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Steven S. Perakis

Publications and source records attributed to Steven S. Perakis.

At least 19 recordsLinked to original sources

Blind spots in our understanding of the tropical nitrogen cycle

Understanding environmental controls on tropical nitrogen (N) cycling is critical for projecting flux rates and modeling ecosystem responses to rising CO 2 . Here, we ask whether geographic bias in field sampling may shape our understanding of N cycling by failing to capture the breadth of environmental heterogeneity within the tropics. We mapped empirical sampling locations of two major ecosystem N inputs and outputs—biological nitrogen fixation and denitrification—paired with a wide range of variables thought to influence N cycling. Both single and multi-variate analyses revealed that sampling density is greater in productive, wet, fertile, warmer tropical ecosystems with low seasonality—areas expected to have higher process rates—but biased away from spatially extensive biomes which may have high process rate variability and thus higher uncertainty. Even within tropical broadleaf forests, sampling is biased toward sites with unusually high soil fertility (N and phosphorus), potentially skewing biome-wide averages. Although some geographically undersampled regions share environmental characteristics with well-studied sites, others are unique, compromising inference from existing data. Finally, we find that while sampled field locations capture the range of N cycle process means/variances predicted by an ensemble of terrestrial biosphere models, this may be the result of circularity in model parameters and driving data rather than true coverage, including overrepresentation of forest plant functional types that may be parameterized from a small subset of sites. Data coverage expansion into areas with novel environmental conditions and higher uncertainty could be used to assess and improve model parameterization, as well as test general hypotheses.

Global Biogeochemical Cycles↗

Global carbon investment in terrestrial biological nitrogen fixation

Biological nitrogen (N) fixation (BNF) provides the N needed to produce proteins and other biological building blocks, helping feed humanity and mitigate climate change. Due to its high energetic cost compared to other forms of N acquisition, biotic investment in BNF indicates N limitation. Globally gridded BNF flux data provide an opportunity to determine the energetic investment in BNF across ecosystems, which would help reconcile conflicting indicators of N limitation. Here, we use a new BNF synthesis to quantify the relative importance of BNF in different N-fixing niches, in different biomes, and across the globe by calculating the fraction of net primary productivity (NPP) invested in BNF and the fraction of plant N acquisition provided by BNF. Larger fractions of non-agricultural NPP were invested in BNF in less-productive, higher-latitude biomes. This pattern was driven by biocrusts and mosses. Similarly, non-agricultural symbiotic N-fixing plants invested relatively more of their own NPP in BNF in less-productive, higher-latitude biomes. This symbiotic plant pattern was driven by shrubs and herbs, overriding the opposite pattern in trees. Symbiotic plants also acquired a higher fraction of their N from BNF at higher latitudes and in less productive biomes. Investments in symbiotic BNF were 10× higher in agricultural (2.9% of NPP) versus natural (0.29%) biomes, providing 26% versus 3.1% of ecosystem-scale plant N acquisition. These results support the paradigm of strong N limitation at higher latitudes, help understand the rarity of N-fixing trees at higher latitudes, underscore the dominance of human activity, and inform terrestrial biosphere models.

Global Biogeochemical Cycles↗

Rising atmospheric CO2 reduces nitrogen availability in boreal forests

Anthropogenic nitrogen (N) pollution has been emphasized as a cause of eutrophication globally. However, several recent datasets have suggested widespread oligotrophication may be occurring in some ecosystems, which is suggested to be a response to rising atmospheric carbon dioxide (eCO 2 ). Plant δ 15 N chronologies have served as primary evidence for oligotrophication, however, there has been wide disagreement whether eCO 2 or temporal changes in N deposition explain these patterns. We constructed δ 15 N tree ring chronologies across Sweden’s 23.5 million hectare productive forest area from the 1950s to 2010s. The study area spans a 1500 km latitudinal distance where N deposition varies four-fold, but where eCO 2 is spatially uniform. Our data revealed negative δ 15 N chronologies throughout Sweden, including forests in the far north where atmospheric N deposition rates are very low. Linear mixed effects models showed that eCO 2 was by far the strongest predictor of δ 15 N values, whereas N deposition variables, temperature, and forest basal area had much lower explanatory power. Our results clarify debates on the interpretation of previous δ 15 N chronologies, and provide clear evidence that eCO 2 is causing oligotrophication in boreal forests, which has implications for predicting their future role as sinks in the global carbon cycle.

