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At least 73 records · Page 4

Cellular remains in a ~3.42-billion-year-old subseafloor hydrothermal environment

Subsurface habitats on Earth host an extensive extant biosphere and likely provided one of Earth’s earliest microbial habitats. Though the site of life’s emergence continues to be debated, evidence of early life provides insights into its early evolution1. Here we present our discovery of exceptionally well-preserved ~3.40-billion-years-old carbonaceous filamentous microfossils that inhabited a paleo-subseafloor hydrothermal vein system of the Barberton greenstone belt (BGB) in South Africa. The distribution of the microfossils in growth position on the walls of the initial conduits created by low- temperature hydrothermal fluid and their morphological and chemical characteristics as investigated over a range of scales, are consistent with an assemblage of methanogens that thrived in an ultramafic volcanic substrate.

Cavalazzi, Barbara↗

Complementary Metagenomic Approaches Improve Reconstruction of Microbial Diversity in a Forest Soil

ABSTRACT Soil ecosystems harbor diverse microorganisms and yet remain only partially characterized as neither single-cell sequencing nor whole-community sequencing offers a complete picture of these complex communities. Thus, the genetic and metabolic potential of this “uncultivated majority” remains underexplored. To address these challenges, we applied a pooled-cell-sorting-based mini-metagenomics approach and compared the results to bulk metagenomics. Informatic binning of these data produced 200 mini-metagenome assembled genomes (sorted-MAGs) and 29 bulk metagenome assembled genomes (MAGs). The sorted and bulk MAGs increased the known phylogenetic diversity of soil taxa by 7.2% with respect to the Joint Genome Institute IMG/M database and showed clade-specific sequence recruitment patterns across diverse terrestrial soil metagenomes. Additionally, sorted-MAGs expanded the rare biosphere not captured through MAGs from bulk sequences, exemplified through phylogenetic and functional analyses of members of the phylum Bacteroidetes . Analysis of 67 Bacteroidetes sorted-MAGs showed conserved patterns of carbon metabolism across four clades. These results indicate that mini-metagenomics enables genome-resolved investigation of predicted metabolism and demonstrates the utility of combining metagenomics methods to tap into the diversity of heterogeneous microbial assemblages. IMPORTANCE Microbial ecologists have historically used cultivation-based approaches as well as amplicon sequencing and shotgun metagenomics to characterize microbial diversity in soil. However, challenges persist in the study of microbial diversity, including the recalcitrance of the majority of microorganisms to laboratory cultivation and limited sequence assembly from highly complex samples. The uncultivated majority thus remains a reservoir of untapped genetic diversity. To address some of the challenges associated with bulk metagenomics as well as low throughput of single-cell genomics, we applied flow cytometry-enabled mini-metagenomics to capture expanded microbial diversity from forest soil and compare it to soil bulk metagenomics. Our resulting data from this pooled-cell sorting approach combined with bulk metagenomics revealed increased phylogenetic diversity through novel soil taxa and rare biosphere members. In-depth analysis of genomes within the highly represented Bacteroidetes phylum provided insights into conserved and clade-specific patterns of carbon metabolism.

59 BASIC BIOLOGICAL SCIENCES↗

Phototrophy and carbon fixation in Chlorobi postdate the rise of oxygen

While most productivity on the surface of the Earth today is fueled by oxygenic photosynthesis, for much of Earth history it is thought that anoxygenic photosynthesis—using compounds like ferrous iron or sulfide as electron donors—drove most global carbon fixation. Anoxygenic photosynthesis is still performed by diverse bacteria in niche environments today. Of these, the Chlorobi (formerly green sulfur bacteria) are often interpreted as being particularly ancient and are frequently proposed to have fueled the biosphere during late Archean and early Paleoproterozoic time before the rise of oxygenic photosynthesis. Here, we perform comparative genomic, phylogenetic, and molecular clock analyses to determine the antiquity of the Chlorobi and their characteristic phenotypes. We show that contrary to common assumptions, the Chlorobi clade is relatively young, with anoxygenic phototrophy, carbon fixation via the rTCA pathway, and iron oxidation all significantly postdating the rise of oxygen ~2.3 billion years ago. The Chlorobi therefore could not have fueled the Archean biosphere, but instead represent a relatively young radiation of organisms which likely acquired the capacity for anoxygenic photosynthesis and other traits via horizontal gene transfer sometime after the evolution of oxygenic Cyanobacteria.

