Dynamics in non-structural carbohydrate (NSC) pools may underlie observed drought legacies in tree growth. We assessed how aridity influences the dynamics of different-aged NSC pools in tree sapwood at two sites with differing climate conditions ('wet' vs. 'dry') that also experienced widespread regional drought five years earlier. We used an incubation method to measure radiocarbon (Δ14C) in CO2 respired from Populus tremuloides (aspen) tree rings to evaluate NSC storage and mixing patterns, coupled with measurements of NSC (soluble sugars, starch) concentrations and respired δ13C-CO2. At a wet site, CO2 respired from rings formed during 1962-1967 was only ~ 11 years old, suggesting deep sapwood mixing of NSCs as starch. At a dry site, total NSC was about one-third of wet site totals, maximum ages in deep rings were lower, and ages more rapidly increased in shallow rings then plateaued. These results suggest historically shallower mixing and/or relatively higher consumption of NSCs under dry conditions. Both sites, however, had similar aged NSC (<1 yr) in the most recent six rings, indicative of deep radial mixing following relatively wet conditions during the sampling year. We suggest significant differences in NSC mixing among sites are driven by moisture stress, where aridity reduces NSC reserves and restricts the depth of radial mixing. However, dynamic climate conditions in the southwestern US resulted in more complex radial patterns of sapwood NSC age than previously described. We suggest a novel conceptual framework to understand how moisture variability might influence the dynamics of NSC mixing in the sapwood.

Northern Arizona University, Flagstaff, Arizona, USA, recently installed a MIni CArbon DAting System (MICADAS) with a gas interface system (GIS) for determining the C-14 content of CO2 gas released by the acid dissolution of biogenic carbonates. We compare 48 paired graphite, GIS, and direct carbonate C-14 determinations of individual mollusk shells and echinoid tests. GIS sample sizes ranged between 0.5 and 1.5 mg and span 0.1 to 45.1 ka BP (n = 42). A reduced major axis regression shows a strong relationship between GIS and graphite percent Modern Carbon (pMC) values (m = 1.011; 95% CI [0.997-1.023], R-2 = 0.999) that is superior to the relationship between the direct carbonate and graphite values (m = 0.978; 95% CI [0.959-0.999], R-2 = 0.997). Sixty percent of GIS pMC values are within +/- 0.5 pMC of their graphite counterparts, compared to 26% of direct carbonate pMC values. The precision of GIS analyses is approximately +/- 70 C-14 yrs to 6.5 ka BP and decreases to approximately +/- 130 C-14 yrs at 12.5 ka BP. This precision is on par with direct carbonate and is approximately five times larger than for graphite. Six Plio-Pleistocene mollusk and echinoid samples yield finite ages when analyzed as direct carbonate but yield non-finite ages when analyzed as graphite or as GIS. Our results show that GIS C-14 dating of biogenic carbonates is preferable to direct carbonate C-14 dating and is an efficient alternative to standard graphite C-14 dating when the precision of graphite C-14 dating is not required.

Radiocarbon (∆14C) measurements of nonstructural carbon enable inference on the age and turnover time of stored photosynthate (e.g., sugars, starch), of which the largest pool in trees resides in the main bole. Because of potential issues with extraction-based methods, we introduce an incubation method to capture the ∆14C of nonstructural carbon via respired CO2. In this study, we compared the ∆14C obtained from these incubations with ∆14C from a well-established extraction method, using increment cores from a mature trembling aspen (Populus tremuloides). To understand any potential ∆14C disagreement, the yields from both methods were also benchmarked against the phenol-sulfuric acid concentration assay. We found incubations captured less than 100% of measured sugar and starch carbon, with recovery ranging from ~ 3% in heartwood to 85% in shallow sapwood. However, extractions universally over-yielded (mean 273 ± 101% expected sugar carbon; as high as 480%), where sugars represented less than half of extracted soluble carbon, indicating very poor specificity. While separation of soluble and insoluble nonstructural carbon is ostensibly a strength of extraction based methods, there was also evidence of poor separation of these two fractions in extractions. The ∆14C of respired CO2 and ∆14C from extractions were similar in the sapwood, while extractions resulted in comparatively higher ∆14C (older carbon) in heartwood and bark. Because yield and ∆14C discrepancies were largest in old tissues, incubations may better capture the ∆14C of nonstructural carbon that is actually metabolically available. That is, we suggest extractions include metabolically irrelevant carbon from dead tissues or cells, as well as carbon that is neither sugar nor starch. In contrast, nonstructural carbon captured by extractions must be respired to be measured. We thus suggest incubations of live tissues are a potentially viable, inexpensive, and versatile method to study the ∆14C of metabolically relevant (available) nonstructural carbon.