Nature↗

Overestimated natural biological nitrogen fixation translates to an exaggerated CO2 fertilization effect in Earth System Models

CO 2 fertilization of the terrestrial biosphere is limited by nitrogen. Biological nitrogen fixation (BNF) is the dominant natural nitrogen source to the terrestrial biosphere and can alleviate nitrogen limitation but is poorly constrained in Earth system models (ESMs). Here, we compare terrestrial BNF from an ensemble of ESMs of the 6th Coupled Model Intercomparison Project to a new global synthesis of observations across natural and agricultural biomes. We find that compared to observations, ESMs underestimate agricultural BNF but overestimate natural BNF in the present day by over 50%. Natural BNF is overestimated in the most productive ecosystems that contribute most to the terrestrial carbon sink (forests and grasslands). ESMs with different BNF representations yield a range of BNF responses to CO 2 enrichment. Some ESMs with phenomenological representations of BNF predict a natural BNF increase in response to a doubling of CO 2 that aligns with a meta-analysis of CO 2 enrichment experiments (31% increase) but fail to account for the substantial carbon cost of BNF. In contrast, ESMs with mechanistic representations of BNF account for its carbon cost as well as its regulation by nitrogen limitation but overestimate the BNF response to a doubling of CO 2 (135% increase). Overall, all current BNF representations in ESMs fall short of fully capturing its response to rising atmospheric CO 2 . Finally, we find a positive correlation between modeled present-day natural BNF and the CO 2 fertilization effect across ESMs, suggesting that overestimated natural BNF translates to an exaggerated CO 2 fertilization effect of approximately 11% in ESMs.

PNAS↗

A global dataset of terrestrial biological nitrogen fixation

Biological nitrogen fixation (BNF) is the main natural source of new nitrogen inputs in terrestrial ecosystems, supporting terrestrial productivity, carbon uptake, and other Earth system processes. We assembled a comprehensive global dataset of field measurements of BNF in all major N-fixing niches across natural terrestrial biomes derived from the analysis of 376 BNF studies. The dataset comprises 32 variables, including site location, biome type, N-fixing niche, sampling year, quantification method, BNF rate (kg N ha −1 y −1 ), the percentage of nitrogen derived from the atmosphere (%N dfa ), N fixer or N-fixing substrate abundance, BNF rate per unit of N fixer abundance, and species identity. Overall, the dataset combines 1,207 BNF rates for trees, shrubs, herbs, soil, leaf litter, woody litter, dead wood, mosses, lichens, and biocrusts, 152 herb %N dfa values, 1,005 measurements of N fixer or N-fixing substrate abundance, and 762 BNF rates per unit of N fixer abundance for a total of 424 species across 66 countries. This dataset facilitates synthesis, meta-analysis, upscaling, and model benchmarking of BNF fluxes at multiple spatial scales.

Scientific Data↗

Global terrestrial nitrogen fixation and its modification by agriculture

Biological nitrogen fixation (BNF) is the largest natural source of new nitrogen (N) that supports terrestrial productivity 1,2 , yet estimates of global terrestrial BNF remain highly uncertain 3,4 . Here we show that this uncertainty is partly because of sampling bias, as field BNF measurements in natural terrestrial ecosystems occur where N fixers are 17 times more prevalent than their mean abundances worldwide. To correct this bias, we develop new estimates of global terrestrial BNF by upscaling field BNF measurements using spatially explicit abundances of all major biogeochemical N-fixing niches. We find that natural biomes sustain lower BNF, 65 (52–77) Tg N yr −1 , than previous empirical bottom-up estimates 3,4 , with most BNF occurring in tropical forests and drylands. We also find high agricultural BNF in croplands and cultivated pastures, 56 (54–58) Tg N yr −1 . Agricultural BNF has increased terrestrial BNF by 64% and total terrestrial N inputs from all sources by 60% over pre-industrial levels. Our results indicate that BNF may impose stronger constraints on the carbon sink in natural terrestrial biomes and represent a larger source of agricultural N than is generally considered in analyses of the global N cycle 5,6 , with implications for proposed safe operating limits for N use 7,8 .