59 BASIC BIOLOGICAL SCIENCES↗

Advances in 3D Geologic Modeling of Alluvial Basins with a Focus on Facies and Property Modeling

The unsaturated zone alluvium reference case is one of several geologic systems under consideration by the U.S. Department of Energy Office of Nuclear Energy for hosting repositories for spent nuclear fuel and associated waste (Sevougian et al., 2019). As noted by Mariner et al. (2018), the generic alluvial basin offers positive attributes that merit its consideration as a reference case by the Spent Fuel and Waste Science and Technology (SFWST) campaign. There are hundreds of alluvial basins and sub-basins scattered across the arid western United States (Figure 1-1). Precipitation and infiltration rates are relatively low with high evapotranspiration, resulting in vertical separation between repository and water table and thus longer transport paths to an aquifer. Accumulations of alluvial sediments within these basins are typically on the order of hundreds of meters, and locally may exceed 1,000 m in thickness, as is the case for the Deming sub-basin in southern New Mexico. A thick geologic host medium, which serves as the natural barrier system (NBS) in the conceptual model framework of a geologic disposal system, isolates the waste packages from receptors in the biosphere. Further, alluvial basin fill is typically comprised of stacked playa and lacustrine deposits along the basin axis (Perry et al., 2018). Characterized by low permeability, these layers protect the biosphere above the repository and the groundwater resources below the repository.

12 MANAGEMENT OF RADIOACTIVE AND NON-RADIOACTIVE W↗

Incorporating Heterogeneity into 3D Geologic Models for the Alluvial Basin Reference Case

The unsaturated zone alluvium reference case is one of several geologic systems under consideration by the U.S. Department of Energy Office of Nuclear Energy for hosting repositories for spent nuclear fuel and associated waste (Sevougian et al., 2019). As noted by Mariner et al. (2018), the generic alluvial basin offers positive attributes that merit its consideration as a reference case by the Spent Fuel and Waste Science and Technology (SFWST) campaign. Hundreds of alluvial basins and sub-basins are scattered across the arid western United States (Figure 1-1). Precipitation and infiltration rates are relatively low with high evapotranspiration, resulting in deep regional water tables. Deep regional water tables allow for significant vertical separation between proposed repository depths and groundwater and thus longer transport paths from a hypothetical repository to an aquifer. Accumulations of alluvial sediments within these basins are typically on the order of hundreds of meters, and locally may exceed 1,000 m in thickness, as is the case for the Deming sub-basin in southern New Mexico. In alluvial basins, alluvium creates a thick geologic host medium that serves as the natural barrier system (NBS) in the conceptual model framework of a geologic disposal system and isolates the waste packages from receptors in the biosphere. Further, alluvial basin fill is typically comprised of stacked playa and lacustrine deposits along the basin axis (Perry et al., 2018). Characterized by low permeability, these layers protect the biosphere above the repository and the groundwater resources below the repository by limiting flow and transport of radionuclides.

12 MANAGEMENT OF RADIOACTIVE AND NON-RADIOACTIVE W↗

Scaling carbon fluxes from eddy covariance sites to globe: synthesis and evaluation of the FLUXCOM approach

