Disturbance-accelerated succession increases the production of a temperate forest.

Many secondary deciduous forests of eastern North America are approaching a transition in which mature early-successional trees are declining, resulting in an uncertain future for this century-long carbon (C) sink. We initiated the Forest Accelerated Succession Experiment (FASET) at the University of Michigan Biological Station to examine the patterns and mechanisms underlying forest C cycling following the stem girdling-induced mortality of >6,700 early-successional Populus spp. (aspen) and Betula papyrifera (paper birch). Meteorological flux tower-based C cycling observations from the 33-ha treatment forest have been paired with those from a nearby unmanipulated forest since 2008. Following over a decade of observations, we revisit our core hypothesis: that net ecosystem production (NEP) would increase following the transition to mid-late-successional species dominance due to increased canopy structural complexity. Supporting our hypothesis, NEP was stable, briefly declined, and then increased relative to the control in the decade following disturbance; however, increasing NEP was not associated with rising structural complexity but rather with a rapid 1-yr recovery of total leaf area index as mid-late-successional Acer, Quercus, and Pinus assumed canopy dominance. The transition to mid-late-successional species dominance improved carbon-use efficiency (CUE = NEP/gross primary production) as ecosystem respiration declined. Similar soil respiration rates in control and treatment forests, along with species differences in leaf physiology and the rising relative growth rates of mid-late-successional species in the treatment forest, suggest changes in aboveground plant respiration and growth were primarily responsible for increases in NEP. We conclude that deciduous forests transitioning from early to middle succession are capable of sustained or increased NEP, even when experiencing extensive tree mortality. This adds to mounting evidence that aging deciduous forests in the region will function as C sinks for decades to come.

COSORE: A community database for continuous soil respiration and other soil-atmosphere greenhouse gas flux data.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil-to-atmosphere CO2 flux, commonly though imprecisely termed soil respiration (RS ), is one of the largest carbon fluxes in the Earth system. An increasing number of high-frequency RS measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open-source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long-term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured RS , the database design accommodates other soil-atmosphere measurements (e.g. ecosystem respiration, chamber-measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package.

3,4-Methylenedioxypyrovalerone: Neuropharmacological Impact of a Designer Stimulant of Abuse on Monoamine Transporters.

Methylenedioxypyrovalerone (MDPV) is an abused synthetic cathinone, commonly referred to as a "bath salt." Because the dopamine (DA) transporter (DAT) and vesicular monoamine transporter-2 (VMAT-2) are key regulators of both the abuse and neurotoxic potential of structurally and behaviorally related agents, the impact of MDPV on these transporters was investigated. Results revealed that a single in vivo MDPV administration rapidly (within 1 hour) and reversibly increased both rat striatal DAT and VMAT-2 activity, as assessed via [3H]DA uptake in synaptosomes and synaptic vesicles, respectively, prepared from treated rats. There was no evidence of an MDPV-induced increase in plasmalemmal membrane DAT surface expression. Plasma concentrations of MDPV increased dose-dependently as assessed 1 hour after 2.5 and 5.0 mg/kg (s.c.) administration and returned to levels less than 10 ng/ml by 18 hours after 2.5 mg/kg (s.c.). Neither pretreatment with a D1 receptor (SCH23390), a D2 receptor (eticlopride), nor a nicotinic receptor (mecamylamine) antagonist attenuated the MDPV-induced increase in DAT activity. In contrast, eticlopride pretreatment attenuated both the MDPV-induced increase in VMAT-2-mediated DA uptake and an associated increase in cytoplasmic-associated vesicle VMAT-2 immunoreactivity. SCH23390 did not attenuate the MDPV-induced increase in VMAT-2 activity. Repeated MDPV injections did not cause persistent DAergic deficits, as assessed 7 to 8 days later. The impact of MDPV on striatal and hippocampal serotonergic assessments was minimal. Taken together, these data contribute to a growing pharmacological rubric for evaluating the ever-growing list of designer cathinone-related stimulants. The profile of MDPV compared with related psychostimulants is discussed. SIGNIFICANCE STATEMENT: Pharmacological characterization of the synthetic cathinone, 3,4-methylenedioxypyrovalerone (MDPV; commonly referred to as a "bath salt"), is critical for understanding the abuse liability and neurotoxic potential of this and related agents. Accordingly, the impact of MDPV on monoaminergic neurons is described and compared with that of related psychostimulants.

Defining a spectrum of integrative trait-based vegetation canopy structural types.

Vegetation canopy structure is a fundamental characteristic of terrestrial ecosystems that defines vegetation types and drives ecosystem functioning. We use the multivariate structural trait composition of vegetation canopies to classify ecosystems within a global canopy structure spectrum. Across the temperate forest sub-set of this spectrum, we assess gradients in canopy structural traits, characterise canopy structural types (CST) and evaluate drivers and functional consequences of canopy structural variation. We derive CSTs from multivariate canopy structure data, illustrating variation along three primary structural axes and resolution into six largely distinct and functionally relevant CSTs. Our results illustrate that within-ecosystem successional processes and disturbance legacies can produce variation in canopy structure similar to that associated with sub-continental variation in forest types and eco-climatic zones. The potential to classify ecosystems into CSTs based on suites of structural traits represents an important advance in understanding and modelling structure-function relationships in vegetated ecosystems.