ABSTRACTWe present a timeseries of 14CO2 for the period 1910–2021 recorded by annual plants collected in the southwestern United States, centered near Flagstaff, Arizona. This timeseries is dominated by five commonly occurring annual plant species in the region, which is considered broadly representative of the southern Colorado Plateau. Most samples (1910–2015) were previously archived herbarium specimens, with additional samples harvested from field experiments in 2015–2021. We used this novel timeseries to develop a smoothed local record with uncertainties for “bomb spike” 14C dating of recent terrestrial organic matter. Our results highlight the potential importance of local records, as we document a delayed arrival of the 1963–1964 bomb spike peak, lower values in the 1980s, and elevated values in the last decade in comparison to the most current Northern Hemisphere Zone 2 record. It is impossible to retroactively collect atmospheric samples, but archived annual plants serve as faithful scribes: samples from herbaria around the Earth may be an under-utilized resource to improve understanding of the modern carbon cycle.

The permafrost region has accumulated organic carbon in cold and waterlogged soils over thousands of years and now contains three times as much carbon as the atmosphere. Global warming is degrading permafrost with the potential to accelerate climate change as increased microbial decomposition releases soil carbon as greenhouse gases. A 19-year time series of soil and ecosystem respiration radiocarbon from Alaska provides long-term insight into changing permafrost soil carbon dynamics in a warmer world. Nine per cent of ecosystem respiration and 23% of soil respiration observations had radiocarbon values more than 50‰ lower than the atmospheric value. Furthermore, the overall trend of ecosystem and soil respiration radiocarbon values through time decreased more than atmospheric radiocarbon values did, indicating that old carbon degradation was enhanced. Boosted regression tree analyses showed that temperature and moisture environmental variables had the largest relative influence on lower radiocarbon values. This suggested that old carbon degradation was controlled by warming/permafrost thaw and soil drying together, as waterlogged soil conditions could protect soil carbon from microbial decomposition even when thawed. Overall, changing conditions increasingly favoured the release of old carbon, which is a definitive fingerprint of an accelerating feedback to climate change as a consequence of warming and permafrost destabilization. This article is part of the Theo Murphy meeting issue 'Radiocarbon in the Anthropocene'.

Abstract Boreal forests harbor as much carbon (C) as the atmosphere and significant amounts of organic nitrogen (N), the nutrient most likely to limit plant productivity in high‐latitude ecosystems. In the boreal biome, the primary disturbance is wildfire, which consumes plant biomass and soil material, emits greenhouse gasses, and influences long‐term C and N cycling. Climate warming and drying is increasing wildfire severity and frequency and is combusting more soil organic matter (SOM). Combustion of surface SOM exposes deeper older layers of accumulated soil material that previously escaped combustion during past fires, here termed legacy SOM. Postfire SOM decomposition and nutrient availability are determined by these layers, but the drivers of legacy SOM decomposition are unknown. We collected soils from plots after the largest fire year on record in the Northwest Territories, Canada, in 2014. We used radiocarbon dating to measure Δ14C (soil age index), soil extractions to quantify N pools and microbial biomass, and a 90‐day laboratory incubation to measure the potential rate of element mineralization and understand patterns and drivers of legacy SOM C decomposition and N availability. We discovered that bulk soil C age predicted C decomposition, where cumulatively, older soil (approximately −450.0‰) produced 230% less C during the incubation than younger soil (~0.0‰). Soil age also predicted C turnover times, with old soil turnover 10 times slower than young soil. We found respired C was younger than bulk soil C, indicating most C enters and leaves relatively quickly, while the older portion remains a stable C sink. Soil age and other indices were unrelated to N availability, but microbial biomass influenced N availability, with more microbial biomass immobilizing soil N pools. Our results stress the importance of legacy SOM as a stable C sink and highlight that soil age drives the pace and magnitude of soil C contributions to the atmosphere between wildfires.

