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At least 91 records · Page 5

Variations in Bedrock and Vegetation Cover Modulate Subsurface Water Flow Dynamics of a Mountainous Hillslope

Abstract Predicting the hydrological response of watersheds to climate disturbances requires a detailed understanding of the processes connecting hillslopes and streams. Using a network of soil moisture and temperature sensors, electrical resistivity tomography monitoring, and a weather station we assess the above and below‐ground processes driving the hydrological response of a hillslope during snowmelt and summer monsoon. The transect covers bedrock and vegetation gradients, with a steep upper part characterized by shallow bedrock, and gentle lower part underlain by colluvium. The main vegetation cover is conifers on the upper, and grass and veratrum on the lower part. Combined with a simplified hydrological model, we show that the thin soil layer of the steep slope acts as a preferential flow path, leading to mostly shallow lateral flow, interrupted by vertical flow, mostly at tree locations, and likely facilitated by flow along fractures and roots. Vertical flow and upstream‐driven groundwater dynamics are prevailing at the colluvium, presenting a very different hydrological behavior compared to the upper part. These results show that subsurface structure and features have a strong control on the hydrological response of a hillslope and that those can create considerably varying hydrological dynamics across small spatial scales.

54 ENVIRONMENTAL SCIENCES↗

Representing Soil Microbial Dynamics and Organo‐Mineral Interactions in the E3SM Land Model (ELM‐ReSOM)

Explicit representation of soil microbial processes and interactions with biotic and abiotic processes in Earth System Models (ESMs) remains limited, despite their importance in biogeochemical cycles. To address this gap, which hinders prediction of global biogeochemial cycling and responses to atmospheric conditions, we integrated a microbe- and mineral-surface-explicit model, the Reaction-network-based model of soil organic matter and Microbes (ReSOM), into the Energy Exascale ESM (E3SM) land model (ELM). Here, we describe ELM-ReSOM and show a case study at a conifer forest in California. ELM-ReSOM accurately simulated surface CO 2 fluxes and SOM stocks, demonstrating improved representations of microbial and mineral interactions compared to the default ELM. We examined ELM-ReSOM sensitivity to microbial traits, enzyme properties, and organo-mineral interactions. Microbial traits such as the maximum mortality rate, transporter-density scaling factor, and maximum monomer assimilation rate were strong controllers of heterotrophic respiration, while these microbial traits and enzyme-related properties collectively influenced SOM stocks. Mineral surfaces primarily affected SOM stocks by adsorbing enzymes, thereby limiting depolymerization. Synergies among processes led to stronger impacts of parameters when evaluated together versus separately (i.e., most parameters had greater indirect than direct effects). For example, due to interactions of microbial necromass with mineral surface adsorption, the indirect effect of the maximum microbial mortality rate was 33% larger than its direct effect on SOM stock. Thus, microbial and enzyme dynamics and their interactions with mineral surfaces play critical roles in SOM cycling. Tackling the challenges of microbe-explicit models will advance understanding and modeling of SOM dynamics.

Tao, Jing [Lawrence Berkeley National Laboratory (↗

No carbon storage in growth-limited trees in a semi-arid woodland

Plant survival depends on a balance between carbon supply and demand. When carbon supply becomes limited, plants buffer demand by using stored carbohydrates (sugar and starch). During drought, NSCs (non-structural carbohydrates) may accumulate if growth stops before photosynthesis. This expectation is pervasive, yet few studies have combined simultaneous measurements of drought, photosynthesis, growth, and carbon storage to test this. Using a field experiment with mature trees in a semi-arid woodland, we show that growth and photosynthesis slow in parallel as ψ pd declines, preventing carbon storage in two species of conifer (J. monosperma and P. edulis). During experimental drought, growth and photosynthesis were frequently co-limited. Our results point to an alternative perspective on how plants use carbon that views growth and photosynthesis as independent processes both regulated by water availability.

