Root distributions are typically based on root mass per soil volume. This plant‐focused approach masks the biogeochemical influence of fine roots, which weigh little. We assert that centimeter‐scale root presence‐absence data from soil profiles provide a more soil‐focused approach for probing depth distributions of root‐regolith interfaces, where microsite‐scale processes drive whole‐ecosystem functioning. In 75 soil pits across the continental USA, Puerto Rico, and the Alps, we quantified fine and coarse root presence as deep as 2 m. In 70 of these pits we estimated root mass and created standardized metrics of both data sets to compare their depth distributions. We addressed whether: (a) depth distributions of root presence‐absence data differ from root mass data, thus implying different degrees of root‐regolith interactions with depth; and (b) if root presence or any depth‐dependent differences between these data sets vary predictably with environmental conditions. Presence of fine roots exhibited diverse depth‐dependent patterns; root mass generally declined with depth. In B and C horizons, standardized root presence was greater than standardized root mass; random forest analyses suggest these discrepancies are greater in B horizons with increasing mean annual precipitation and in C horizons with increasing mean annual temperature. Our work suggests that deep in the subsurface, biogeochemical and reactive transport processes result from more numerous root‐regolith interfaces than mass data suggest. We present a new paradigm for discerning patterns in depth distributions of root‐regolith interfaces across multiple biomes and land uses that promotes understanding of the roles of those interfaces in driving key critical zone processes.

Soil carbon-nitrogen (C:N) stoichiometry acts as a control over decomposition and soil organic matter formation and loss, making it a key soil property for understanding ecosystem dynamics and projected ecosystems responses to global environmental change. However, the controls of soil C:N and how they respond to increasing pressures from global change agents are not fully understood. The “foundational” controls on soil C:N, namely plant and microbial C:N, have been used to predict soil C:N, but fail to accurately simulate all ecosystems and may be insufficient for predictions under global environmental change. We present an “emerging” representation of controls of soil C:N that includes plant-microbe-mineral feedbacks that have been shown to regulate soil C:N. We argue that including representation of these emerging drivers in process-based terrestrial biogeochemistry models, which include biological N fixation, mycorrhizae, priming, root exudation of organic acids, and mineralogy (including soil texture, mineral composition, and aggregation), will improve mechanistic representation of soil C:N and associated processes. Such improvements will produce models that will better simulate a variety of ecological states and predict soil C:N when global changes modify plant-microbe-mineral interactions. Here, we align our empirical understanding of controls of soil C:N with those controls represented in models, identifying contexts where emerging drivers might be particularly important to represent (e.g., priming and root exudation in nutrient-limited conditions) and areas of future work. Additionally, we show that implementing emerging drivers of soil C:N results in different simulated outcomes at steady state and in response to elevated atmospheric CO2. Our review and preliminary simulations support the need to incorporate emerging drivers of soil C:N into process-based terrestrial biogeochemistry models, allowing for both theoretical exploration of mechanisms and potentially more accurate predictions of land biogeochemical responses to global change.

Abstract Increased plant growth under elevated carbon dioxide (CO2) slows the pace of climate warming and underlies projections of terrestrial carbon (C) and climate dynamics. However, this important ecosystem service may be diminished by concurrent changes to vegetation carbon‐to‐nitrogen (C:N) ratios. Despite clear observational evidence of increasing foliar C:N under elevated CO2, our understanding of potential ecological consequences of foliar stoichiometric flexibility is incomplete. Here, we illustrate that when we incorporated CO2‐driven increases in foliar stoichiometry into the Community Land Model the projected land C sink decreased two‐fold by the end of the century compared to simulations with fixed foliar chemistry. Further, CO2‐driven increases in foliar C:N profoundly altered Earth's hydrologic cycle, reducing evapotranspiration and increasing runoff, and reduced belowground N cycling rates. These findings underscore the urgency of further research to examine both the direct and indirect effects of changing foliar stoichiometry on soil N cycling and plant productivity.

Nearly 50 y ago, Walker and Syers hypothesized that sources of most terrestrial nutrients shift in dominance from mineral- to organic matter-derived over millennia as soils weather. We investigated how overlaying this soil development framework with vegetation dynamics that can feed back to soil development on relatively short timescales offers insight into ecosystem functioning. To test the hypothesis that forest nutrient economies mediate the nutritional importance of organic matter as mineral weathering proceeds, we paired litterfall decay experiments with soil mineralogical data from diverse forests across the Critical Zone (CZ) Observatory Network, USA. Our findings suggest that dominant sources of tree P may shift from organic matter-bound stocks to minerals as roots expand during the transition from mid to late stages of forest growth and encounter deeper soils that have experienced a lesser degree of weathering. Thus, plants may develop nutritional strategies that do not necessarily rely most heavily on the dominant P form present in an ecosystem, typically driven by stage of soil development, but rather on root proliferation over time, which governs the ability of plants to mine soil volumes at a diversity of depths. Ecosystem P nutrition therefore depends strongly on the interaction between dominant P form and root system growth, particularly as it reflects past land use for both plants and soils. We use these findings to produce a novel framework of vegetative nutrient economics that highlights how root system growth and land use change can influence nutrient transformations and bioavailability, and soil development, across Earth's critical zones.

