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Microbial transformations of nitrogen (N) largely determine whether N is retained in ecosystems via net primary productivity or lost via gaseous emissions and leaching. The controls on soil N cycling are often studied at single locales, making it difficult to predict N cycling at regional to global scales. We hypothesized that contemporary soil properties exhibit consistent relationships with instantaneous gross N cycling rates across diverse biomes that create a continuum in these properties. We measured ex situ gross N cycling rates and soil properties at 33 study sites representing five biome classifications in California including deserts, grasslands, shrublands, forest, and wetlands. Desert soils had significantly lower total N, organic carbon (C), microbial biomass N, and soil moisture as well as higher pH than all other biomes, whereas forests and wetlands had significantly lower soil nitrate (NO3−) concentrations (P < 0.001 for all). Gross mineralization rates were best predicted by the combination of soil moisture and soil C:N ratios (R2 = 0.46), which exerted positive and negative controls, respectively. Grasslands exhibited marginally higher gross mineralization than all other biomes, whereas deserts had the lowest rates due to low soil moisture (P = 0.09). Gross nitrification rates were positively correlated to soil NO3− concentrations (R2 = 0.34) and negatively correlated to soil C:N ratios (R2 = 0.31). The negative relationship between gross nitrification and soil C:N ratios was driven by forest soils, which had significantly higher C:N ratios and lower gross nitrification than all other biomes (P < 0.05). Dissimilatory NO3− reduction to NH4+ (DNRA) occurred in soils from all biomes. The strong positive correlation between DNRA rates and soil NO3− (R2 = 0.41) suggests NO3− limitation of DNRA. Predictable patterns in gross N cycling across biomes in California suggest that contemporary soil properties are important drivers of instantaneous soil N cycling rates that integrate over differences in vegetation type, atmospheric N deposition rates, and local climate.

Anthropogenically derived nitrogen (N) has a central role in global environmental changes, including climate change, biodiversity loss, air pollution, greenhouse gas emission, water pollution, as well as food production and human health. Current understanding of the biogeochemical processes that govern the N cycle in coupled human–ecological systems around the globe is drawn largely from the long-term ecological monitoring and experimental studies. Here, we review spatial and temporal patterns and trends in reactive N emissions, and the interactions between N and other important elements that dictate their delivery from terrestrial to aquatic ecosystems, and the impacts of N on biodiversity and human society. Integrated international and long-term collaborative studies covering research gaps will reduce uncertainties and promote further understanding of the nitrogen cycle in various ecosystems.

Urban expansion is accelerating in the tropics, and may promote the spread of introduced plant species into urban-proximate forests. For example, soil disturbance can deplete the naturally high soil nitrogen pools in wet tropical soils, favoring introduced species with nitrogen-fixing capabilities. Also, forest fragmentation and canopy disturbance are likely to favor high-light species over shade-adapted rainforest species. We measured understory woody diversity, the abundance of introduced species, and soil nitrogen and carbon in urban, suburban, and rural secondary forests in Puerto Rico with canopies dominated by (1) native species, (2) introduced Fabaceae (potential nitrogen-fixers), and (3) introduced non-Fabaceae species. We hypothesized that forest stands with introduced Fabaceae in the canopy have higher soil nitrogen levels than stands with other introduced canopy species, and that this higher nitrogen is linked to increased native woody species diversity in the understory. We also predicted that more open canopies and smaller fragment sizes would be positively related with introduced species in the understory, and negatively related with total understory diversity. We found that stands with introduced Fabaceae in the canopy had significantly higher soil nitrogen levels than stands with other non-nitrogen fixing introduced species, and understory woody diversity in Fabaceae stands approached similar diversity levels as stands with native-dominated canopies. As predicted, aboveground stand structure and fragment size were also significantly associated with understory woody diversity across stands. These results suggest that introduced nitrogen-fixing trees may improve recruitment of native woody species in degraded tropical sites where native soil nitrogen is naturally high, particularly as Fabaceae stands mature and canopies close. © 2014 Elsevier B.V.

