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Summary Carbon reserves are distributed throughout plant cells allowing past photosynthesis to fuel current metabolism. In trees, comparing the radiocarbon (Δ¹⁴C) of reserves to the atmospheric bomb spike can trace reserve ages.We synthesized Δ¹⁴C observations of stem reserves in nine tree species, fitting a new process model of reserve building. We asked how the distribution, mixing, and turnover of reserves vary across trees and species. We also explored how stress (drought and aridity) and disturbance (fire and bark beetles) perturb reserves.Given sufficient sapwood, young (< 1 yr) and old (20–60+ yr) reserves were simultaneously present in single trees, including ‘prebomb’ reserves in two conifers. The process model suggested that most reserves are deeply mixed (30.2 ± 21.7 rings) and then respired (2.7 ± 3.5‐yr turnover time). Disturbance strongly increased Δ¹⁴C mean ages of reserves (+15–35 yr), while drought and aridity effects on mixing and turnover were species‐dependent. Fire recovery inSequoia sempervirensalso appears to involve previously unobserved outward mixing of old reserves.Deep mixing and rapid turnover indicate most photosynthate is rapidly metabolized. Yet ecological variation in reserve ages is enormous, perhaps driven by stress and disturbance. Across species, maximum reserve ages appear primarily constrained by sapwood longevity, and thus old reserves are probably widespread. 

Summary Tree rings are considered the gold standard for observing variation in past atmospheric radiocarbon (¹⁴C), yet little attention has been paid to whether different trees record tropospheric¹⁴C evenly. The discovery of Miyake events, rapid increases in¹⁴C production occurring in past millennia, has led to repeated measurements of¹⁴C in tree rings across species and locations. These records demonstrate remarkable synchrony, yet significant variability between samples remains, limiting reliable use of tree rings as a precise indicator of the timing and scale of past¹⁴C production. Understanding how trees and species record tropospheric¹⁴C would improve reconstructions of past¹⁴C production events, refine geochronological control, and because of the pulse‐like nature of past¹⁴C production events, shed light on a fundamental question in tree physiology and dendrochronology: what is the lag time between photosynthesis, storage, and allocation to wood and how might this lag blur isotopic signals in tree rings? Here, we evaluate the implicit assumption that all trees record tropospheric¹⁴C evenly by focusing on the path carbon takes within trees including: phenology of carbon uptake and wood formation, storage and use of nonstructural carbohydrates, and how these factors interact to affect the tropospheric¹⁴C signal in wood. 

ABSTRACT We present a timeseries of¹⁴CO₂for 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. 

Abstract 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. 

Biological soil crusts (biocrusts) are critical components of dryland and other ecosystems worldwide, and are increasingly recognized as novel model ecosystems from which more general principles of ecology can be elucidated. Biocrusts are often diverse communities, comprised of both eukaryotic and prokaryotic organisms with a range of metabolic lifestyles that enable the fixation of atmospheric carbon and nitrogen. However, how the function of these biocrust communities varies with succession is incompletely characterized, especially in comparison to more familiar terrestrial ecosystem types such as forests. We conducted a greenhouse experiment to investigate how community composition and soil-atmosphere trace gas fluxes of CO₂, CH₄, and N₂O varied from early-successional light cyanobacterial biocrusts to mid-successional dark cyanobacteria biocrusts and late-successional moss-lichen biocrusts and as biocrusts of each successional stage matured. Cover type richness increased as biocrusts developed, and richness was generally highest in the late-successional moss-lichen biocrusts. Microbial community composition varied in relation to successional stage, but microbial diversity did not differ significantly among stages. Net photosynthetic uptake of CO₂by each biocrust type also increased as biocrusts developed but tended to be moderately greater (by up to ≈25%) for the mid-successional dark cyanobacteria biocrusts than the light cyanobacterial biocrusts or the moss-lichen biocrusts. Rates of soil C accumulation were highest for the dark cyanobacteria biocrusts and light cyanobacteria biocrusts, and lowest for the moss-lichen biocrusts and bare soil controls. Biocrust CH₄and N₂O fluxes were not consistently distinguishable from the same fluxes measured from bare soil controls; the measured rates were also substantially lower than have been reported in previous biocrust studies. Our experiment, which uniquely used greenhouse-grown biocrusts to manipulate community composition and accelerate biocrust development, shows how biocrust function varies along a dynamic gradient of biocrust successional stages. 

Abstract Nonstructural carbohydrates (NSCs) play a critical role in plant physiology and metabolism, yet we know little about their distribution within individual organs such as the stem. This leaves many open questions about whether reserves deep in the stem are metabolically active and available to support functional processes. To gain insight into the availability of reserves, we measured radial patterns of NSCs over the course of a year in the stemwood of temperate trees with contrasting wood anatomy (ring porous vs diffuse porous). In a subset of trees, we estimated the mean age of soluble sugars within and between different organs using the radiocarbon (14C) bomb spike approach. First, we found that NSC concentrations were the highest and most seasonally dynamic in the outermost stemwood segments for both ring-porous and diffuse-porous trees. However, while the seasonal fluctuation of NSCs was dampened in deeper stemwood segments for ring-porous trees, it remained high for diffuse-porous trees. These NSC dynamics align with differences in the proportion of functional sapwood and the arrangement of vessels between ring-porous and diffuse-porous trees. Second, radial patterns of 14C in the stemwood showed that sugars became older when moving toward the pith. The same pattern was found in the coarse roots. Finally, when taken together, our results highlight how the radial distribution and age of NSCs relate to wood anatomy and suggest that while deeper, and likely older, reserves in the stemwood fluctuated across the seasons, the deepest reserves at the center of the stem were not used to support tree metabolism under usual environmental conditions.