Rooting depth is an ecosystem trait that determines the extent of soil development and carbon (C) and water cycling. Recent hypotheses propose that human-induced changes to Earth's biogeochemical cycles propagate deeply into Earth's subsurface due to rooting depth changes from agricultural and climate-induced land cover changes. Yet, the lack of a global-scale quantification of rooting depth responses to human activity limits knowledge of hydrosphere-atmosphere-lithosphere feedbacks in the Anthropocene. Here we use land cover data sets to demonstrate that root depth distributions are changing globally as a consequence of agricultural expansion truncating depths above which 99% of root biomass occurs (D99) by ∼60 cm, and woody encroachment linked to anthropogenic climate change extending D99 in other regions by ∼38 cm. The net result of these two opposing drivers is a global reduction of D99 by 5%, or ∼8 cm, representing a loss of ∼11,600 km 3 of rooted volume. Projected land cover scenarios in 2100 suggest additional future D99 shallowing of up to 30 cm, generating further losses of rooted volume of ∼43,500 km 3 , values exceeding root losses experienced to date and suggesting that the pace of root shallowing will quicken in the coming century. Losses of Earth's deepest roots—soil-forming agents—suggest unanticipated changes in fluxes of water, solutes, and C. Two important messages emerge from our analyses: dynamic, human-modified root distributions should be incorporated into earth systems models, and a significant gap in deep root research inhibits accurate projections of future root distributions and their biogeochemical consequences.

Abstract Rooting depth is an ecosystem trait that determines the extent of soil development and carbon (C) and water cycling. Recent hypotheses propose that human‐induced changes to Earth's biogeochemical cycles propagate deeply into Earth's subsurface due to rooting depth changes from agricultural and climate‐induced land cover changes. Yet, the lack of a global‐scale quantification of rooting depth responses to human activity limits knowledge of hydrosphere‐atmosphere‐lithosphere feedbacks in the Anthropocene. Here we use land cover data sets to demonstrate that root depth distributions are changing globally as a consequence of agricultural expansion truncating depths above which 99% of root biomass occurs (D99) by ∼60 cm, and woody encroachment linked to anthropogenic climate change extending D99 in other regions by ∼38 cm. The net result of these two opposing drivers is a global reduction of D99 by 5%, or ∼8 cm, representing a loss of ∼11,600 km3 of rooted volume. Projected land cover scenarios in 2100 suggest additional future D99 shallowing of up to 30 cm, generating further losses of rooted volume of ∼43,500 km3, values exceeding root losses experienced to date and suggesting that the pace of root shallowing will quicken in the coming century. Losses of Earth's deepest roots—soil‐forming agents—suggest unanticipated changes in fluxes of water, solutes, and C. Two important messages emerge from our analyses: dynamic, human‐modified root distributions should be incorporated into earth systems models, and a significant gap in deep root research inhibits accurate projections of future root distributions and their biogeochemical consequences.

Most terrestrial nutrient sources are hypothesized to shift in dominance from mineral- to organic matter (OM)-derived over millennia. We investigated how overlaying this hypothesis with plant rooting dynamics that can feedback to soil development offers insight into ecosystem functioning. To test the hypothesis that the nutritional importance of OM as mineral weathering proceeds is mediated by rooting system nutrient economies that vary with vegetation development, we paired litterfall decay experiments with soil mineralogical data from diverse forests across the Critical Zone (CZ) Observatory Network. We demonstrate that sources of phosphorus shift from OM-bound stocks to minerals as the rooting zone expands during the transition from mid to late stages of forest growth. Root-driven, plant-soil feedbacks thus can prompt inconsistencies with soil development models that posit a unidirectional transition from mineral to organic nutrient dominance, and illuminate how forest growth and land use influence nutrient bioavailability in Earth’s CZ.

Biotically-mediated weathering helps to shape Earth's surface. For example, plants expend carbon (C) to mobilize nutrients in forms whose relative abundances vary with depth. It thus is likely that trees' nutrient acquisition strategies-their investment in rooting systems and exudates-may function differently following disturbance-induced changes in depth of rooting zones and soil nutrient stocks. These changes may persist across centuries. We test the hypothesis that plant C allocation for nutrient acquisition is depth dependent as a function of rooting system development and relative abundances of organic vs. mineral nutrient stocks. We further posit that patterns of belowground C allocation to nutrient acquisition reveal anthropogenic signatures through many decades of forest regeneration. To test this idea, we examined fine root abundances and rooting system C in organic acid exudates and exo-enzymes in tandem with depth distributions of organically- and mineral-bound P stocks. Our design permitted us to estimate C tradeoffs between organic vs. mineral nutrient benefits in paired forests with many similar aboveground traits but different ages: post-agricultural mixed-pine forests and older reference hardwoods. Fine roots were more abundant throughout the upper 2 m in reference forest soils than in regenerating stands. Rooting systems in all forests exhibited depth-dependent C allocations to nutrient acquisition reflecting relative abundances of organic vs. mineral bound P stocks. Further, organic vs. mineral stocks underwent redistribution with historic land use, producing distinct ecosystem nutritional economies. In reference forests, rooting systems are allocating C to relatively deep fine roots and low-C exudation strategies that can increase mobility of mineral-bound P stocks. Regenerating forests exhibit relatively shallower fine root distributions and more diverse exudation strategies reflecting more variable nutrient stocks. We observed these disparities in rooting systems' depth and nutritional mechanisms even though the regenerating forests have attained aboveground biomass stocks similar to those in reference hardwood forests. These distinctions offer plausible belowground mechanisms for observations of continued C sink strength in relatively old forests, and have implications for soil C fates and soil development on timescales relevant to human lifetimes. As such, depth-dependent nutrient returns on plant C investments represent a subtle but consequential signal of the Anthropocene.