Mitigating further anthropogenic changes to the global climate will require reducing greenhouse-gas emissions ("abatement"), or else removing carbon dioxide from the atmosphere and/or diminishing solar input ("climate engineering"). Here, we develop and apply criteria to measure technical, economic, ecological, institutional, and ethical dimensions of, and public acceptance for, climate engineering strategies; provide a relative rating for each dimension; and offer a new interdisciplinary framework for comparing abatement and climate engineering options. While abatement remains the most desirable policy, certain climate engineering strategies, including forest and soil management for carbon sequestration, merit broad-scale application. Other proposed strategies, such as biochar production and geological carbon capture and storage, are rated somewhat lower, but deserve further research and development. Iron fertilization of the oceans and solar radiation management, although cost-effective, received the lowest ratings on most criteria. We conclude that although abatement should remain the central climate-change response, some low-risk, cost-effective climate engineering approaches should be applied as complements. The framework presented here aims to guide and prioritize further research and analysis, leading to improvements in climate engineering strategies. © The Ecological Society of America.

The increasing prevalence of drought events in grasslands and shrublands worldwide potentially has impacts on soil organic carbon (SOC). We leveraged the International Drought Experiment to study how SOC, including particulate organic carbon (POC) and mineral-associated organic carbon (MAOC) concentrations, responds to extreme drought treatments (1-in-100-year) for 1 to 5 years at 19 sites worldwide. In more mesic areas (aridity index > 0.65), SOC and POC concentrations decreased by 7.9% (±3.9) and 15.9% (±6.2) with drought, respectively, but there were no impacts on MAOC concentrations. However, drought had no impact on SOC, POC, or MAOC concentrations in drylands (aridity index < 0.65). The response of SOC to drought varied along an aridity gradient, concomitant with interannual precipitation variability and standing SOC concentration gradients. These findings highlight the differing response magnitudes of POC and MAOC concentrations to drought and the key regulating role of aridity.

Tropical forests account for over 50% of the global terrestrial carbon sink, but climate change threatens to alter the carbon balance of these ecosystems. We show that warming and drying of tropical forest soils may increase soil carbon vulnerability, by increasing degradation of older carbon. In situ whole-profile heating by 4 °C and 50% throughfall exclusion each increased the average radiocarbon age of soil CO 2 efflux by ~2-3 years, but the mechanisms underlying this shift differed. Warming accelerated decomposition of older carbon as increased CO 2 emissions depleted newer carbon. Drying suppressed decomposition of newer carbon inputs and decreased soil CO 2 emissions, thereby increasing contributions of older carbon to CO 2 efflux. These findings imply that both warming and drying, by accelerating the loss of older soil carbon or reducing the incorporation of fresh carbon inputs, will exacerbate soil carbon losses and negatively impact carbon storage in tropical forests under climate change., (© 2024. The Author(s).)

Tropical ecosystems face escalating global change. These shifts can disrupt tropical forests' carbon (C) balance and impact root dynamics. Since roots perform essential functions such as resource acquisition and tissue protection, root responses can inform about the strategies and vulnerabilities of ecosystems facing present and future global changes. However, root trait dynamics are poorly understood, especially in tropical ecosystems. We analyzed existing research on tropical root responses to key global change drivers: warming, drought, flooding, cyclones, nitrogen (N) deposition, elevated (e) CO 2 , and fires. Based on tree species- and community-level literature, we obtained 266 root trait observations from 93 studies across 24 tropical countries. We found differences in the proportion of root responsiveness to global change among different global change drivers but not among root categories. In particular, we observed that tropical root systems responded to warming and eCO 2 by increasing root biomass in species-scale studies. Drought increased the root: shoot ratio with no change in root biomass, indicating a decline in aboveground biomass. Despite N deposition being the most studied global change driver, it had some of the most variable effects on root characteristics, with few predictable responses. Episodic disturbances such as cyclones, fires, and flooding consistently resulted in a change in root trait expressions, with cyclones and fires increasing root production, potentially due to shifts in plant community and nutrient inputs, while flooding changed plant regulatory metabolisms due to low oxygen conditions. The data available to date clearly show that tropical forest root characteristics and dynamics are responding to global change, although in ways that are not always predictable. This synthesis indicates the need for replicated studies across root characteristics at species and community scales under different global change factors., (© 2024 Oak Ridge National Laboratory, managed by UT‐Battelle, LLC and The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