FLUXNET comprises globally distributed eddy-covariance-based estimates of carbon fluxes between the biosphere and the atmosphere. Since eddy covariance flux towers have a relatively small footprint and are distributed unevenly across the world, upscaling the observations is necessary to obtain global-scale estimates of biosphere–atmosphere exchange. Based on cross-consistency checks with atmospheric inversions, sun-induced fluorescence (SIF) and dynamic global vegetation models (DGVMs), here we provide a systematic assessment of the latest upscaling efforts for gross primary production (GPP) and net ecosystem exchange (NEE) of the FLUXCOM initiative, where different machine learning methods, forcing data sets and sets of predictor variables were employed. Spatial patterns of mean GPP are consistent across FLUXCOM and DGVM ensembles (R 2 >0.94 at 1° spatial resolution) while the majority of DGVMs show, for 70 % of the land surface, values outside the FLUXCOM range. Global mean GPP magnitudes for 2008–2010 from FLUXCOM members vary within 106 and 130 PgC yr -1 with the largest uncertainty in the tropics. Seasonal variations in independent SIF estimates agree better with FLUXCOM GPP (mean global pixel-wise R 2 ~0.75) than with GPP from DGVMs (mean global pixel-wise R 2 ~0.6). Seasonal variations in FLUXCOM NEE show good consistency with atmospheric inversion-based net land carbon fluxes, particularly for temperate and boreal regions (R 2 >0.92). Interannual variability of global NEE in FLUXCOM is underestimated compared to inversions and DGVMs. The FLUXCOM version which also uses meteorological inputs shows a strong co-variation in interannual patterns with inversions (R 2 =0.87 for 2001–2010). Mean regional NEE from FLUXCOM shows larger uptake than inversion and DGVM-based estimates, particularly in the tropics with discrepancies of up to several hundred grammes of carbon per square metre per year. These discrepancies can only partly be reconciled by carbon loss pathways that are implicit in inversions but not captured by the flux tower measurements such as carbon emissions from fires and water bodies. We hypothesize that a combination of systematic biases in the underlying eddy covariance data, in particular in tall tropical forests, and a lack of site history effects on NEE in FLUXCOM are likely responsible for the too strong tropical carbon sink estimated by FLUXCOM. Furthermore, as FLUXCOM does not account for CO 2 fertilization effects, carbon flux trends are not realistic. Overall, current FLUXCOM estimates of mean annual and seasonal cycles of GPP as well as seasonal NEE variations provide useful constraints of global carbon cycling, while interannual variability patterns from FLUXCOM are valuable but require cautious interpretation. Exploring the diversity of Earth observation data and of machine learning concepts along with improved quality and quantity of flux tower measurements will facilitate further improvements of the FLUXCOM approach overall.

54 ENVIRONMENTAL SCIENCES↗

Carbon cycle extremes accelerate weakening of the land carbon sink in the late 21st century

Increasing surface temperature could lead to enhanced evaporation, reduced soil moisture availability, and more frequent droughts and heat waves. The spatiotemporal co-occurrence of such effects further drives extreme anomalies in vegetation productivity and net land carbon storage. However, the impacts of climate change on extremes in net biospheric production (NBP) over longer time periods are unknown. Using the percentile threshold on the probability distribution curve of NBP anomalies, we computed negative and positive extremes in NBP. Here we show that due to climate warming, about 88% of global regions will experience a larger magnitude of negative NBP extremes than positive NBP extremes toward the end of 2100, which accelerate the weakening of the land carbon sink. Our analysis indicates the frequency of negative extremes associated with declines in biospheric productivity was larger than positive extremes, especially in the tropics. While the overall impact of warming at high latitudes is expected to increase plant productivity and carbon uptake, high-temperature anomalies increasingly induce negative NBP extremes toward the end of the 21st century. Using regression analysis, we found soil moisture anomalies to be the most dominant individual driver of NBP extremes. The compound effect of hotness, dryness, and fire caused extremes at more than 50% of the total grid cells. The larger proportion of negative NBP extremes raises a concern about whether the Earth is capable of increasing vegetation production with a growing human population and rising demand for plant material for food, fiber, fuel, and building materials. The increasing proportion of negative NBP extremes highlights the consequences not only of reduction in total carbon uptake capacity but also of conversion of land to a carbon source.