Plant, soil, and aquatic microbiomes interact, but scientists often study them independently. Integrating knowledge across these traditionally separate subdisciplines will generate better understanding of microbial ecological properties. Interactions among plant, soil, and aquatic microbiomes, as well as anthropogenic factors, influence important ecosystem processes, including greenhouse gas fluxes, crop production, nonnative species control, and nutrient flux from terrestrial to aquatic habitats. Terrestrial microbiomes influence nutrient retention and particle movement, thereby influencing the composition and functioning of aquatic microbiomes, which, themselves, govern water quality, and the potential for harmful algal blooms. Understanding how microbiomes drive links among terrestrial (plant and soil) and aquatic habitats will inform management decisions influencing ecosystem services. In the present article, we synthesize knowledge of microbiomes from traditionally disparate fields and how they mediate connections across physically separated systems. We identify knowledge gaps currently limiting our abilities to actualize microbiome management approaches for addressing environmental problems and optimize ecosystem services.

Roots and associated microbes generate acid-forming CO2 and organic acids and accelerate mineral weathering deep within Earth’s critical zone (CZ). At the Calhoun CZ Observatory in the USA’s Southern Piedmont, we tested the hypothesis that deforestation-induced deep root losses reduce root- and microbially-mediated weathering agents well below maximum root density (to 5 m), and impart land-use legacies even after ~70 y of forest regeneration. In forested plots, root density declined with depth to 200 cm; in cultivated plots, roots approached zero at depths >70 cm. Below 70 cm, root densities in old-growth forests averaged 2.1 times those in regenerating forests. Modeled root distributions suggest declines in density with depth were steepest in agricultural plots, and least severe in old-growth forests. Root densities influenced biogeochemical environments in multiple ways. Microbial community composition varied with land use from surface horizons to 500 cm; relative abundance of root-associated bacteria was greater in old-growth soils than in regenerating forests, particularly at 100–150 cm. At 500 cm in old-growth forests, salt-extractable organic C (EOC), an organic acid proxy, was 8.8 and 12.5 times that in regenerating forest and agricultural soils, respectively. The proportion of soil organic carbon comprised of EOC was greater in old-growth forests (20.0 ± 2.6%) compared to regenerating forests (2.1 ± 1.1) and agricultural soils (1.9 ± 0.9%). Between 20 and 500 cm, [EOC] increased more with root density in old-growth relative to regenerating forests. At 300 cm, in situ growing season [CO2] was significantly greater in old-growth forests relative to regenerating forests and cultivated plots; at 300 and 500 cm, cultivated soil [CO2] was significantly lower than in forests. Microbially-respired δ13C-CO2 suggests that microbes may rely partially on crop residue even after ~70 y of forest regeneration. We assert that forest conversion to frequently disturbed ecosystems limits deep roots and reduces biotic generation of downward-propagating weathering agents.

Microbial transformation of reactive nitrogen in agricultural fertilizers through nitrification and denitrification is responsible for emissions of greenhouse gases like N2O, NO, and CO2. This study investigates the relationship between the form of nitrogen in fertilizer, the transformations that nitrogen undergoes in the soil, and the corresponding release of N2O and CO2, with the ultimate goal of determining which nitrogen fertilizers contribute the most greenhouse gases to the atmosphere. I applied ammonium, nitrate, and organic nitrogen fertilizers to a cornfield in South Deerfield, MA and measured the N2O and CO2 flux from those fields. In addition, I tested the soils from fertilized plots for nitrate and ammonium content. To further clarify the relationship between nitrate and ammonium fertilizers and greenhouse gas emissions, I also measured the N2O and CO2 flux from soils fertilized and incubated in a lab. In South Deerfield soils, ammonium fertilizer application correlated with high rates of nitrification and high N2O fluxes. This result indicates that in Massachusetts soils and similar soil types, nitrification is the dominant process that produces N2O, and ammonium-based fertilizers contribute more N2O to the atmosphere on a sub-daily scale. In the field, results of CO2 flux tests were inconclusive, but in the lab nitrogen fertilization suppressed CO2 emissions. Further testing would help establish the long-term effects of nitrogen fertilizers on greenhouse gas emissions and the effects in different soil types.