Fast-decaying plant litter enhances soil carbon in temperate forests but not through microbial physiological traits.

Conceptual and empirical advances in soil biogeochemistry have challenged long-held assumptions about the role of soil micro-organisms in soil organic carbon (SOC) dynamics; yet, rigorous tests of emerging concepts remain sparse. Recent hypotheses suggest that microbial necromass production links plant inputs to SOC accumulation, with high-quality (i.e., rapidly decomposing) plant litter promoting microbial carbon use efficiency, growth, and turnover leading to more mineral stabilization of necromass. We test this hypothesis experimentally and with observations across six eastern US forests, using stable isotopes to measure microbial traits and SOC dynamics. Here we show, in both studies, that microbial growth, efficiency, and turnover are negatively (not positively) related to mineral-associated SOC. In the experiment, stimulation of microbial growth by high-quality litter enhances SOC decomposition, offsetting the positive effect of litter quality on SOC stabilization. We suggest that microbial necromass production is not the primary driver of SOC persistence in temperate forests. Factors such as microbial necromass origin, alternative SOC formation pathways, priming effects, and soil abiotic properties can strongly decouple microbial growth, efficiency, and turnover from mineral-associated SOC.

Biological mechanisms may contribute to soil carbon saturation patterns.

Increasing soil organic carbon (SOC) storage is a key strategy to mitigate rising atmospheric CO2 , yet SOC pools often appear to saturate, or increase at a declining rate, as carbon (C) inputs increase. Soil C saturation is commonly hypothesized to result from the finite amount of reactive mineral surface area available for retaining SOC, and is accordingly represented in SOC models as a physicochemically determined SOC upper limit. However, mineral-associated SOC is largely microbially generated. In this perspective, we present the hypothesis that apparent SOC saturation patterns could emerge as a result of ecological constraints on microbial biomass-for example, via competition or predation-leading to reduced C flow through microbes and a reduced rate of mineral-associated SOC formation as soil C inputs increase. Microbially explicit SOC models offer an opportunity to explore this hypothesis, yet most of these models predict linear microbial biomass increases with C inputs and insensitivity of SOC to input rates. Synthesis of 54 C addition studies revealed constraints on microbial biomass as C inputs increase. Different hypotheses limiting microbial density were embedded in a three-pool SOC model without explicit limits on mineral surface area. As inputs increased, the model demonstrated either no change, linear, or apparently saturating increases in mineral-associated and particulate SOC pools. Taken together, our results suggest that microbial constraints are common and could lead to reduced mineral-associated SOC formation as input rates increase. We conclude that SOC responses to altered C inputs-or any environmental change-are influenced by the ecological factors that limit microbial populations, allowing for a wider range of potential SOC responses to stimuli. Understanding how biotic versus abiotic factors contribute to these patterns will better enable us to predict and manage soil C dynamics.

Root-derived inputs are major contributors to soil carbon in temperate forests, but vary by mycorrhizal type.

Roots promote the formation of slow-cycling soil carbon (C), yet we have a limited understanding of the magnitude and controls on this flux. We hypothesised arbuscular mycorrhizal (AM)- and ectomycorrhizal (ECM)-associated trees would exhibit differences in root-derived C accumulation in the soil, and that much of this C would be transferred into mineral-associated pools. We installed δ13 C-enriched ingrowth cores across mycorrhizal gradients in six Eastern U.S. forests (n = 54 plots). Overall, root-derived C was 54% greater in AM versus ECM-dominated plots. This resulted in nearly twice as much root-derived C in putatively slow-cycling mineral-associated pools in AM compared to ECM plots. Given that our estimates of root-derived inputs were often equal to or greater than leaf litter inputs, our results suggest that variation in root-derived soil C accumulation due to tree mycorrhizal dominance may be a key control of soil C dynamics in forests.

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2.

Atmospheric carbon dioxide concentration ([CO2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2 ] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2 ]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2 ] (iCO2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Nitrogen cycling microbiomes are structured by plant mycorrhizal associations with consequences for nitrogen oxide fluxes in forests.