54 ENVIRONMENTAL SCIENCES↗

Global Carbon Budget 2021

Abstract. Accurate assessment of anthropogenic carbon dioxide (CO2) emissions and their redistribution among the atmosphere, ocean, and terrestrial biosphere in a changing climate is critical to better understand the global carbon cycle, support the development of climate policies, and project future climate change. Here we describe and synthesize datasets and methodology to quantify the five major components of the global carbon budget and their uncertainties. Fossil CO2 emissions (EFOS) are based on energy statistics and cement production data, while emissions from land-use change (ELUC), mainly deforestation, are based on land use and land-use change data and bookkeeping models. Atmospheric CO2 concentration is measured directly, and its growth rate (GATM) is computed from the annual changes in concentration. The ocean CO2 sink (SOCEAN) is estimated with global ocean biogeochemistry models and observation-based data products. The terrestrial CO2 sink (SLAND) is estimated with dynamic global vegetation models. The resulting carbon budget imbalance (BIM), the difference between the estimated total emissions and the estimated changes in the atmosphere, ocean, and terrestrial biosphere, is a measure of imperfect data and understanding of the contemporary carbon cycle. All uncertainties are reported as ±1σ. For the first time, an approach is shown to reconcile the difference in our ELUC estimate with the one from national greenhouse gas inventories, supporting the assessment of collective countries' climate progress. For the year 2020, EFOS declined by 5.4 % relative to 2019, with fossil emissions at 9.5 ± 0.5 GtC yr−1 (9.3 ± 0.5 GtC yr−1 when the cement carbonation sink is included), and ELUC was 0.9 ± 0.7 GtC yr−1, for a total anthropogenic CO2 emission of 10.2 ± 0.8 GtC yr−1 (37.4 ± 2.9 GtCO2). Also, for 2020, GATM was 5.0 ± 0.2 GtC yr−1 (2.4 ± 0.1 ppm yr−1), SOCEAN was 3.0 ± 0.4 GtC yr−1, and SLAND was 2.9 ± 1 GtC yr−1, with a BIM of −0.8 GtC yr−1. The global atmospheric CO2 concentration averaged over 2020 reached 412.45 ± 0.1 ppm. Preliminary data for 2021 suggest a rebound in EFOS relative to 2020 of +4.8 % (4.2 % to 5.4 %) globally. Overall, the mean and trend in the components of the global carbon budget are consistently estimated over the period 1959–2020, but discrepancies of up to 1 GtC yr−1 persist for the representation of annual to semi-decadal variability in CO2 fluxes. Comparison of estimates from multiple approaches and observations shows (1) a persistent large uncertainty in the estimate of land-use changes emissions, (2) a low agreement between the different methods on the magnitude of the land CO2 flux in the northern extra-tropics, and (3) a discrepancy between the different methods on the strength of the ocean sink over the last decade. This living data update documents changes in the methods and datasets used in this new global carbon budget and the progress in understanding of the global carbon cycle compared with previous publications of this dataset (Friedlingstein et al., 2020, 2019; Le Quéré et al., 2018b, a, 2016, 2015b, a, 2014, 2013). The data presented in this work are available at https://doi.org/10.18160/gcp-2021 (Friedlingstein et al., 2021).

Friedlingstein, Pierre (ORCID:0000000333094739)↗

Global Carbon Budget 2022

Abstract. Accurate assessment of anthropogenic carbon dioxide (CO2) emissions and their redistribution among the atmosphere, ocean, and terrestrial biosphere in a changing climate is critical to better understand the global carbon cycle, support the development of climate policies, and project future climate change. Here we describe and synthesize data sets and methodologies to quantify the five major components of the global carbon budget and their uncertainties. Fossil CO2 emissions (EFOS) are based on energy statistics and cement production data, while emissions from land-use change (ELUC), mainly deforestation, are based on land use and land-use change data and bookkeeping models. Atmospheric CO2 concentration is measured directly, and its growth rate (GATM) is computed from the annual changes in concentration. The ocean CO2 sink (SOCEAN) is estimated with global ocean biogeochemistry models and observation-based data products. The terrestrial CO2 sink (SLAND) is estimated with dynamic global vegetation models. The resulting carbon budget imbalance (BIM), the difference between the estimated total emissions and the estimated changes in the atmosphere, ocean, and terrestrial biosphere, is a measure of imperfect data and understanding of the contemporary carbon cycle. All uncertainties are reported as ±1σ. For the year 2021, EFOS increased by 5.1 % relative to 2020, with fossil emissions at 10.1 ± 0.5 GtC yr−1 (9.9 ± 0.5 GtC yr−1 when the cement carbonation sink is included), and ELUC was 1.1 ± 0.7 GtC yr−1, for a total anthropogenic CO2 emission (including the cement carbonation sink) of 10.9 ± 0.8 GtC yr−1 (40.0 ± 2.9 GtCO2). Also, for 2021, GATM was 5.2 ± 0.2 GtC yr−1 (2.5 ± 0.1 ppm yr−1), SOCEAN was 2.9 ± 0.4 GtC yr−1, and SLAND was 3.5 ± 0.9 GtC yr−1, with a BIM of −0.6 GtC yr−1 (i.e. the total estimated sources were too low or sinks were too high). The global atmospheric CO2 concentration averaged over 2021 reached 414.71 ± 0.1 ppm. Preliminary data for 2022 suggest an increase in EFOS relative to 2021 of +1.0 % (0.1 % to 1.9 %) globally and atmospheric CO2 concentration reaching 417.2 ppm, more than 50 % above pre-industrial levels (around 278 ppm). Overall, the mean and trend in the components of the global carbon budget are consistently estimated over the period 1959–2021, but discrepancies of up to 1 GtC yr−1 persist for the representation of annual to semi-decadal variability in CO2 fluxes. Comparison of estimates from multiple approaches and observations shows (1) a persistent large uncertainty in the estimate of land-use change emissions, (2) a low agreement between the different methods on the magnitude of the land CO2 flux in the northern extratropics, and (3) a discrepancy between the different methods on the strength of the ocean sink over the last decade. This living data update documents changes in the methods and data sets used in this new global carbon budget and the progress in understanding of the global carbon cycle compared with previous publications of this data set. The data presented in this work are available at https://doi.org/10.18160/GCP-2022 (Friedlingstein et al., 2022b).