54 ENVIRONMENTAL SCIENCES↗

Land management explains major trends in forest structure and composition over the last millennium in California’s Klamath Mountains

Significance We provide the first assessment of aboveground live tree biomass in a mixed conifer forest over the late Holocene. The biomass record, coupled with local Native oral history and fire scar records, shows that Native burning practices, along with a natural lightning-based fire regime, promoted long-term stability of the forest structure and composition for at least 1 millennium in a California forest. This record demonstrates that climate alone cannot account for observed forest conditions. Instead, forests were also shaped by a regime of frequent fire, including intentional ignitions by Native people. This work suggests a large-scale intervention could be required to achieve the historical conditions that supported forest resiliency and reflected Indigenous influence.

Knight, Clarke A.↗

Effects of short-interval reburns in the boreal forest on soil bacterial communities compared to long-interval reburns

Abstract Increasing fire frequency in some biomes is leading to fires burning in close succession, triggering rapid vegetation change and altering soil properties. We studied the effects of short-interval (SI) reburns on soil bacterial communities of the boreal forest of northwestern Canada using paired sites (n = 44). Both sites in each pair had burned in a recent fire; one site had burned within the previous 20 years before the recent fire (SI reburn) and the other had not. Paired sites were closely matched in prefire ecosite characteristics, prefire tree species composition, and stand structure. We hypothesized that there would be a significant effect of short vs. long fire-free intervals on community composition and that richness would not be consistently different between paired sites. We found that Blastococcus sp. was consistently enriched in SI reburns, indicating its role as a strongly ‘pyrophilous’ bacterium. Caballeronia sordidicola was consistently depleted in SI reburns. The depletion of this endophytic diazotroph raises questions about whether this is contributing to—or merely reflects—poor conifer seedling recolonization post-fire at SI reburns. While SI reburns had no significant effect on richness, dissimilarity between short- and long-interval pairs was significantly correlated with difference in soil pH, and there were small significant changes in overall community composition.

54 ENVIRONMENTAL SCIENCES↗

Soil microbiome feedbacks during disturbance-driven forest ecosystem conversion

Abstract Disturbances cause rapid changes to forests, with different disturbance types and severities creating unique ecosystem trajectories that can impact the underlying soil microbiome. Pile burning—the combustion of logging residue on the forest floor—is a common fuel reduction practice that can have impacts on forest soils analogous to those following high-severity wildfire. Further, pile burning following clear-cut harvesting can create persistent openings dominated by nonwoody plants surrounded by dense regenerating conifer forest. A paired 60-year chronosequence of burn scar openings and surrounding regenerating forest after clear-cut harvesting provides a unique opportunity to assess whether belowground microbial processes mirror aboveground vegetation during disturbance-induced ecosystem shifts. Soil ectomycorrhizal fungal diversity was reduced the first decade after pile burning, which could explain poor tree seedling establishment and subsequent persistence of herbaceous species within the openings. Fine-scale changes in the soil microbiome mirrored aboveground shifts in vegetation, with short-term changes to microbial carbon cycling functions resembling a postfire microbiome (e.g. enrichment of aromatic degradation genes) and respiration in burn scars decoupled from substrate quantity and quality. Broadly, however, soil microbiome composition and function within burn scar soils converged with that of the surrounding regenerating forest six decades after the disturbances, indicating potential microbial resilience that was disconnected from aboveground vegetation shifts. This work begins to unravel the belowground microbial processes that underlie disturbance-induced ecosystem changes, which are increasing in frequency tied to climate change.

Nelson, Amelia R.↗

Peak radial growth of diffuse-porous species occurs during periods of lower water availability than for ring-porous and coniferous trees

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over 6 years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16–18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 ± 4 days; mean ± 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 ± 19 mm (mean ± SE) during their peak growth than ring-porous and coniferous species (15 ± 35 mm and 30 ± 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.