Tropical forest root characteristics and resource acquisition strategies are underrepresented in vegetation and global models, hampering the prediction of forest-climate feedbacks for these carbon-rich ecosystems. Lowland tropical forests often have globally unique combinations of high taxonomic and functional biodiversity, rainfall seasonality, and strongly weathered infertile soils, giving rise to distinct patterns in root traits and functions compared with higher latitude ecosystems. We provide a roadmap for integrating recent advances in our understanding of tropical forest belowground function into vegetation models, focusing on water and nutrient acquisition. We offer comparisons of recent advances in empirical and model understanding of root characteristics that represent important functional processes in tropical forests. We focus on: (1) fine-root strategies for soil resource exploration, (2) coupling and trade-offs in fine-root water vs nutrient acquisition, and (3) aboveground-belowground linkages in plant resource acquisition and use. We suggest avenues for representing these extremely diverse plant communities in computationally manageable and ecologically meaningful groups in models for linked aboveground-belowground hydro-nutrient functions. Tropical forests are undergoing warming, shifting rainfall regimes, and exacerbation of soil nutrient scarcity caused by elevated atmospheric CO 2 . The accurate model representation of tropical forest functions is crucial for understanding the interactions of this biome with the climate., (© 2024 The Authors New Phytologist © 2024 New Phytologist Foundation.)

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought., Competing Interests: Competing interests statement:The authors declare no competing interest.

The global demand for beef is rapidly increasing (FAO, 2019), raising concern about climate change impacts (Clark et al., 2020; Leip et al., 2015; Springmann et al., 2018). Beef and dairy contribute over 70% of livestock greenhouse gas emissions (GHG), which collectively contribute ~6.3 Gt CO 2 -eq/year (Gerber et al., 2013; Herrero et al., 2016) and account for 14%-18% of human GHG emissions (Friedlingstein et al., 2019; Gerber et al., 2013). The utility of beef GHG mitigation strategies, such as land-based carbon (C) sequestration and increased production efficiency, are actively debated (Garnett et al., 2017). We compiled 292 local comparisons of "improved" versus "conventional" beef production systems across global regions, assessing net GHG emission data from Life Cycle Assessment (LCA) studies. Our results indicate that net beef GHG emissions could be reduced substantially via changes in management. Overall, a 46 % reduction in net GHG emissions per unit of beef was achieved at sites using carbon (C) sequestration management strategies on grazed lands, and an 8% reduction in net GHGs was achieved at sites using growth efficiency strategies. However, net-zero emissions were only achieved in 2% of studies. Among regions, studies from Brazil had the greatest improvement, with management strategies for C sequestration and efficiency reducing beef GHG emissions by 57%. In the United States, C sequestration strategies reduced beef GHG emissions by over 100% (net-zero emissions) in a few grazing systems, whereas efficiency strategies were not successful at reducing GHGs, possibly because of high baseline efficiency in the region. This meta-analysis offers insight into pathways to substantially reduce beef production's global GHG emissions. Nonetheless, even if these improved land-based and efficiency management strategies could be fully applied globally, the trajectory of growth in beef demand will likely more than offset GHG emissions reductions and lead to further warming unless there is also reduced beef consumption., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Soils represent the largest terrestrial reservoir of organic carbon, and the balance between soil organic carbon (SOC) formation and loss will drive powerful carbon-climate feedbacks over the coming century. To date, efforts to predict SOC dynamics have rested on pool-based models, which assume classes of SOC with internally homogenous physicochemical properties. However, emerging evidence suggests that soil carbon turnover is not dominantly controlled by the chemistry of carbon inputs, but rather by restrictions on microbial access to organic matter in the spatially heterogeneous soil environment. The dynamic processes that control the physicochemical protection of carbon translate poorly to pool-based SOC models; as a result, we are challenged to mechanistically predict how environmental change will impact movement of carbon between soils and the atmosphere. Here, we propose a novel conceptual framework to explore controls on belowground carbon cycling: Probabilistic Representation of Organic Matter Interactions within the Soil Environment (PROMISE). In contrast to traditional model frameworks, PROMISE does not attempt to define carbon pools united by common thermodynamic or functional attributes. Rather, the PROMISE concept considers how SOC cycling rates are governed by the stochastic processes that influence the proximity between microbial decomposers and organic matter, with emphasis on their physical location in the soil matrix. We illustrate the applications of this framework with a new biogeochemical simulation model that traces the fate of individual carbon atoms as they interact with their environment, undergoing biochemical transformations and moving through the soil pore space. We also discuss how the PROMISE framework reshapes dialogue around issues related to SOC management in a changing world. We intend the PROMISE framework to spur the development of new hypotheses, analytical tools, and model structures across disciplines that will illuminate mechanistic controls on the flow of carbon between plant, soil, and atmospheric pools., (© 2020 John Wiley & Sons Ltd.)

