Jared L. DeForest

Drought during canopy development has lasting effect on annual carbon balance in a deciduous temperate forest

* Climate change projections predict an intensifying hydrologic cycle and an increasing frequency of droughts, yet quantitative understanding of the effects on ecosystem carbon exchange remains limited. * Here, the effect of contrasting precipitation and soil moisture dynamics were evaluated on forest carbon exchange using 2 yr of eddy covariance and microclimate data from a 50-yr-old mixed oak woodland in northern Ohio, USA. * The stand accumulated 40% less carbon in a year with drought between bud-break and full leaf expansion (354 +/- 81 g C m(-2) yr(-1) in 2004 and 252 +/- 45 g C m(-2) yr(-1) in 2005). This was caused by greater suppression of gross ecosystem productivity (GEP; 16% = 200 g) than of ecosystem respiration (ER; 11% = 100 g) by drought. Suppressed GEP was traced to lower leaf area, lower apparent quantum yield and lower canopy conductance. The moisture sensitivity of ER may have been mediated by GEP. * The results highlight the vulnerability of the ecosystem to even a moderate drought, when it affects a critical aspect of development. Although the drought was preceded by rain, the storage capacity of the soil seemed limited to 1-2 wk, and therefore droughts longer than this are likely to impair productivity in the region.

Soil microbial responses to elevated phosphorus and pH in acidic temperate deciduous forests

Although northern temperate forests are generally not considered phosphorus (P) limited, ecosystem P limitation may occur on highly weathered or strongly acidic soils where bioavailable inorganic P is low. In such environments, soil organisms may compensate by increasing the utilization of organic P via the production of extracellular enzymes to prevent limitation. In this study, we experimentally increased available P and/or pH in several acidic eastern deciduous forests underlain by glaciated and unglaciated soils in eastern Ohio, USA. We hypothesized that where inorganic P is low; soil microbes are able to access organic P by increasing production of phosphatase enzymes, thereby overcoming biogeochemical P limitations. We measured surface soil for: available P pools, N mineralization and nitrification rates, total C and N, enzymes responsible for C, N, and P hydrolysis, and microbial community composition (PLFA). Increasing surface soil pH a whole unit had little effect on microbial community composition, but increased N cycling rates in unglaciated soils. Phosphorus additions suppressed phosphatase activities over 60% in the unglaciated soils but were unchanged in the glaciated soils. All treatments had minimal influence on microbial biomass, but available pools of P strongly correlated with microbial composition. Microbes may be dependent on sources of organic P in some forest ecosystems and from a microbial perspective soil pH might be less important overall than P availability. Although our sampling was conducted less than 1 year after treatment initiation, microbial community composition was strongly influenced by available P pools and these effects may be greater than short-term increases in soil pH.

Root morphology and mycorrhizal symbioses together shape nutrient foraging strategies of temperate trees

Significance Plant growth requires acquisition of soil nutrients in a patchy environment. Nutrient patches may be actively foraged by symbioses comprising roots and mycorrhizal fungi. Here, we show that thicker root tree species (e.g., tulip poplar, pine) respond weakly or not at all to nutrient heterogeneity. In contrast, thinner root tree species readily respond by selectively growing roots [arbuscular mycorrhizal trees (e.g., maple)] or mycorrhizal fungal hyphae [ectomycorrhizal trees (e.g., oak)] in nutrient-rich “hotspots.” Our results thus indicate predictable patterns of nutrient foraging among tree species with contrasting mycorrhiza types and root morphologies. These findings can pave the way for a more holistic understanding of root-microbial function, which is critical to plant growth and biogeochemical cycles in forested ecosystems. Photosynthesis by leaves and acquisition of water and minerals by roots are required for plant growth, which is a key component of many ecosystem functions. Although the role of leaf functional traits in photosynthesis is generally well understood, the relationship of root functional traits to nutrient uptake is not. In particular, predictions of nutrient acquisition strategies from specific root traits are often vague. Roots of nearly all plants cooperate with mycorrhizal fungi in nutrient acquisition. Most tree species form symbioses with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi. Nutrients are distributed heterogeneously in the soil, and nutrient-rich “hotspots” can be a key source for plants. Thus, predicting the foraging strategies that enable mycorrhizal root systems to exploit these hotspots can be critical to the understanding of plant nutrition and ecosystem carbon and nutrient cycling. Here, we show that in 13 sympatric temperate tree species, when nutrient availability is patchy, thinner root species alter their foraging to exploit patches, whereas thicker root species do not. Moreover, there appear to be two distinct pathways by which thinner root tree species enhance foraging in nutrient-rich patches: AM trees produce more roots, whereas EM trees produce more mycorrhizal fungal hyphae. Our results indicate that strategies of nutrient foraging are complementary among tree species with contrasting mycorrhiza types and root morphologies, and that predictable relationships between below-ground traits and nutrient acquisition emerge only when both roots and mycorrhizal fungi are considered together.

Single-walled carbon nanotubes alter soil microbial community composition.