54 ENVIRONMENTAL SCIENCES↗

Organic carbon burial by river meandering partially offsets bank erosion carbon fluxes in a discontinuous permafrost floodplain

Abstract. Arctic river systems erode permafrost in their banks and mobilize particulate organic carbon (OC). Meandering rivers can entrain particulate OC from permafrost many meters below the depth of annual thaw, potentially enabling the production of greenhouse gases. However, the amount and fate of permafrost OC that is mobilized by river erosion is uncertain. To constrain OC fluxes due to riverbank erosion and deposition, we collected riverbank and floodplain sediment samples along the Koyukuk River, which meanders through discontinuous permafrost in the Yukon River watershed, Alaska, USA, with an average migration rate of 0.52 m yr−1. We measured sediment total OC (TOC) content, radiocarbon activity, water content, bulk density, grain size, and floodplain stratigraphy. Radiocarbon activity and TOC content were higher in samples dominated by silt as compared to sand, which we used to map OC content onto floodplain stratigraphy and estimate carbon fluxes due to river meandering. Results showed that the Koyukuk River erodes and re-deposits a substantial flux of OC each year due to its depth and high migration rate, generating a combined OC flux of a similar magnitude to the floodplain net ecological productivity. However, sediment being eroded from cutbanks and deposited as point bars had similar OC stocks (mean ± 1 SD of 125.3±13.1 kg OC m−2 in cutbanks versus 114.0±15.7 kg OC m−2 in point bars) whether or not the banks contained permafrost. We also observed radiocarbon-depleted biospheric OC in both cutbanks and permafrost-free point bars. These results indicate that a substantial fraction of aged biospheric OC that is liberated from floodplains by bank erosion is subsequently re-deposited in point bars rather than being oxidized. The process of aging, erosion, and re-deposition of floodplain organic material may be intrinsic to river–floodplain dynamics, regardless of permafrost content.

54 ENVIRONMENTAL SCIENCES↗

Assessing dynamic vegetation model parameter uncertainty across Alaskan arctic tundra plant communities