Tropical forests are expected to green up with increasing atmospheric CO 2 concentrations, but primary productivity may be limited by soil nutrient availability. However, rarely have canopy-scale measurements been assessed against soil measurements in the tropics. Here, we sought to assess remotely sensed canopy greenness against steep soil nutrient gradients across 50 1-ha mature forest plots in Panama. Contrary to expectations, increases in in situ extractable soil phosphorus (P) and base cations (K, Mg) corresponded to declines in remotely sensed mean annual canopy greenness (r 2  = 0.77-0.85; p < 0.1), controlling for precipitation. The reason for this inverse relationship appears to be that litterfall also increased with increasing soil P and cation availability (r 2  = 0.88-0.98; p < 0.1), resulting in a decline in greenness with increasing annual litterfall (r 2  = 0.94; p < 0.1). As such, greater soil nutrient availability corresponded to greater leaf turnover, resulting in decreased greenness. However, these decreases in greenness with increasing soil P and cations were countered by increases in greenness with increasing soil nitrogen (N) (r 2  = 0.14; p < 0.1), which had no significant relationship with litterfall, likely reflecting a direct effect of soil N on leaf chlorophyll content, but not on litterfall rates. In addition, greenness increased with extractable soil aluminum (Al) (r 2  = 0.97; p < 0.1), but Al had no significant relationship with litterfall, suggesting a physiological adaptation of plants to high levels of toxic metals. Thus, spatial gradients in canopy greenness are not necessarily positive indicators of soil nutrient scarcity. Using a novel remote sensing index of canopy greenness limitation, we assessed how observed greenness compares with potential greenness. We found a strong relationship with soil N only (r 2  = 0.65; p < 0.1), suggesting that tropical canopy greenness in Panama is predominantly limited by soil N, even if plant productivity (e.g., litterfall) responds to rock-derived nutrients. Moreover, greenness limitation was also significantly correlated with fine root biomass and soil carbon stocks (r 2  = 0.62-0.71; p < 0.1), suggesting a feedback from soil N to canopy greenness to soil carbon storage. Overall, these data point to the potential utility of a remote sensing product for assessing belowground properties in tropical ecosystems.

Urban areas are expanding rapidly in tropical regions, with potential to alter ecosystem dynamics. In particular, exotic grasses and atmospheric nitrogen (N) deposition simultaneously affect tropical urbanized landscapes, with unknown effects on properties like soil carbon (C) storage. We hypothesized that (H1) soil nitrate (NO3 (-) ) is elevated nearer to the urban core, reflecting N deposition gradients. (H2) Exotic grasslands have elevated soil NO3 (-) and decreased soil C relative to secondary forests, with higher N promoting decomposer activity. (H3) Exotic grasslands have greater seasonality in soil NO3 (-) vs. secondary forests, due to higher sensitivity of grassland soil moisture to rainfall. We predicted that NO3 (-) would be positively related to dissolved organic C (DOC) production via changes in decomposer activity. We measured six paired grassland/secondary forest sites along a tropical urban-to-rural gradient during the three dominant seasons (hurricane, dry, and early wet). We found that (1) soil NO3 (-) was generally elevated nearer to the urban core, with particularly clear spatial trends for grasslands. (2) Exotic grasslands had lower soil C than secondary forests, which was related to elevated decomposer enzyme activities and soil respiration. Unexpectedly, soil NO3 (-) was negatively related to enzyme activities, and was lower in grasslands than forests. (3) Grasslands had greater soil NO3 (-) seasonality vs. forests, but this was not strongly linked to shifts in soil moisture or DOC. Our results suggest that exotic grasses in tropical regions are likely to drastically reduce soil C storage, but that N deposition may have an opposite effect via suppression of enzyme activities. However, soil NO3 (-) accumulation here was higher in urban forests than grasslands, potentially related to of aboveground N interception. Net urban effects on C storage across tropical landscapes will likely vary depending on the mosaic of grass cover, rates of N deposition, and responses by local decomposer communities., (© 2015 John Wiley & Sons Ltd.)

Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.