1.ABSTRACTBackgroundDetermining the life-history traits of extinct species is often difficult from skeletal remains alone, limiting the accuracy of studies modeling past ecosystems. However, the analysis of the degraded endogenous bacterial DNA present in paleontological fecal matter (coprolites) may enable the characterization of specific traits such as the host’s digestive physiology and diet. An issue when evaluating the microbial composition of coprolites is the degree to which the microbiome is representative of the host’s original gut community versus the changes that occur in the weeks following deposition due to desiccation. Analyses of paleontological microorganisms are also relevant in the light of recent studies linking the Late Pleistocene and Early Holocene extinctions with modern-day zoonotic pathogen outbreaks.MethodsShotgun sequencing was performed on ancient DNA (aDNA) extracted from coprolites of the Columbian mammoth (Mammuthus Columbi), Shasta ground sloth (Nothrotheriops shastensis) and paleontological bison (Bison sp.) collected from caves on the Colorado Plateau, Southwestern USA. The novel metagenomic classifier MTSv, parameterized for studies of aDNA, was used to assign bacterial taxa to sequencing reads. The resulting bacterial community of coprolites was then compared to those from modern fecal specimens of the African savannah elephant (Loxodonta africana), the brown-throated sloth (Bradypus variegatus) and the modern bison (Bison bison). Both paleontological and modern bison fecal bacterial communities were also compared to those of progressively dried cattle feces to determine whether endogenous DNA from coprolites had a microbiome signal skewed towards aerobic microorganisms typical of desiccated fecal matter.ResultsThe diversity of phyla identified from coprolites was lower than modern specimens. The relative abundance of Actinobacteria was increased in coprolites compared to modern specimens, with fewer Bacteroidetes and Euryarchaeota. Firmicutes had a reduced relative abundance in the mammoth and bison coprolites, compared to the African savanna elephants and modern bison. There was a significant separation of samples in NMDS plots based on their classification as either paleontological or modern, and to a lesser extent, based on the host species. Increasingly dried cattle feces formed a continuum between the modern and paleontological bison samples.ConclusionOur results reveal that any coprolite metagenomes should always be compared to desiccated modern fecal samples from closely related hosts fed a comparable diet to determine the degree to which the coprolite metagenome is a result of desiccation versus true dissimilarities between the modern and paleontological hosts. Also, a large-scale desiccation study including a variety of modern species may shed light on life-history traits of extinct species without close extant relatives, by establishing the proximity of coprolite metagenomes with those from dried modern samples.

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Under increasingly frequent, persistent, and severe drought events, predicting future forest carbon dynamics necessitates quantitative understanding of the physiological processes leading to tree mortality and physiological impairment. The responses of non-structural carbon (NSC; primarily sugars and starch) pools in mature trees is particularly important, as dynamics in NSC interact with hydraulic damage to perturb future tree growth. However, NSC concentration measurements alone are not suUcient to understand the stress responses of tree NSC pools formed over years to decades. Thus, we are using radiocarbon (14C) to quantify the age of NSC stored within, and used by, piñon pine trees exposed to either severe or long-term drought stress at the Sevilleta LTER, in New Mexico, USA. Measuring the age of NSC allows inference on the storage history of a tree, and how different NSC pools may be altered by drought. Experimental plots are subjected to either 0% (control) or 90% reduction in precipitation. A 45% precipitation reduction plot has also been in place since 2009, offering a chance to study the impacts of a decade of drought. We are measuring Δ14C of NSC in twigs, bole sapwood, and coarse roots, as well as in CO2 respired from the bole and branches. Our goal is to quantify the role of different-aged NSC pools across tree organs in driving whole-tree physiological responses to drought. Preliminary results show that the long-term droughted trees store and respire on average younger NSC than control trees. Ongoing drought treatments and sampling will provide additional information on how NSC dynamics in these trees are influenced by drought.