Abstract As the Arctic region moves into uncharted territory under a warming climate, it is important to refine the terrestrial biosphere models (TBMs) that help us understand and predict change. One fundamental uncertainty in TBMs relates to model parameters, configuration variables internal to the model whose value can be estimated from data. We incorporate a version of the Terrestrial Ecosystem Model (TEM) developed for arctic ecosystems into the Predictive Ecosystem Analyzer (PEcAn) framework. PEcAn treats model parameters as probability distributions, estimates parameters based on a synthesis of available field data, and then quantifies both model sensitivity and uncertainty to a given parameter or suite of parameters. We examined how variation in 21 parameters in the equation for gross primary production influenced model sensitivity and uncertainty in terms of two carbon fluxes (net primary productivity and heterotrophic respiration) and two carbon (C) pools (vegetation C and soil C). We set up different parameterizations of TEM across a range of tundra types (tussock tundra, heath tundra, wet sedge tundra, and shrub tundra) in northern Alaska, along a latitudinal transect extending from the coastal plain near Utqiaġvik to the southern foothills of the Brooks Range, to the Seward Peninsula. TEM was most sensitive to parameters related to the temperature regulation of photosynthesis. Model uncertainty was mostly due to parameters related to leaf area, temperature regulation of photosynthesis, and the stomatal responses to ambient light conditions. Our analysis also showed that sensitivity and uncertainty to a given parameter varied spatially. At some sites, model sensitivity and uncertainty tended to be connected to a wider range of parameters, underlining the importance of assessing tundra community processes across environmental gradients or geographic locations. Generally, across sites, the flux of net primary productivity (NPP) and pool of vegetation C had about equal uncertainty, while heterotrophic respiration had higher uncertainty than the pool of soil C. Our study illustrates the complexity inherent in evaluating parameter uncertainty across highly heterogeneous arctic tundra plant communities. It also provides a framework for iteratively testing how newly collected field data related to key parameters may result in more effective forecasting of Arctic change.

54 ENVIRONMENTAL SCIENCES↗

Diffuse Radiation Forcing Constraints on Gross Primary Productivity and Global Terrestrial Evapotranspiration

Abstract The diffuse radiation fertilization effect—the increase in plant productivity in the presence of higher diffuse radiation ( K ↓,d )—is an important yet understudied aspect of atmosphere‐biosphere interactions and can modify the terrestrial carbon, energy, and water budgets. The K ↓,d fertilization effect links the carbon cycle with clouds and aerosols, all of which are large sources of uncertainties for our current understanding of the Earth system and for future climate projections. Here we establish to what extent observational and modeling uncertainty in sunlight's diffuse fraction ( k d ) affects simulated gross primary productivity (GPP) and terrestrial evapotranspiration ( λE ). We find only 48 eddy covariance sites with simultaneous sufficient measurements of K ↓,d with none in the tropical climate zone, making it difficult to constrain this mechanism globally using observations. Using a land modeling framework based on the latest version of the Community Land Model, we find that global GPP ranges from 114 Pg C year −1 when using k d forcing from the Modern‐Era Retrospective analysis for Research and Applications, version 2 reanalysis to a ∼7% higher value of 122 Pg C year −1 when using the Clouds and the Earth's Radiant Energy System satellite product, with especially strong differences apparent over the tropical region (mean increase ∼9%). The differences in λE , although smaller (−0.4%) due to competing changes in shaded and sunlit leaf transpiration, can be greater than regional impacts of individual forcing agents like aerosols. Our results demonstrate the importance of comprehensively and systematically validating the simulated k d by atmosphere modules as well as the response to differences in k d within land modules across Earth System Models.

54 ENVIRONMENTAL SCIENCES↗

Systematic Underestimation of Canopy Conductance Sensitivity to Drought by Earth System Models

The response of vegetation canopy conductance (g c ) to changes in moisture availability ($γ^{m}_{gc}$) is a major source of uncertainty in climate projections. While vegetation typically reduces stomatal conductance during drought, accurately modeling how and to what degree stomata respond to changes in moisture availability at global scales is particularly challenging, because no global scale g c observations exist. Here, we leverage a collection of satellite, reanalysis and station-based near-surface air and surface temperature estimates, which are physically and statistically linked to $γ^{m}_{gc}$ due to the local cooling effect of g c through transpiration, to develop a novel emergent constraint of $γ^{m}_{gc}$ in an ensemble of Earth System Models (ESMs). We find that ESMs systematically underestimate $γ^{m}_{gc}$ by ~33%, particularly in grasslands, croplands, and savannas in semi-arid and bordering regions of the Central United States, Central Europe, Southeastern South America, Southern Africa, Eastern Australia, and parts of East Asia. We show that this underestimation occurs because ESMs inadequately reduce g c when soil moisture decreases. As g c controls carbon, water and energy fluxes, the misrepresentation of modeled $γ^{m}_{gc}$ contributes to biases in ESM projections of gross primary production, transpiration, and temperature during droughts. Our results suggest that the severity and duration of droughts may be misrepresented in ESMs due to the impact of sustained g c on both soil moisture dynamics and the biosphere-atmosphere feedbacks that affect local temperatures and regional weather patterns.