Recent developments in nanotechnology may lead to the release of nanomaterials into the natural environment, such as soils, with largely unknown consequences. We investigated the effects of single-walled carbon nanotubes (SWCNTs), one of the most widely used nanomaterials, on soil microbial communities by incubation of soils to which powder or suspended forms of SWCNTs were added (0.03 to 1 mg g(-1) soil). To determine changes in soil microbial community composition, phospholipid fatty acid (PLFA) profiles were analyzed at 25th day of the incubation experiment. The biomass of major microbial groups including Gram-positive and Gram-negative bacteria, and fungi showed a significant negative relationship with SWCNT concentration, while the relative abundance of bacteria showed a positive relationship with SWCNT concentration. Furthermore, soils under distinct concentrations of SWCNT treatments had PLFA profiles that were significantly different from one another. Our results indicate that the biomass of a broad range of soil microbial groups is negatively related with SWCNT concentration and upon entry into soils, SWCNTs may alter microbial community composition. Our results may serve as foundation for scientific guideline on regulating the discharge of nanomaterials such as SWCNTs to the soil ecosystem.

Effects of timber harvest on carbon pools in Ozark forests

We quantified and compared carbon (C) pools at a Missouri Ozark experimental forest 8 years after different har- vest treatments. Total C pools were 182, 170, and 130 Mg Cha -1 for the control (no-harvest management; NHM), single- tree, uneven-age management (UAM), and clearcut even-age management (EAM) stands, respectively. Harvesting reduced the live tree C pool by 31% in the UAM, 93% in EAM stands, and increased the coarse woody debris (CWD) C pool by 50% in UAM and 176% for EAM, compared with NHM stands. UAM significantly (p = 0.02) increased the mineral soil C pool by 14%, whereas EAM had no effect. More interestingly, the distribution of C among various components (i.e., live, dead wood, CWD, litter, and soil) ranged from 0.7% to 29% on NHM stands and from 0.1% to 43% on EAM stands. Soil ni- trogen (N) (%) was significantly correlated with soil C (%) in the UAM stands, whereas soil temperature was negatively re- lated to live tree C. Soil N (%) and canopy cover were significantly correlated with live tree and soil C (%) pools at EAM stands. Our results revealed that the largest C pool in these forests was living trees. The soil and CWD C pool sizes suggest the importance of dynamics of decaying harvest debris, which influences N retention.

Mycorrhizal Response to Experimental pH and P Manipulation in Acidic Hardwood Forests

Many temperate forests of the Northeastern United States and Europe have received significant anthropogenic acid and nitrogen (N) deposition over the last century. Although temperate hardwood forests are generally thought to be N-limited, anthropogenic deposition increases the possibility of phosphorus (P) limiting productivity in these forest ecosystems. Moreover, inorganic P availability is largely controlled by soil pH and biogeochemical theory suggests that forests with acidic soils (i.e., <pH 5) are particularly vulnerable to P limitation. Results from previous studies in these systems are mixed with evidence both for and against P limitation. We hypothesized that shifts in mycorrhizal colonization and community structure help temperate forest ecosystems overcome an underlying P limitation by accessing mineral and organic P sources that are otherwise unavailable for direct plant uptake. We examined arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) communities and soil microbial activity in an ecosystem-level experiment where soil pH and P availability were manipulated in mixed deciduous forests across eastern Ohio, USA. One year after treatment initiation, AM root biomass was positively correlated with the most available P pool, resin P, while AM colonization was negatively correlated. In total, 15,876 EcM root tips were identified and assigned to 26 genera and 219 operational taxonomic units (97% similarity). Ectomycorrhizal richness and root tip abundance were negatively correlated with the moderately available P pools, while the relative percent of tips colonized by Ascomycetes was positively correlated with soil pH. Canonical correspondence analysis revealed regional, but not treatment, differences in AM communities, while EcM communities had both treatment and regional differences. Our findings highlight the complex interactions between mycorrhizae and the soil environment and further underscore the fact that mycorrhizal communities do not merely reflect the host plant community.

Soil Microbial Responses to Elevated CO2 and O3 in a Nitrogen-Aggrading Agroecosystem

Climate change factors such as elevated atmospheric carbon dioxide (CO2) and ozone (O3) can exert significant impacts on soil microbes and the ecosystem level processes they mediate. However, the underlying mechanisms by which soil microbes respond to these environmental changes remain poorly understood. The prevailing hypothesis, which states that CO2- or O3-induced changes in carbon (C) availability dominate microbial responses, is primarily based on results from nitrogen (N)-limiting forests and grasslands. It remains largely unexplored how soil microbes respond to elevated CO2 and O3 in N-rich or N-aggrading systems, which severely hinders our ability to predict the long-term soil C dynamics in agroecosystems. Using a long-term field study conducted in a no-till wheat-soybean rotation system with open-top chambers, we showed that elevated CO2 but not O3 had a potent influence on soil microbes. Elevated CO2 (1.5×ambient) significantly increased, while O3 (1.4×ambient) reduced, aboveground (and presumably belowground) plant residue C and N inputs to soil. However, only elevated CO2 significantly affected soil microbial biomass, activities (namely heterotrophic respiration) and community composition. The enhancement of microbial biomass and activities by elevated CO2 largely occurred in the third and fourth years of the experiment and coincided with increased soil N availability, likely due to CO2-stimulation of symbiotic N2 fixation in soybean. Fungal biomass and the fungi∶bacteria ratio decreased under both ambient and elevated CO2 by the third year and also coincided with increased soil N availability; but they were significantly higher under elevated than ambient CO2. These results suggest that more attention should be directed towards assessing the impact of N availability on microbial activities and decomposition in projections of soil organic C balance in N-rich systems under future CO2 scenarios.