ABSTRACT The direct carbonate procedure for accelerator mass spectrometry radiocarbon (AMS 14 C) dating of submilligram samples of biogenic carbonate without graphitization is becoming widely used in a variety of studies. We compare the results of 153 paired direct carbonate and standard graphite 14 C determinations on single specimens of an assortment of biogenic carbonates. A reduced major axis regression shows a strong relationship between direct carbonate and graphite percent Modern Carbon (pMC) values (m = 0.996; 95% CI [0.991–1.001]). An analysis of differences and a 95% confidence interval on pMC values reveals that there is no significant difference between direct carbonate and graphite pMC values for 76% of analyzed specimens, although variation in direct carbonate pMC is underestimated. The difference between the two methods is typically within 2 pMC, with 61% of direct carbonate pMC measurements being higher than their paired graphite counterpart. Of the 36 specimens that did yield significant differences, all but three missed the 95% significance threshold by 1.2 pMC or less. These results show that direct carbonate 14 C dating of biogenic carbonates is a cost-effective and efficient complement to standard graphite 14 C dating.

Warming of the Arctic can stimulate microbial decomposition and release of permafrost soil carbon (C) as greenhouse gases, and thus has the potential to influence climate change. At the same time, plant growth can be stimulated and offset C release. This study presents a 15‐year time series comprising chamber and eddy covariance measurements of net ecosystem C exchange in a tundra ecosystem in Alaska where permafrost has been degrading due to regional warming. The site was a carbon dioxide source to the atmosphere with a cumulative total loss of 781.6 g C m −2 over the study period. Both gross primary productivity (GPP) and ecosystem respiration ( R eco ) were already likely higher than historical levels such that increases in R eco losses overwhelmed GPP gains in most years. This shift to a net C source to the atmosphere likely started in the early 1990s when permafrost was observed to warm and thaw at the site. Shifts in the plant community occur more slowly and are likely to constrain future GPP increases as compared to more rapid shifts in the microbial community that contribute to increased R eco . Observed rates suggest that cumulative net soil C loss of 4.18–10.00 kg C m −2 —8%–20% of the current active layer soil C pool—could occur from 2020 to the end of the century. This amount of permafrost C loss to the atmosphere represents a significant accelerating feedback to climate change if it were to occur at a similar magnitude across the permafrost region.

Isotopic radiocarbon (Δ 14 C) signatures of ecosystem respiration (Reco) can identify old soil carbon (C) loss and serve as an early indicator of permafrost destabilization in a warming climate. Warming also stimulates plant productivity causing plant respiration to dominate Reco Δ 14 C signatures and potentially obscuring old soil C loss. Here, we investigate how a wide spatio‐temporal gradient of permafrost thaw and plant productivity affects Reco Δ 14 C patterns and isotopic partitioning. Spatial gradients came from a warming experiment with doubling thaw depth and variable biomass, and a vegetation removal manipulation to eliminate plant contributions. We sampled in August and September to capture transitions from high to low plant productivity, decreased surface soil temperature, and relatively small seasonal thaw extensions. We found that surface processes dominate spatial variation in old soil C loss and a process‐based partitioning approach was crucial for constraining old soil C loss. Resampling the same plots in different times of the year revealed that old soil C losses tripled with cooling surface temperature, and the largest old soil C losses were detected when the organic‐to‐mineral soil horizons thawed (∼50–60 cm). We suggest that the measured increase in old soil respiration over the season and when the organic‐to‐mineral horizon thawed, may be explained by mobilization of nitrogen that stimulates microbial decomposition at depth. Our results suggest that soil C in the organic to mineral horizon may be an important source of soil C loss as the entire Arctic region warms and could lead to nonlinearities in projected permafrost climate feedbacks.

Horizontal ICP with large droplet cloud showing YO and neutral Y emission.