54 ENVIRONMENTAL SCIENCES↗

The Modeled Seasonal Cycles of Surface N 2 O Fluxes and Atmospheric N 2 O

Nitrous oxide (N 2 O) is a greenhouse gas and stratospheric ozone-depleting substance with large and growing anthropogenic emissions. Previous studies identified the influx of N 2 O-depleted air from the stratosphere to partly cause the seasonality in tropospheric N 2 O (aN 2 O), but other contributions remain unclear. Here, we combine surface fluxes from eight land and four ocean models from phase 2 of the Nitrogen/N 2 O Model Intercomparison Project with tropospheric transport modeling to simulate aN 2 O at eight remote air sampling sites for modern and pre-industrial periods. Models show general agreement on the seasonal phasing of zonal-average N 2 O fluxes for most sites, but seasonal peak-to-peak amplitudes differ several-fold across models. The modeled seasonal amplitude of surface aN 2 O ranges from 0.25 to 0.80 ppb (interquartile ranges 21%–52% of median) for land, 0.14–0.25 ppb (17%–68%) for ocean, and 0.28–0.77 ppb (23%–52%) for combined flux contributions. The observed seasonal amplitude ranges from 0.34 to 1.08 ppb for these sites. The stratospheric contributions to aN 2 O, inferred by the difference between the surface-troposphere model and observations, show 16%–126% larger amplitudes and minima delayed by ~1 month compared to Northern Hemisphere site observations. Land fluxes and their seasonal amplitude have increased since the pre-industrial era and are projected to grow further under anthropogenic activities. Our results demonstrate the increasing importance of land fluxes for aN 2 O seasonality. Considering the large model spread, in situ aN 2 O observations and atmospheric transport-chemistry models will provide opportunities for constraining terrestrial and oceanic biosphere models, critical for projecting carbon-nitrogen cycles under ongoing global warming.

54 ENVIRONMENTAL SCIENCES↗

Evidence and attribution of the enhanced land carbon sink

Climate change has been partially mitigated by an increasing net land carbon sink in the terrestrial biosphere; understanding the processes that drive this sink is thus essential for protecting, managing and projecting this important ecosystem service. In this Review, we examine evidence for an enhanced land carbon sink and attribute the observed response to drivers and processes. This sink has doubled from 1.2 ± 0.5 PgC yr -1 in the 1960s to 3.1 ± 0.6 PgC yr -1 in the 2010s. This trend results largely from carbon dioxide fertilization increasing photosynthesis (driving an increase in the annual land carbon sink of >2 PgC globally since 1900), mainly in tropical forest regions, and elevated temperatures reducing cold limitation, mainly at higher latitudes. Continued long-term land carbon sequestration is possible through the end of this century under multiple emissions scenarios, especially if nature-based climate solutions and appropriate ecosystem management are used. In conclusion, a new generation of globally distributed field experiments is needed to improve understanding of future carbon sink potential by measuring belowground carbon release, the response to carbon dioxide enrichment, and long-term shifts in carbon allocation and turnover.

54 ENVIRONMENTAL SCIENCES↗

Climate change increases carbon allocation to leaves in early leaf green-up

Global greening, characterized by an increase in leaf area index (LAI), implies an increase in foliar carbon (C). Whether this increase in foliar C under climate change is due to higher photosynthesis or to higher allocation of C to leaves remains unknown. Here, we explored the trends in foliar C accumulation and allocation during leaf green-up from 2000 to 2017 using satellite-derived LAI and solar-induced chlorophyll fluorescence (SIF) across the Northern Hemisphere. The accumulation of foliar C accelerated in the early green-up period due to both increased photosynthesis and higher foliar C allocation driven by climate change. In the late stage of green-up, however, we detected decreasing trends in foliar C accumulation and foliar C allocation. Such stage-dependent trends in the accumulation and allocation of foliar C are not represented in current terrestrial biosphere models. Further, our results highlight that a better representation of C allocation should be incorporated into models.