Mycorrhizal fungi and roots are complementary in foraging within nutrient patches

The roots of the majority of tree species are associated with either arbuscular mycorrhizal (AM) or ectomycorrhizal (EM) fungi. The absorptive roots of tree species also vary widely in their diameter. The linkages between root thickness, mycorrhiza type and nutrient foraging are poorly understood. We conducted a large root ingrowth experiment in the field to investigate how absorptive roots of varying thickness and their associated fungi (AM vs. EM) exploit different nutrient patches (inorganic and organic) in a common garden. In nutrient-rich patches, thin-root tree species more effectively proliferated absorptive roots than thick-root tree species, whereas thick-root tree species proliferated more mycorrhizal fungal biomass than thin-root tree species. Moreover, nutrient patches enriched with organic materials resulted in greater root and mycorrhizal fungal proliferation compared to those enriched with inorganic nutrients. Irrespective of root morphology, AM tree species had higher root foraging precision than mycorrhizal hyphae foraging precision within organic patches, whereas EM tree species exhibited the opposite. Our findings that roots and mycorrhizal fungi are complementary in foraging within nutrient patches provide new insights into species coexistence and element cycling in terrestrial ecosystems.

Mycorrhizal fungal communities respond to experimental elevation of soil pH and P availability in temperate hardwood forests

Many forests are affected by chronic acid deposition, which can lower soil pH and limit the availability of nutrients such as phosphorus (P), but the response of mycorrhizal fungi to changes in soil pH and P availability and how this affects tree acquisition of nutrients is not well understood. Here, we describe an ecosystem-level manipulation in 72 plots, which increased pH and/or P availability across six forests in Ohio, USA. Two years after treatment initiation, mycorrhizal fungi on roots were examined with molecular techniques, including 454-pyrosequencing. Elevating pH significantly increased arbuscular mycorrhizal (AM) fungal colonization and total fungal biomass, and affected community structure of AM and ectomycorrhizal (EcM) fungi, suggesting that raising soil pH altered both mycorrhizal fungal communities and fungal growth. AM fungal taxa were generally negatively correlated with recalcitrant P pools and soil enzyme activity, whereas EcM fungal taxa displayed variable responses, suggesting that these groups respond differently to P availability. Additionally, the production of extracellular phosphatase enzymes in soil decreased under elevated pH, suggesting a shift in functional activity of soil microbes with pH alteration. Thus, our findings suggest that elevating pH increased soil P availability, which may partly underlie the mycorrhizal fungal responses we observed.

Diminished Soil Quality in an Old-Growth, Mixed Mesophytic Forest Following Chronic Acid Deposition

Abstract Human activities have increased acid deposition throughout the Ohio River Valley due to the large number of coal-fired power-generating facilities. The long-term effects of chronic acid deposition can include a decrease in soil pH, loss of soil fertility, and a decrease in base saturation—all of which can reduce forest productivity. Dysart Woods, a remnant old-growth, mixed mesophytic forest in eastern Ohio, has experienced a decrease in soil pH from 5.0 in 1971 to 4.6 in 1997, which may be due to chronic acid deposition. The objective of this study was to utilize a long-term study to evaluate how soil quality has changed due to chronic acid deposition. To meet the study objectives, a variety of soil chemical properties (pH, base saturation, C, N, P, etc.) were measured from surface soil within two stands of opposing aspect at Dysart Woods within the unglaciated Allegheny Plateau. Because soil pH correlates strongly with other soil chemical properties, we used pH data from 1971 to estimate how soil quality has changed over time. Mean soil pH from the south-facing stand was 5.0 in 1971, 4.6 in 1997, and 4.3 in 2007. While soil pH was not measured in the north-facing stand in 1971, pH was 4.6 in 1997 and 4.7 in 2007. Using changes in pH to estimate past soil properties, our results suggest that available base cations in the south-facing stand have been reduced from ≈12 to 6 (cmolc kg-1), with a 50% reduction in base saturation since 1971. Considering that both stands receive the same amount of acid deposition due to their close proximity to each other, results raise an interesting question: Are unglaciated Allegheny Plateau south-facing soils more susceptible to the effects of acid deposition than north-facing soils?