Nature↗

Nitrogen deposition weakens soil carbon control of nitrogen dynamics across the contiguous United States

Anthropogenic nitrogen (N) deposition is unequally distributed across space and time, with inputs to terrestrial ecosystems impacted by industry regulations and variations in human activity. Soil carbon (C) content normally controls the fraction of mineralized N that is nitrified ( ƒ nitrified ), affecting N bioavailability for plants and microbes. However, it is unknown whether N deposition has modified the relationships among soil C, net N mineralization, and net nitrification. To test whether N deposition alters the relationship between soil C and net N transformations, we collected soils from coniferous and deciduous forests, grasslands, and residential yards in 14 regions across the contiguous United States that vary in N deposition rates. We quantified rates of net nitrification and N mineralization, soil chemistry (soil C, N, and pH), and microbial biomass and function (as beta-glucosidase (BG) and N -acetylglucosaminidase (NAG) activity) across these regions. Following expectations, soil C was a driver of ƒ nitrified across regions, whereby increasing soil C resulted in a decline in net nitrification and ƒ nitrified . The ƒ nitrified value increased with lower microbial enzymatic investment in N acquisition (increasing BG:NAG ratio) and lower active microbial biomass, providing some evidence that heterotrophic microbial N demand controls the ammonium pool for nitrifiers. However, higher total N deposition increased ƒ nitrified , including for high soil C sites predicted to have low ƒ nitrified , which decreased the role of soil C as a predictor of ƒ nitrified . Notably, the drop in contemporary atmospheric N deposition rates during the 2020 COVID-19 pandemic did not weaken the effect of N deposition on relationships between soil C and ƒ nitrified . Our results suggest that N deposition can disrupt the relationship between soil C and net N transformations, with this change potentially explained by weaker microbial competition for N. Therefore, past N inputs and soil C should be used together to predict N dynamics across terrestrial ecosystems.

contiguous United States↗

Nitrogen fixation and fertilization have similar effects on biomass allocation in nitrogen-fixing plants

Plants adjust their allocation to different organs based on nutrient supply. In some plant species, symbioses with nitrogen-fixing bacteria that live in root nodules provide an alternate pathway for nitrogen acquisition. Does access to nitrogen-fixing bacteria modify plants' biomass allocation? We hypothesized that access to nitrogen-fixing bacteria would have the same effect on allocation to aboveground versus belowground tissues as access to plentiful soil nitrogen. To test this hypothesis and related hypotheses about allocation to stems versus leaves and roots versus nodules, we conducted experiments with 15 species of nitrogen-fixing plants in two separate greenhouses. In each, we grew seedlings with and without access to symbiotic bacteria across a wide gradient of soil nitrogen supply. As is common, uninoculated plants allocated relatively less biomass belowground when they had more soil nitrogen. As we hypothesized, nitrogen fixation had a similar effect as the highest level of fertilization on allocation aboveground versus belowground. Both nitrogen fixation and high fertilization led to ~10% less biomass allocated belowground (~10% more aboveground) than the uninoculated, lowest fertilization treatment. Fertilization reduced allocation to nodules relative to roots. The responses for allocation of aboveground tissues to leaves versus stems were not as consistent across greenhouses or species as the other allocation trends, though more nitrogen fixation consistently led to relatively more allocation to leaves when soil nitrogen supply was low. Synthesis: Our results suggest that symbiotic nitrogen fixation causes seedlings to allocate relatively less biomass belowground, with potential implications for competition and carbon storage in early forest development.

Ecology and Evolution↗

Leaf nitrogen affects photosynthesis and water use efficiency similarly in nitrogen-fixing and non-fixing trees

Nitrogen (N)-fixing trees are thought to break a basic rule of leaf economics: higher leaf N concentrations do not translate into higher rates of carbon assimilation. Understanding how leaf N affects photosynthesis and water use efficiency (WUE) in this ecologically important group is critical. We grew six N-fixing and four non-fixing tree species for 4–5 years at four fertilization treatments in field experiments in temperate and tropical regions to assess how functional type (N fixer vs. non-fixer) and N limitation affected leaf N and how leaf N affected light-saturated photosynthesis ( A sat ), stomatal conductance ( g sw ) and WUE (WUE i and δ 13 C). A sat , WUE i and δ 13 C, but not g sw , increased with higher leaf N. Surprisingly, N-fixing and non-fixing trees displayed similar scaling between leaf N and these physiological variables, and this finding was supported by reanalysis of a global dataset. N fixers generally had higher leaf N than non-fixers, even when non-fixers were not N-limited at the leaf level. Leaf-level N limitation did not alter the relationship of A sat , g sw , WUE i and δ 13 C with leaf N, although it did affect the photosynthetic N use efficiency. Higher WUE was associated with higher productivity, whereas higher A sat was not. Synthesis : The ecological success of N-fixing trees depends on the effect of leaf N on carbon gain and water loss. Using a field fertilization experiment and reanalysis of a global dataset, we show that high leaf-level photosynthesis and WUE in N fixers stems from their higher average leaf N, rather than a difference between N fixers and non-fixers in the scaling of photosynthesis and WUE with leaf N. By clarifying the mechanism by which N fixers achieve and benefit from high WUE, our results further the understanding of global N fixer distributions.