59 BASIC BIOLOGICAL SCIENCES↗

Imprint of tree species mycorrhizal association on microbial‐mediated enzyme activity and stoichiometry

Abstract Understanding the effects of tree species and their mycorrhizal association on soil processes is critical for predicting the ecosystem consequences of species shifts owing to global change and forest management decisions. While it is well established that forests dominated by different mycorrhizal types can vary in how they cycle carbon (C), nitrogen (N) and phosphorus (P), the degree to which these patterns are driven by microbial‐mediated enzyme activity (EA) and ecoenzymatic stoichiometry (ES) remains elusive. Here, we synthesized the effects of mycorrhizal association on seven soil enzymes involved in microbial C, N and P acquisition and ES using data from 56 peer‐reviewed papers. We found that relative to soil in ectomycorrhizal (EcM) trees, soil in arbuscular mycorrhizal (AM) trees exhibited greater activity of some C acquisition enzymes (e.g. beta‐glucosidase; BG) and higher ecoenzymatic ratios of BG/NAG (N‐acetyl‐glucosaminidase) and BG/AP (acid phosphatase). These results supported that AM trees had rapid C and nutrient turnover rates, inorganic nutrient economics and high soil microbial C limitation. We also found evidence for an organic nutrient economy and greater soil microbial demand for nutrients in EcM trees compared to AM trees. In addition, the effect of mycorrhizal association on the activity of certain soil enzymes and enzymatic stoichiometry (i.e. BG and BG/NAG ratio) appeared to be associated with the differences in soil pH, phylogenetic group (i.e. conifers and broadleaves) and leaf habit (i.e. evergreen and deciduous) between AM and EcM trees. The results from the global meta‐analysis suggested that soil EA and ES appear to play critical roles in shaping the differences in the nutrient economy between AM and EcM tree species, but leaf morphology and soil conditions should be considered in evaluations of soil processes in forests of different mycorrhizal associations. Given that most of the studies in the database were from the temperate and subtropical regions, further research in other biomes is needed to elucidate the underlying mechanisms driving the mycorrhizal effect at the global scale. Read the free Plain Language Summary for this article on the Journal blog.

54 ENVIRONMENTAL SCIENCES↗

Montane forest productivity across a semiarid climatic gradient

Abstract High‐elevation montane forests are disproportionately important to carbon sequestration in semiarid climates where low elevations are dry and characterized by low carbon density ecosystems. However, these ecosystems are increasingly threatened by climate change with seasonal implications for photosynthesis and forest growth. As a result, we leveraged eddy covariance data from six evergreen conifer forest sites in the semiarid western United States to extrapolate the status of carbon sequestration within a framework of projected warming and drying. At colder locations, the seasonal evolution of gross primary productivity (GPP) was characterized by a single broad maximum during the summer that corresponded to snow melt‐derived moisture and a transition from winter dormancy to spring activity. Conversely, winter dormancy was transient at warmer locations, and GPP was responsive to both winter and summer precipitation such that two distinct GPP maxima were separated by a period of foresummer drought. This resulted in a predictable sequence of primary limiting factors to GPP beginning with air temperature in winter and proceeding to moisture and leaf area during the summer. Due to counteracting winter (positive) and summer (negative) GPP responses to warming, leaf area index and moisture availability were the best predictors of annual GPP differences across sites. Overall, mean annual GPP was greatest at the warmest site due to persistent vegetation photosynthetic activity throughout the winter. These results indicate that the trajectory of this region's carbon sequestration will be sensitive to reduced or delayed summer precipitation, especially if coupled to snow drought and earlier soil moisture recession, but summer precipitation changes remain highly uncertain. Given the demonstrated potential for seasonally offsetting responses to warming, we project that decadal semiarid montane forest carbon sequestration will remain relatively stable in the absence of severe disturbance.

Knowles, John F.↗

Wildfire‐induced increases in photosynthesis in boreal forest ecosystems of North America