Volatile nitrogen oxides (N2 O, NO, NO2 , HONO, …) can negatively impact climate, air quality, and human health. Using soils collected from temperate forests across the eastern United States, we show microbial communities involved in nitrogen (N) cycling are structured, in large part, by the composition of overstory trees, leading to predictable N-cycling syndromes, with consequences for emissions of volatile nitrogen oxides to air. Trees associating with arbuscular mycorrhizal (AM) fungi promote soil microbial communities with higher N-cycle potential and activity, relative to microbial communities in soils dominated by trees associating with ectomycorrhizal (ECM) fungi. Metagenomic analysis and gene expression studies reveal a 5 and 3.5 times greater estimated N-cycle gene and transcript copy numbers, respectively, in AM relative to ECM soil. Furthermore, we observe a 60% linear decrease in volatile reactive nitrogen gas flux (NOy  ≡ NO, NO2 , HONO) as ECM tree abundance increases. Compared to oxic conditions, gas flux potential of N2 O and NO increase significantly under anoxic conditions for AM soil (30- and 120-fold increase), but not ECM soil-likely owing to small concentrations of available substrate ( NO 3 - ) in ECM soil. Linear mixed effects modeling shows that ECM tree abundance, microbial process rates, and geographic location are primarily responsible for variation in peak potential NOy flux. Given that nearly all tree species associate with either AM or ECM fungi, our results indicate that the consequences of tree species shifts associated with global change may have predictable consequences for soil N cycling.

Tree mycorrhizal type predicts within-site variability in the storage and distribution of soil organic matter.

Forest soils store large amounts of carbon (C) and nitrogen (N), yet how predicted shifts in forest composition will impact long-term C and N persistence remains poorly understood. A recent hypothesis predicts that soils under trees associated with arbuscular mycorrhizas (AM) store less C than soils dominated by trees associated with ectomycorrhizas (ECM), due to slower decomposition in ECM-dominated forests. However, an incipient hypothesis predicts that systems with rapid decomposition-e.g. most AM-dominated forests-enhance soil organic matter (SOM) stabilization by accelerating the production of microbial residues. To address these contrasting predictions, we quantified soil C and N to 1 m depth across gradients of ECM-dominance in three temperate forests. By focusing on sites where AM- and ECM-plants co-occur, our analysis controls for climatic factors that covary with mycorrhizal dominance across broad scales. We found that while ECM stands contain more SOM in topsoil, AM stands contain more SOM when subsoil to 1 m depth is included. Biomarkers and soil fractionations reveal that these patterns are driven by an accumulation of microbial residues in AM-dominated soils. Collectively, our results support emerging theory on SOM formation, demonstrate the importance of subsurface soils in mediating plant effects on soil C and N, and indicate that shifts in the mycorrhizal composition of temperate forests may alter the stabilization of SOM.

Plant-microbial competition for nitrogen increases microbial activities and carbon loss in invaded soils.

Many invasive plant species show high rates of nutrient acquisition relative to their competitors. Yet the mechanisms underlying this phenomenon, and its implications for ecosystem functioning, are poorly understood, particularly in nutrient-limited systems. Here, we test the hypothesis that an invasive plant species (Microstegium vimineum) enhances its rate of nitrogen (N) acquisition by outcompeting soil organic matter-degrading microbes for N, which in turn accelerates soil N and carbon (C) cycling. We estimated plant cover as an indicator of plant N acquisition rate and quantified plant tissue N, soil C and N content and transformations, and extracellular enzyme activities in invaded and uninvaded plots. Under low ambient N availability, invaded plots had 77% higher plant cover and lower tissue C:N ratios, suggesting that invasion increased rates of plant N acquisition. Concurrent with this pattern, we observed significantly higher mass-specific enzyme activities in invaded plots as well as 71% higher long-term N availability, 21% lower short-term N availability, and 16% lower particulate organic matter N. A structural equation model showed that these changes were interrelated and associated with 27% lower particulate organic matter C in invaded areas. Our findings suggest that acquisition of N by this plant species enhances microbial N demand, leading to an increased flux of N from organic to inorganic forms and a loss of soil C. We conclude that high N acquisition rates by invasive plants can drive changes in soil N cycling that are linked to effects on soil C.

