen fertilization.

A major frontier in global change research is predicting how multiple agents of global change will alter plant productivity, a critical component of the carbon cycle. Recent research has shown that plant responses to climate change are phylogenetically conserved such that species within some lineages are more productive than those within other lineages in changing environments. However, it remains unclear how phylogenetic patterns in plant responses to changing abiotic conditions may be altered by another agent of global change, the introduction of non-native species. Using a system of 28 native Tasmanian Eucalyptus species belonging to two subgenera, Symphyomyrtus and Eucalyptus, we hypothesized that productivity responses to abiotic agents of global change (elevated CO2 and increased soil N) are unique to lineages, but that novel interactions with a non-native species mediate these responses. We tested this hypothesis by examining productivity of 1) native species monocultures and 2) mixtures of native species with an introduced hardwood plantation species, Eucalyptus nitens, to experimentally manipulated soil N and atmospheric CO2. Consistent with past research, we found that N limits productivity overall, especially in elevated CO2 conditions. However, monocultures of species within the Symphyomyrtus subgenus showed the strongest response to N (gained 127% more total biomass) in elevated CO2 conditions, whereas those within the Eucalyptus subgenus did not respond to N. Root:shoot ratio (an indicator of resource use) was on average greater in species pairs containing Symphyomyrtus species, suggesting that functional traits important for resource uptake are phylogenetically conserved and explaining the phylogenetic pattern in plant response to changing environmental conditions. Yet, native species mixtures with E. nitens exhibited responses to CO2 and N that differed from those of monocultures, supporting our hypothesis and highlighting that both plant evolutionary history and introduced species will shape community productivity in a changing world.

Plant community feedbacks and long-term ecosystem responses to multi-factored global change.

While short-term plant responses to global change are driven by physiological mechanisms, which are represented relatively well by models, long-term ecosystem responses to global change may be determined by shifts in plant community structure resulting from other ecological phenomena such as interspecific interactions, which are represented poorly by models. In single-factor scenarios, plant communities often adjust to increase ecosystem response to that factor. For instance, some early global change experiments showed that elevated CO2 favours plants that respond strongly to elevated CO2, generally amplifying the response of ecosystem productivity to elevated CO2, a positive community feedback. However, most ecosystems are subject to multiple drivers of change, which can complicate the community feedback effect in ways that are more difficult to generalize. Recent studies have shown that (i) shifts in plant community structure cannot be reliably predicted from short-term plant physiological response to global change and (ii) that the ecosystem response to multi-factored change is commonly less than the sum of its parts. Here, we survey results from long-term field manipulations to examine the role community shifts may play in explaining these common findings. We use a simple model to examine the potential importance of community shifts in governing ecosystem response. Empirical evidence and the model demonstrate that with multi-factored change, the ecosystem response depends on community feedbacks, and that the magnitude of ecosystem response will depend on the relationship between plant response to one factor and plant response to another factor. Tradeoffs in the ability of plants to respond positively to, or to tolerate, different global change drivers may underlie generalizable patterns of covariance in responses to different drivers of change across plant taxa. Mechanistic understanding of these patterns will help predict the community feedbacks that determine long-term ecosystem responses.

Fire, hurricane and carbon dioxide: effects on net primary production of a subtropical woodland.

Disturbance affects most terrestrial ecosystems and has the potential to shape their responses to chronic environmental change. Scrub-oak vegetation regenerating from fire disturbance in subtropical Florida was exposed to experimentally elevated carbon dioxide (CO2) concentration (+350 µl l(-1)) using open-top chambers for 11 yr, punctuated by hurricane disturbance in year 8. Here, we report the effects of elevated CO2 on aboveground and belowground net primary productivity (NPP) and nitrogen (N) cycling during this experiment. The stimulation of NPP and N uptake by elevated CO2 peaked within 2 yr after disturbance by fire and hurricane, when soil nutrient availability was high. The stimulation subsequently declined and disappeared, coincident with low soil nutrient availability and with a CO2 -induced reduction in the N concentration of oak stems. These findings show that strong growth responses to elevated CO2 can be transient, are consistent with a progressively limited response to elevated CO2 interrupted by disturbance, and illustrate the importance of biogeochemical responses to extreme events in modulating ecosystem responses to global environmental change.

Ecosystem response to elevated CO(2) levels limited by nitrogen-induced plant species shift.

Terrestrial ecosystems gain carbon through photosynthesis and lose it mostly in the form of carbon dioxide (CO(2)). The extent to which the biosphere can act as a buffer against rising atmospheric CO(2) concentration in global climate change projections remains uncertain at the present stage. Biogeochemical theory predicts that soil nitrogen (N) scarcity may limit natural ecosystem response to elevated CO(2) concentration, diminishing the CO(2)-fertilization effect on terrestrial plant productivity in unmanaged ecosystems. Recent models have incorporated such carbon-nitrogen interactions and suggest that anthropogenic N sources could help sustain the future CO(2)-fertilization effect. However, conclusive demonstration that added N enhances plant productivity in response to CO(2)-fertilization in natural ecosystems remains elusive. Here we manipulated atmospheric CO(2) concentration and soil N availability in a herbaceous brackish wetland where plant community composition is dominated by a C(3) sedge and C(4) grasses, and is capable of responding rapidly to environmental change. We found that N addition enhanced the CO(2)-stimulation of plant productivity in the first year of a multi-year experiment, indicating N-limitation of the CO(2) response. But we also found that N addition strongly promotes the encroachment of C(4) plant species that respond less strongly to elevated CO(2) concentrations. Overall, we found that the observed shift in the plant community composition ultimately suppresses the CO(2)-stimulation of plant productivity by the third and fourth years. Although extensive research has shown that global change factors such as elevated CO(2) concentrations and N pollution affect plant species differently and that they may drive plant community changes, we demonstrate that plant community shifts can act as a feedback effect that alters the whole ecosystem response to elevated CO(2) concentrations. Moreover, we suggest that trade-offs between the abilities of plant taxa to respond positively to different perturbations may constrain natural ecosystem response to global change.

Elevated CO2 stimulates marsh elevation gain, counterbalancing sea-level rise.

Tidal wetlands experiencing increased rates of sea-level rise (SLR) must increase rates of soil elevation gain to avoid permanent conversion to open water. The maximal rate of SLR that these ecosystems can tolerate depends partly on mineral sediment deposition, but the accumulation of organic matter is equally important for many wetlands. Plant productivity drives organic matter dynamics and is sensitive to global change factors, such as rising atmospheric CO(2) concentration. It remains unknown how global change will influence organic mechanisms that determine future tidal wetland viability. Here, we present experimental evidence that plant response to elevated atmospheric [CO(2)] stimulates biogenic mechanisms of elevation gain in a brackish marsh. Elevated CO(2) (ambient + 340 ppm) accelerated soil elevation gain by 3.9 mm yr(-1) in this 2-year field study, an effect mediated by stimulation of below-ground plant productivity. Further, a companion greenhouse experiment revealed that the CO(2) effect was enhanced under salinity and flooding conditions likely to accompany future SLR. Our results indicate that by stimulating biogenic contributions to marsh elevation, increases in the greenhouse gas, CO(2), may paradoxically aid some coastal wetlands in counterbalancing rising seas.