Abstract Observations of the annual cycle of atmospheric CO 2 in high northern latitudes provide evidence for an increase in terrestrial metabolism in Arctic tundra and boreal forest ecosystems. However, the mechanisms driving these changes are not yet fully understood. One proposed hypothesis is that ecological change from disturbance, such as wildfire, could increase the magnitude and change the phase of net ecosystem exchange through shifts in plant community composition. Yet, little quantitative work has evaluated this potential mechanism at a regional scale. Here we investigate how fire disturbance influences landscape‐level patterns of photosynthesis across western boreal North America. We use Alaska and Canadian large fire databases to identify the perimeters of wildfires, a Landsat‐derived land cover time series to characterize plant functional types (PFTs), and solar‐induced fluorescence (SIF) from the Orbiting Carbon Observatory‐2 (OCO‐2) as a proxy for photosynthesis. We analyze these datasets to characterize post‐fire changes in plant succession and photosynthetic activity using a space‐for‐time approach. We find that increases in herbaceous and sparse vegetation, shrub, and deciduous broadleaf forest PFTs during mid‐succession yield enhancements in SIF by 8–40% during June and July for 2‐ to 59‐year stands relative to pre‐fire controls. From the analysis of post‐fire land cover changes within individual ecoregions and modeling, we identify two mechanisms by which fires contribute to long‐term trends in SIF. First, increases in annual burning are shifting the stand age distribution, leading to increases in the abundance of shrubs and deciduous broadleaf forests that have considerably higher SIF during early‐ and mid‐summer. Second, fire appears to facilitate a long‐term shift from evergreen conifer to broadleaf deciduous forest in the Boreal Plain ecoregion. These findings suggest that increasing fire can contribute substantially to positive trends in seasonal CO 2 exchange without a close coupling to long‐term increases in carbon storage.

Biodiversity & Conservation↗

A functional trait framework for integrating nitrogen‐fixing cover crops into short‐rotation woody crop systems

Abstract Developing approaches to simultaneously maximize short‐rotation woody crop (SRWC) productivity while minimizing footprints associated with intensive management is imperative to profitable and sustainable bioenergy production systems. Intercropping nitrogen (N)‐fixing cover crops in SRWC systems is an overlooked approach to sustainably intensify SRWC production by increasing N availability using less environmentally costly inputs. Here, we discuss how functional traits (e.g., seasonal activity, lifespan, leaf habit, soil exploration) of cover crops and SRWCs may interact through space and time influencing access to light, water, and nutrients to provide a framework for successful integration of cover crops into SRWCs. Next, we summarize the literature on intercropping forest plantations with N‐fixing cover crops to identity research gaps and outline future research needs and opportunities. And then, using empirical N demand and productivity data from SRWCs and cover crop N inputs from the literature, we illustrate how SRWC leaf habit (conifer evergreens and deciduous hardwoods) would influence successful integration of cover crops and potential N fixation. We estimate that integrating cover crops into SRWCs could supply 27% and 72% of the N demand across a 10‐year rotation for an evergreen and a deciduous hardwood, respectively. These figures suggest these integrated SRWC systems may approach a virtual minimal external N input when other biogeochemical cycles are considered. The guiding principles presented here are grounded in ecological theory and provide a framework for sustainable intensification of forest production.

60 APPLIED LIFE SCIENCES↗

Tree water uptake patterns across the globe

Plant water uptake from the soil is a crucial element of the global hydrological cycle and essential for vegetation drought resilience. Yet, knowledge of how the distribution of water uptake depth (WUD) varies across species, climates, and seasons is scarce relative to our knowledge of aboveground plant functions. With a global literature review, we found that average WUD varied more among biomes than plant functional types (i.e. deciduous/evergreen broadleaves and conifers), illustrating the importance of the hydroclimate, especially precipitation seasonality, on WUD. By combining records of rooting depth with WUD, we observed a consistently deeper maximum rooting depth than WUD with the largest differences in arid regions – indicating that deep taproots act as lifelines while not contributing to the majority of water uptake. The most ubiquitous observation across the literature was that woody plants switch water sources to soil layers with the highest water availability within short timescales. Hence, seasonal shifts to deep soil layers occur across the globe when shallow soils are drying out, allowing continued transpiration and hydraulic safety. While there are still significant gaps in our understanding of WUD, the consistency across global ecosystems allows integration of existing knowledge into the next generation of vegetation process models.