Journal of Ecology↗

Nitrogen-bedrock interactions regulate multi-element nutrient limitation and sustainability in forests

Nutrient limitation of tree growth can intensify when nutrients are lost to forest harvest, creating challenges for forest growth and sustainability. Forest harvest accelerates nutrient loss by removing nutrient-containing biomass and by increasing nutrient leaching, shaping patterns of nutrient depletion that cause long-term shifts in nutrient limitation. Nitrogen most frequently limits tree growth, but where nitrogen is abundant, nutrient limitation often shifts to phosphorus and base cations, depending on soil mineralogy. We used the process-based biogeochemical model NutsFor to evaluate how multiple elements can limit long-term forest growth via interactions between soil nitrogen (low vs. high nitrogen) and soil mineralogy (sedimentary vs. basaltic bedrock). Simulations were run for 525 years with 40-year harvest intervals for managed Douglas-fir forests of the Oregon Coast Range. In low nitrogen sites, nutrient limitation switched after several centuries from nitrogen to phosphorus, as cycles of forest growth and harvest depleted soil organic phosphorus pools. In contrast, high nitrogen sites displayed severe base cation depletion and reduced tree growth within only one to two rotations, with sedimentary bedrock sites limited by calcium and basaltic sites by both calcium and potassium. Harvesting stimulated the largest fractional losses of nitrogen and potassium across all simulations, and additionally of calcium in high nitrogen sites. These multi-element simulations of interactions among harvesting, soil nitrogen, and bedrock type provide a set of testable predictions to guide monitoring and changes in management aimed at sustaining long-term forest productivity across a wide range of soil biogeochemical conditions.

Biogeochemistry↗

Symbiotic nitrogen fixation does not stimulate soil phosphatase activity under temperate and tropical trees

Symbiotic nitrogen (N)-fixing plants can enrich ecosystems with N, which can alter the cycling and demand for other nutrients. Researchers have hypothesized that fixed N could be used by plants and soil microbes to produce extracellular phosphatase enzymes, which release P from organic matter. Consistent with this speculation, the presence of N-fixing plants is often associated with high phosphatase activity, either in the soil or on root surfaces, although other studies have not found this association, and the connection between phosphatase and rates of N fixation—the mechanistic part of the argument—is tenuous. Here, we measured soil phosphatase activity under N-fixing trees and non-fixing trees transplanted and grown in tropical and temperate sites in the USA: two sites in Hawaii, and one each in New York and Oregon. This provides a rare example of phosphatase activity measured in a multi-site field experiment with rigorously quantified rates of N fixation. We found no difference in soil phosphatase activity under N-fixing vs. non-fixing trees nor across rates of N fixation, though we note that no sites were P limited and only one was N limited. Our results add to the literature showing no connection between N fixation rates and phosphatase activity.

Oecologia↗

Tree symbioses sustain nitrogen fixation despite excess nitrogen supply

Symbiotic nitrogen fixation (SNF) is a key ecological process whose impact depends on the strategy of SNF regulation—the degree to which rates of SNF change in response to limitation by N versus other resources. SNF that is obligate or exhibits incomplete downregulation can result in excess N fixation, whereas a facultative SNF strategy does not. We hypothesized that tree-based SNF strategies differed by latitude (tropical vs. temperate) and symbiotic type (actinorhizal vs. rhizobial). Specifically, we expected tropical rhizobial symbioses to display strongly facultative SNF as an explanation of their success in low-latitude forests. In this study we used 15 N isotope dilution field experiments in New York, Oregon, and Hawaii to determine SNF strategies in six N-fixing tree symbioses. Nitrogen fertilization with +10 and +15 g N m −2 year −1 for 4–5 years alleviated N limitation in all taxa, paving the way to determine SNF strategies. Contrary to our hypothesis, all six of the symbioses we studied sustained SNF even at high N. Robinia pseudoacacia (temperate rhizobial) fixed 91% of its N (%N dfa ) in controls, compared to 64% and 59% in the +10 and +15 g N m −2 year −1 treatments. For Alnus rubra (temperate actinorhizal), %N dfa was 95%, 70%, and 60%. For the tropical species, %N dfa was 86%, 80%, and 82% for Gliricidia sepium (rhizobial); 79%, 69%, and 67% for Casuarina equisetifolia (actinorhizal); 91%, 42%, and 67% for Acacia koa (rhizobial); and 60%, 51%, and 19% for Morella faya (actinorhizal). Fertilization with phosphorus did not stimulate tree growth or SNF. These results suggest that the latitudinal abundance distribution of N-fixing trees is not caused by a shift in SNF strategy. They also help explain the excess N in many forests where N fixers are common.