Grass invasion effects on forest soil carbon depend on landscape-level land use patterns.

Plant invasions can alter the quality and quantity of detrital and root-derived inputs entering a system, thereby influencing the activities of microbial decomposers and affecting the soil carbon cycle. The effect of these inputs on soil carbon storage is often conflicting, suggesting strong context dependency in the plant-decomposer relationship. Whether there is a generalizable pattern that explains this dependency remains relatively unexplored. Here, we (1) examine how invasion by the exotic grass Microstegium vimineum affects carbon cycling across a land use gradient, and (2) evaluate the importance of inorganic nitrogen availability and other environmental variables for explaining patterns in soil carbon. Using paired invaded and uninvaded plots, we quantified invasion effects on belowground carbon pools, extracellular enzyme activities, and native leaf litter decomposition in forests embedded in an urban, agricultural, or forested landscape matrix. Compared to the urban matrix, invasion-associated declines in total soil organic carbon in the forested and agricultural landscapes were 3.5 and 2.5 times greater, respectively. Inorganic nitrogen availability and M. vimineum biomass interacted to explain these patterns: when both nitrogen availability and M. vimineum biomass were high, invaded soils exhibited higher total organic carbon, unchanged particulate organic matter carbon, and higher mineral-associated organic matter carbon compared to adjacent uninvaded soils. Consistent with these patterns, activities of carbon-mineralizing enzymes were lower in invaded than in uninvaded soils when both nitrogen availability and M. vimineum biomass were high. By contrast,. decomposition of native leaf litter was faster when inorganic nitrogen availability and M. vimineum biomass were high. Our findings suggest that, although this invader may accelerate carbon cycling in forest soils, its effects on soil carbon storage largely depend on nitrogen availability and invader biomass, which can be altered by landscape-level patterns of land use. Additional research is needed to determine whether land use or other broad-scale processes such as atmospheric nitrogen deposition can explain context dependence in plant invasion effects on other ecosystem processes.

Bird-window collisions in the summer breeding season.

Birds that reside in urban settings face numerous human-related threats to survival, including mortality from bird-window collisions (BWCs). Our current understanding of this issue has largely been driven by data collected during spring and fall migration, and patterns of collision mortality during the summer breeding season remain relatively unexplored. We assessed BWCs during four breeding seasons (2009-2012) at a site in northwestern Illinois, USA, by comparing the abundance, richness, migratory class, and age of the species living around buildings to species mortally wounded by window collisions. We also systematically assessed the daily timing of BWCs throughout the breeding season. We documented BWCs in 4 of 25 (16%) species and 7 of 21 (33%) species in 2009 and 2010, respectively. The relationship between BWCs and abundance depended on age. For adults, BWCs were highest in the least abundant species, e.g., Red-eyed Vireo (Vireo olivaceus), and lowest in species with high abundance values, e.g., Chipping Sparrow (Spizella passerina). For juveniles, mortality was greatest for the most abundant species, and the American Robin (Turdus migratorius) accounted for 62% of all juvenile carcasses. Early in the breeding season, collision mortality was restricted to adults of Long-distance Migrants, whereas juveniles of all three migratory guilds (Long-distance and Short-distance Migrants and Permanent Residents) died at windows from late June through early August. Daily mortality for all species was highest between sunrise-1600 h and lowest from 1600 h-sunrise the next day. Generally, the species observed as carcasses matched birds considered a 'high risk' for BWCs, e.g., Ruby-throated Hummingbird (Archilochus colubris), and those considered 'low risk' were not observed as carcasses, e.g., Blue-gray Gnatcatcher (Polioptila caerulea). Our results suggest that the number of BWCs during the breeding season does not necessarily increase with abundance, but rather appears related to variation among species and age classes, which may have important implications on the population health of affected species. The mechanisms driving these differences are unknown, but may be related reproductive behavior, flight speed, distance movements, and dispersal patterns.