54 ENVIRONMENTAL SCIENCES↗

Dynamic regulation of water potential in Juniperus osteosperma mediates ecosystem carbon fluxes

Summary Some plants exhibit dynamic hydraulic regulation, in which the strictness of hydraulic regulation (i.e. iso/anisohydry) changes in response to environmental conditions. However, the environmental controls over iso/anisohydry and the implications of flexible hydraulic regulation for plant productivity remain unknown. InJuniperus osteosperma, a drought‐resistant dryland conifer, we collected a 5‐month growing season time series ofin situ, high temporal‐resolution plant water potential () and stand gross primary productivity (GPP). We quantified the stringency of hydraulic regulation associated with environmental covariates and evaluated how predawn water potential contributes to empirically predicting carbon uptake. Juniperus osteospermashowed less stringent hydraulic regulation (more anisohydric) after monsoon precipitation pulses, when soil moisture and atmospheric demand were high, and corresponded with GPP pulses. Predawn water potential matched the timing of GPP fluxes and improved estimates of GPP more strongly than soil and/or atmospheric moisture, notably resolving GPP underestimation before vegetation green‐up. Flexible hydraulic regulation appears to allowJ. osteospermato prolong soil water extraction and, therefore, the period of high carbon uptake following monsoon precipitation pulses. Water potential and its dynamic regulation may account for why process‐based and empirical models commonly underestimate the magnitude and temporal variability of dryland GPP.

Plant Sciences↗

Local carbon reserves are insufficient for phloem terpene induction during drought in Pinus edulis in response to bark beetle–associated fungi

Stomatal closure during drought inhibits carbon uptake and may reduce a tree's defensive capacity. Limited carbon availability during drought may increase a tree's mortality risk, particularly if drought constrains trees' capacity to rapidly produce defenses during biotic attack. We parameterized a new model of conifer defense using physiological data on carbon reserves and chemical defenses before and after a simulated bark beetle attack in mature Pinus edulis under experimental drought. Attack was simulated using inoculations with a consistent bluestain fungus (Ophiostoma sp.) of Ips confusus, the main bark beetle colonizing this tree, to induce a defensive response. Trees with more carbon reserves produced more defenses but measured phloem carbon reserves only accounted for c. 23% of the induced defensive response. Our model predicted universal mortality if local reserves alone supported defense production, suggesting substantial remobilization and transport of stored resin or carbon reserves to the inoculation site. Our results show that de novo terpene synthesis represents only a fraction of the total measured phloem terpenes in P. edulis following fungal inoculation. Without direct attribution of phloem terpene concentrations to available carbon, many studies may be overestimating the scale and importance of de novo terpene synthesis in a tree's induced defense response.

59 BASIC BIOLOGICAL SCIENCES↗

Evolutionary constraints and climate variability jointly shape starch–sugar balance in woody plants

Nonstructural carbohydrates (NSC) buffer plants against carbon imbalances, yet their partitioning between storage and soluble pools remains elusive at global scales. Here, we compiled a dataset of starch to soluble sugar ratio (St : Su) for 308 woody species across 220 sites world-wide and introduce a dimensionless index that integrates storage and demand while minimizing methodological artifacts. St : Su was strongly associated with growth, identifying it as a key axis of carbon allocation. Foliage consistently exhibited lower St : Su than lignified organs, reflecting a division between transient and conservative pools. Conifers accumulated more starch in foliage but less in stems relative to angiosperms, while leaf habits and mycorrhizal associations further modulated organ-specific strategies. Contrary to expectation, foliar and root St : Su varied little among biomes, but stems exhibited higher ratios in tropical rainforests than in boreal or arid regions, reflecting differences in species composition and adaptive storage under disturbance. Phylogeny constrained stem storage, whereas climatic variability, rather than mean conditions, dominated allocation in leaves and roots. These findings establish St : Su as a robust functional trait linking allocation strategies, growth, and resilience, which can be used to improve vegetation model prediction of forest productivity and mortality under climate variability.