Hawai'i, New York, Oregon↗

Exploring the role of cryptic nitrogen fixers in terrestrial ecosystems: A frontier in nitrogen cycling research

Biological nitrogen fixation represents the largest natural flux of new nitrogen (N) into terrestrial ecosystems, providing a critical N source to support net primary productivity of both natural and agricultural systems. When they are common, symbiotic associations between plants and bacteria can add more than 100 kg N ha −1 y −1 to ecosystems. Yet, these associations are uncommon in many terrestrial ecosystems. In most cases, N inputs derive from more cryptic sources, including mutualistic and/or free-living microorganisms in soil, plant litter, decomposing roots and wood, lichens, insects, and mosses, among others. Unfortunately, large gaps remain in the understanding of cryptic N fixation. We conducted a literature review to explore rates, patterns, and controls of cryptic N fixation in both unmanaged and agricultural ecosystems. Our analysis indicates that, as is common with N fixation, rates are highly variable across most cryptic niches, with N inputs in any particular cryptic niche ranging from near zero to more than 20 kg ha −1 y −1 . Such large variation underscores the need for more comprehensive measurements of N fixation by organisms not in symbiotic relationships with vascular plants in terrestrial ecosystems, as well as identifying the factors that govern cryptic N fixation rates. We highlight several challenges, opportunities, and priorities in this important research area, and we propose a conceptual model that posits an interacting hierarchy of biophysical and biogeochemical controls over N fixation that should generate valuable new hypotheses and research.

Ecosystems↗

N and P constrain C in ecosystems under climate change: Role of nutrient redistribution, accumulation, and stoichiometry

We use the Multiple Element Limitation (MEL) model to examine responses of twelve ecosystems to elevated carbon dioxide (CO 2 ), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO 2 , warming, and decreased precipitation combined because higher water-use efficiency with elevated CO 2 and higher fertility with warming compensate for responses to drought. Response to elevated CO 2 , warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO 2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C-nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO 2 and climate change.

Ecological Applications↗

Hypotheses and lessons from a native moth outbreak in a low-diversity, tropical rainforest

Outbreaks of defoliating insects in low-diversity tropical forests occur infrequently but provide valuable insights about outbreak ecology in temperate environments and in general. We investigated an extensive outbreak of the endemic koa moth ( Scotorythra paludicola ), which defoliated endemic koa trees ( Acacia koa ) over a third of their range on Hawai‘i Island during 2013 and 2014. At Hakalau Forest National Wildlife Refuge, we observed the dynamics of the outbreak and its effects on host trees, nutrient cycling, and insectivorous consumers in reforestation stands of densely planted koa and in natural forest stands of mixed koa and ‘ōhi‘a ( Metrosideros polymorpha ). Contrary to predictions of the resource concentration hypothesis, caterpillar biomass and defoliation severity were greater in the natural forest sites, where koa density was relatively low. Caterpillars preferentially consumed the most palatable koa foliage type (phyllodes), and koa initially refoliated with the least palatable foliage type (true leaves). Lightly defoliated small trees refoliated more quickly than did heavily defoliated ones but the opposite was true for large trees, which also produced a greater proportion of phyllodes. Mortality was greatest for heavily defoliated small koa. Caterpillar frass caused larger increases in soil nitrogen (N) than phosphorus (P) availability, with the greatest N increases in fine-textured soils. Foliar N increased in alien grasses under koa canopies compared to grasses away from koa and to native woody understory species. Bird activity was influenced by ‘ōhi‘a flower abundance and the severity of koa defoliation; birds switched to outbreaking caterpillar prey, and they gained weight during the outbreak. Bat foraging times decreased during the outbreak, apparently because they became satiated quickly each night. Parasitoid wasps increased with caterpillar abundance but had little influence on outbreak dynamics. Reducing alien grass cover and increasing tree diversity would likely reduce the impacts of insect outbreaks and similar perturbations to native forests.