54 ENVIRONMENTAL SCIENCES↗

Multi‐hypothesis comparison of Farquhar and Collatz photosynthesis models reveals the unexpected influence of empirical assumptions at leaf and global scales

Abstract Mechanistic photosynthesis models are at the heart of terrestrial biosphere models (TBMs) simulating the daily, monthly, annual and decadal rhythms of carbon assimilation ( A ). These models are founded on robust mathematical hypotheses that describe how A responds to changes in light and atmospheric CO 2 concentration. Two predominant photosynthesis models are in common usage: Farquhar (FvCB) and Collatz (CBGB). However, a detailed quantitative comparison of these two models has never been undertaken. In this study, we unify the FvCB and CBGB models to a common parameter set and use novel multi‐hypothesis methods (that account for both hypothesis and parameter variability) for process‐level sensitivity analysis. These models represent three key biological processes: carboxylation, electron transport, triose phosphate use (TPU) and an additional model process: limiting‐rate selection. Each of the four processes comprises 1–3 alternative hypotheses giving 12 possible individual models with a total of 14 parameters. To broaden inference, TBM simulations were run and novel, high‐resolution photosynthesis measurements were made. We show that parameters associated with carboxylation are the most influential parameters but also reveal the surprising and marked dominance of the limiting‐rate selection process (accounting for 57% of the variation in A vs. 22% for carboxylation). The limiting‐rate selection assumption proposed by CBGB smooths the transition between limiting rates and always reduces A below the minimum of all potentially limiting rates, by up to 25%, effectively imposing a fourth limitation on A . Evaluation of the CBGB smoothing function in three TBMs demonstrated a reduction in global A by 4%–10%, equivalent to 50%–160% of current annual fossil fuel emissions. This analysis reveals a surprising and previously unquantified influence of a process that has been integral to many TBMs for decades, highlighting the value of multi‐hypothesis methods.

54 ENVIRONMENTAL SCIENCES↗

Defining the Sphagnum Core Microbiome across the North American Continent Reveals a Central Role for Diazotrophic Methanotrophs in the Nitrogen and Carbon Cycles of Boreal Peatland Ecosystems

Peat mosses of the genus Sphagnum are ecosystem engineers that frequently predominate over photosynthetic production in boreal peatlands. Sphagnum spp. host diverse microbial communities capable of nitrogen fixation (diazotrophy) and methane oxidation (methanotrophy), thereby potentially supporting plant growth under severely nutrient-limited conditions. Moreover, diazotrophic methanotrophs represent a possible “missing link” between the carbon and nitrogen cycles, but the functional contributions of the Sphagnum-associated microbiome remain in question. A combination of metagenomics, metatranscriptomics, and dual-isotope incorporation assays was applied to investigate Sphagnum microbiome community composition across the North American continent and provide empirical evidence for diazotrophic methanotrophy in Sphagnum-dominated ecosystems. Remarkably consistent prokaryotic communities were detected in over 250 Sphagnum SSU rRNA libraries from peatlands across the United States (5 states, 17 bog/fen sites, 18 Sphagnum species), with 12 genera of the core microbiome comprising 60% of the relative microbial abundance. Additionally, nitrogenase (nifH) and SSU rRNA gene amplicon analysis revealed that nitrogen-fixing populations made up nearly 15% of the prokaryotic communities, predominated by Nostocales cyanobacteria and Rhizobiales methanotrophs. While cyanobacteria comprised the vast majority (>95%) of diazotrophs detected in amplicon and metagenome analyses, obligate methanotrophs of the genus Methyloferula (order Rhizobiales) accounted for one-quarter of transcribed nifH genes. Furthermore, in dual-isotope tracer experiments, members of the Rhizobiales showed substantial incorporation of 13 CH 4 and 15 N 2 isotopes into their rRNA. Our study characterizes the core Sphagnum microbiome across large spatial scales and indicates that diazotrophic methanotrophs, here defined as obligate methanotrophs of the rare biosphere (Methyloferula spp. of the Rhizobiales) that also carry out diazotrophy, play a keystone role in coupling of the carbon and nitrogen cycles in nutrient-poor peatlands.

54 ENVIRONMENTAL SCIENCES↗