Almost half of the global terrestrial soil carbon (C) is stored in the northern circumpolar permafrost region, where air temperatures are increasing two times faster than the global average. As climate warms, permafrost thaws and soil organic matter becomes vulnerable to greater microbial decomposition. Long‐term soil warming of ice‐rich permafrost can result in thermokarst formation that creates variability in environmental conditions. Consequently, plant and microbial proportional contributions to ecosystem respiration may change in response to long‐term soil warming. Natural abundance δ13C and Δ14C of aboveground and belowground plant material, and of young and old soil respiration were used to inform a mixing model to partition the contribution of each source to ecosystem respiration fluxes. We employed a hierarchical Bayesian approach that incorporated gross primary productivity and environmental drivers to constrain source contributions. We found that long‐term experimental permafrost warming introduced a soil hydrology component that interacted with temperature to affect old soil C respiration. Old soil C loss was suppressed in plots with warmer deep soil temperatures because they tended to be wetter. When soil volumetric water content significantly decreased in 2018 relative to 2016 and 2017, the dominant respiration sources shifted from plant aboveground and young soil respiration to old soil respiration. The proportion of ecosystem respiration from old soil C accounted for up to 39% of ecosystem respiration and represented a 30‐fold increase compared to the wet‐year average. Our findings show that thermokarst formation may act to moderate microbial decomposition of old soil C when soil is highly saturated. However, when soil moisture decreases, a higher proportion of old soil C is vulnerable to decomposition and can become a large flux to the atmosphere. As permafrost systems continue to change with climate, we must understand the thresholds that may propel these systems from a C sink to a source.

Boreal forest fires emit large amounts of carbon into the atmosphere primarily through the combustion of soil organic matter 1 – 3 . During each fire, a portion of this soil beneath the burned layer can escape combustion, leading to a net accumulation of carbon in forests over multiple fire events 4 . Climate warming and drying has led to more severe and frequent forest fires 5 – 7 , which threaten to shift the carbon balance of the boreal ecosystem from net accumulation to net loss 1 , resulting in a positive climate feedback 8 . This feedback will occur if organic-soil carbon that escaped burning in previous fires, termed ‘legacy carbon’, combusts. Here we use soil radiocarbon dating to quantitatively assess legacy carbon loss in the 2014 wildfires in the Northwest Territories of Canada 2 . We found no evidence for the combustion of legacy carbon in forests that were older than the historic fire-return interval of northwestern boreal forests 9 . In forests that were in dry landscapes and less than 60 years old at the time of the fire, legacy carbon that had escaped burning in the previous fire cycle was combusted. We estimate that 0.34 million hectares of young forests (<60 years) that burned in the 2014 fires could have experienced legacy carbon combustion. This implies a shift to a domain of carbon cycling in which these forests become a net source—instead of a sink—of carbon to the atmosphere over consecutive fires. As boreal wildfires continue to increase in size, frequency and intensity 7 , the area of young forests that experience legacy carbon combustion will probably increase and have a key role in shifting the boreal carbon balance.

High latitude warming and permafrost thaw will expose vast stores of deep soil organic carbon (SOC) to decomposition. Thaw also changes water movement causing either wetter or drier soil. The fate of deep SOC under different thaw and moisture conditions is unclear. We measured weekly growing‐season δ13C of ecosystem respiration (Recoδ13C) across thaw and moisture conditions (Shallow‐Dry; Deep‐Dry; Deep‐Wet) in a soil warming manipulation. Deep SOC loss was inferred from known δ13C signatures of plant shoot, root, surface soil, and deep soil respiration. In addition, a 2‐year‐old vegetation removal treatment (No Veg) was used to isolate surface and deep SOC decomposition contributions to Reco. In No Veg, seasonal Recoδ13C indicated that deep SOC loss increased as the soil column thawed, while in vegetated areas, root contributions appeared to dominate Reco. The Recoδ13C differences between Shallow‐Dry and Deep‐Dry were significant but surprisingly small. This most likely suggests that, under dry conditions, soil warming stimulates root and surface SOC respiration with a negative 13C signature that opposes the more positive 13C signal from increased deep SOC respiration. In Deep‐Wet conditions, Recoδ13C suggests reduced deep SOC loss but could also reflect altered diffusion or methane (CH4) dynamics. Together, these results demonstrate that frequent Recoδ13C measurements can detect deep SOC loss and that plants confound the signal. In future studies, soil profile δ13C measurements, vegetation removal across thaw gradients, and isotopic effects of CH4 dynamics could further deconvolute deep SOC loss via surface Reco.