Hawai'i↗

Climatic aridity shapes post-fire interactions between Ceanothus spp. and Douglas-fir (Pseudotsuga menziesii) across the Klamath Mountains

Climate change is leading to increased drought intensity and fire frequency, creating early-successional landscapes with novel disturbance–recovery dynamics. In the Klamath Mountains of northwestern California and southwestern Oregon, early-successional interactions between nitrogen (N)-fixing shrubs ( Ceanothus spp.) and long-lived conifers (Douglas-fir) are especially important determinants of forest development. We sampled post-fire vegetation and soil biogeochemistry in 57 plots along gradients of time since fire (7–28 years) and climatic water deficit (aridity). We found that Ceanothus biomass increased, and Douglas-fir biomass decreased with increasing aridity. High aridity and Ceanothus biomass interacted with lower soil C:N more than either factor alone. Ceanothus biomass was initially high after fire and declined with time, suggesting a large initial pulse of N-fixation that could enhance N availability for establishing Douglas-fir. We conclude that future increases in aridity and wildfire frequency will likely limit post-fire Douglas-fir establishment, though Ceanothus may ameliorate some of these impacts through benefits to microclimate and soils. Results from this study contribute to our understanding of the effects of climate change and wildfires on interspecific interactions and forest dynamics. Management seeking to accelerate forest recovery after high-severity fire should emphasize early-successional conifer establishment while maintaining N-fixing shrubs to enhance soil fertility.

California, Oregon↗

Early growth and ecophysiological responses of Koa (Acacia koa A. Gray) seedlings to reduced water and phosphorus

Sites in need of restoration typically have one or more environmental factors that limit seedling establishment. Identifying ecophysiological responses to environmental stressors can provide important insights into mitigating measures that would allow seedlings to overcome such constraints to survival. Koa ( Acacia koa A. Gray) is a nitrogen-fixing tree species endemic to Hawaiʻi that is highly valued in restoring degraded forest ecosystems, which are often limited in available water and phosphorus. This study examined how koa seedlings respond to conditions of reduced water (65 W) and no phosphorus (0P). After 17 weeks, seedlings subjected to 65 W or 0P accumulated less biomass, smaller root-collar diameters, and lower nitrogen and phosphorus contents. Combined reductions in water and P resulted in seedlings with increased root to shoot dry biomass and shorter shoots. Seedlings subjected to 65 W also had lower instantaneous rates of CO 2 assimilation, but higher instantaneous water-use efficiencies following irrigation, suggesting that koa responds to water deficits by decreasing water loss via reduced stomatal conductance. Seedlings subjected to 0P had similar rates of CO 2 assimilation relative to those grown with adequate P, suggesting that koa is able to employ strategies to avoid physiological impairment from conditions of inadequate P. Future research should assess whether subjecting koa seedlings to reduced water before planting on water-limited sites cues increased drought resistance and whether uptake and storage of P by seedlings in the nursery better supports growth following outplanting, particularly on sites with anticipated low plant-available water.

Hawai'i↗

N supply mediates the radiative balance of N2O emissions and CO2 sequestration driven by N-fixing vs. non-fixing trees

Forests are a significant CO 2 sink. However, CO 2 sequestration in forests is radiatively offset by emissions of nitrous oxide (N 2 O), a potent greenhouse gas, from forest soils. Reforestation, an important strategy for mitigating climate change, has focused on maximizing CO 2 sequestration in plant biomass without integrating N 2 O emissions from soils. Although nitrogen (N)-fixing trees are often recommended for reforestation because of their rapid growth on N-poor soil, they can stimulate significant N 2 O emissions from soils. Here, we first used a field experiment to show that a N-fixing tree ( Robinia pseudoacacia ) initially mitigated climate change more than a non-fixing tree ( Betula nigra ). We then used our field data to parameterize a theoretical model to investigate these effects over time. Under lower N supply, N-fixers continued to mitigate climate change more than non-fixers by overcoming N limitation of plant growth. However, under higher N supply, N-fixers ultimately mitigated climate change less than non-fixers by enriching soil N and stimulating N 2 O emissions from soils. These results have implications for reforestation, suggesting that N-fixing trees are more effective at mitigating climate change at lower N supply, whereas non-fixing trees are more effective at mitigating climate change at higher N supply.

New York↗