Li, Weibin [Lanzhou Univ. (China)] (ORCID:00000001↗

Photosynthetic capacity in middle-aged larch and spruce acclimates independently to experimental warming and elevated CO 2

Photosynthetic acclimation to both warming and elevated CO 2 of boreal trees remains a key uncertainty in modelling the response of photosynthesis to future climates. We investigated the impact of increased growth temperature and elevated CO 2 on photosynthetic capacity (V cmax and J max ) in mature trees of two North American boreal conifers, tamarack and black spruce. We show that V cmax and J max at a standard temperature of 25°C did not change with warming, while V cmax and J max at their thermal optima (T opt ) and growth temperature (T g ) increased. Moreover, V cmax and J max at either 25°C, T opt or T g decreased with elevated CO 2 . The J max /V cmax ratio decreased with warming when assessed at both T opt and T g but did not significantly vary at 25°C. The J max /V cmax increased with elevated CO 2 at either reference temperature. We found no significant interaction between warming and elevated CO 2 on all traits. If this lack of interaction between warming and elevated CO 2 on the V cmax , J max and J max /V cmax ratio is a general trend, it would have significant implications for improving photosynthesis representation in vegetation models. However, future research is required to investigate the widespread nature of this response in a larger number of species and biomes.

60 APPLIED LIFE SCIENCES↗

Characterization of switchgrass ( Panicum virgatum L.) PvKSL1 as a levopimaradiene/abietadiene‐type diterpene synthase

Abstract The diverse class of plant diterpenoid metabolites serves important functions in mediating growth, chemical defence, and ecological adaptation. In major monocot crops, such as maize (Zea mays), rice (Oryza sativa), and barley (Hordeum vulgare), diterpenoids function as core components of biotic and abiotic stress resilience. Switchgrass (Panicum virgatum) is a perennial grass valued as a stress‐resilient biofuel model crop. Previously we identified an unusually large diterpene synthase family that produces both common and species‐specific diterpenoids, several of which accumulate in response to abiotic stress. Here, we report discovery and functional characterization of a previously unrecognized monofunctional class I diterpene synthase (PvKSL1) viain vivoco‐expression assays with different copalyl pyrophosphate (CPP) isomers, structural and mutagenesis studies, as well as genomic and transcriptomic analyses. In particular, PvKSL1 convertsent‐CPP intoent‐abietadiene,ent‐palustradiene,ent‐levopimaradiene, andent‐neoabietadiene via a 13‐hydroxy‐8(14)‐ent‐abietene intermediate. Notably, although featuring a distinctent‐stereochemistry, this product profile is near‐identical to bifunctional (+)‐levopimaradiene/abietadiene synthases occurring in conifer trees. PvKSL1 has three of four active site residues previously shown to control (+)‐levopimaradiene/abietadiene synthase catalytic specificity. However, mutagenesis studies suggest a distinct catalytic mechanism in PvKSL1. Genome localization ofPvKSL1distant from other diterpene synthases, and its phylogenetic distinctiveness from known abietane‐forming diterpene synthases, support an independent evolution of PvKSL1 activity. Albeit at low levels,PvKSL1gene expression predominantly in roots suggests a role of diterpenoid formation in belowground tissue. Together, these findings expand the known chemical and functional space of diterpenoid metabolism in monocot crops.

Plant Sciences↗

Blodgett Forest CA Warming Experiment Soil Gas Well Data March 2014 to June 2017

These are soil gas data from the whole soil profile warming experiment at the University of California Blodgett Forest Research Station, in the Sierra Nevada foothills near Georgetown, CA at 1370 m above sea level. Mean annual precipitation is 1774 mm with most of it occurring from November through April and mean annual temperature is about 12.5°C . The experiment is in a thinned 80-year-old stand of mixed conifers including ponderosa pine (Pinus ponderosa), sugar pine (Pinus lambertiana), incense cedar (Calodefrus decurrens), white fir (Abies concolor), and douglas fir (Pseudotsuga menziesii). The soils are Holland series: fine-loamy, mixed, superactive, mesic Ultic Haploxeralfs of granitic origin with thick, >5 cm O horizons, minimal carbonates , and a pH that ranges from 5.6 to 6.5. The warming experiment warms the top meter of the soil profile to +4°C above ambient. These data include depth-resolved measurements of CO2 concentrations, 13CO2, apparent respiration quotients, soil temperature, and soil moisture (VWC). These data can be used to model CO2 production rates across the soil profile or to track how the sources and processes contributing to soil respiration change seasonally and with warming.

13C↗