Forest aging, disturbance and the carbon cycle.

Contents Summary 1188 I. Introduction 1188 II. Forest aging and carbon storage 1189 III. Successional trends of NEP in northern deciduous forests 1190 IV. Mechanisms sustaining NEP in aging deciduous forests 1191 Acknowledgements 1192 References 1192 SUMMARY: Large areas of forestland in temperate North America, as well as in other parts of the world, are growing older and will soon transition into middle and then late successional stages exceeding 100 yr in age. These ecosystems have been important regional carbon sinks as they recovered from prior anthropogenic and natural disturbance, but their future sink strength, or annual rate of carbon storage, is in question. Ecosystem development theory predicts a steady decline in annual carbon storage as forests age, but newly available, direct measurements of forest net CO2 exchange challenge that prediction. In temperate deciduous forests, where moderate severity disturbance regimes now often prevail, there is little evidence for any marked decline in carbon storage rate during mid-succession. Rather, an increase in physical and biological complexity under these disturbance regimes may drive increases in resource-use efficiency and resource availability that help to maintain significant carbon storage in these forests well past the century mark. Conservation of aging deciduous forests may therefore sustain the terrestrial carbon sink, whilst providing other goods and services afforded by these biologically and structurally complex ecosystems.

Effects of structural complexity on within-canopy light environments and leaf traits in a northern mixed deciduous forest.

Canopy structure influences forest productivity through its effects on the distribution of radiation and the light-induced changes in leaf physiological traits. Due to the difficulty of accessing and measuring forest canopies, few field-based studies have quantitatively linked these divergent scales of canopy functioning. The objective of our study was to investigate how canopy structure affects light profiles within a forest canopy and whether leaves of mature trees adjust morphologically and biochemically to the light environments characteristic of canopies with different structural complexity. We used a combination of light detection and ranging (LiDAR) data and hemispherical photographs to quantify canopy structure and light environments, respectively, and a telescoping pole to sample leaves. Leaf mass per area (LMA), nitrogen on an area basis (Narea) and chlorophyll on a mass basis (Chlmass) were measured in red maple (Acer rubrum), american beech (Fagus grandifolia), white pine (Pinus strobus), and northern red oak (Quercus rubra) at different heights in plots with similar leaf area index but contrasting canopy complexity (rugosity). We found that more complex canopies had greater porosity and reduced light variability in the midcanopy while total light interception was unchanged relative to less complex canopies. Leaf phenotypes of F. grandifolia, Q. rubra and P. strobus were more sun-acclimated in the midstory of structurally complex canopies while leaf phenotypes of A. rubrum were more shade-acclimated (lower LMA) in the upper canopy of more complex stands, despite no differences in total light interception. Broadleaf species showed further differences in acclimation with increased Narea and reduced Chlmass in leaves with higher LMA, while P. strobus showed no change in Narea and Chlmass with higher LMA. Our results provide new insight on how light distribution and leaf acclimation in mature trees might be altered when natural and anthropogenic disturbances cause structural changes in the canopy.

Net primary production of a temperate deciduous forest exhibits a threshold response to increasing disturbance severity.

The global carbon (C) balance is vulnerable to disturbances that alter terrestrial C storage. Disturbances to forests occur along a continuum of severity, from low-intensity disturbance causing the mortality or defoliation of only a subset of trees to severe stand- replacing disturbance that kills all trees; yet considerable uncertainty remains in how forest production changes across gradients of disturbance intensity. We used a gradient of tree mortality in an upper Great Lakes forest ecosystem to: (1) quantify how aboveground wood net primary production (ANPP,) responds to a range of disturbance severities; and (2) identify mechanisms supporting ANPPw resistance or resilience following moderate disturbance. We found that ANPPw declined nonlinearly with rising disturbance severity, remaining stable until >60% of the total tree basal area senesced. As upper canopy openness increased from disturbance, greater light availability to the subcanopy enhanced the leaf-level photosynthesis and growth of this formerly light-limited canopy stratum, compensating for upper canopy production losses and a reduction in total leaf area index (LAI). As a result, whole-ecosystem production efficiency (ANPPw/LAI) increased with rising disturbance severity, except in plots beyond the disturbance threshold. These findings provide a mechanistic explanation for a nonlinear relationship between ANPPw, and disturbance severity, in which the physiological and growth enhancement of undisturbed vegetation is proportional to the level of disturbance until a threshold is exceeded. Our results have important ecological and management implications, demonstrating that in some ecosystems moderate levels of disturbance minimally alter forest production.

Joint control of terrestrial gross primary productivity by plant phenology and physiology.

Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate-carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy-covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000-2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r(2) = 0.90) and GPP recovery after a fire disturbance in South Dakota (r(2) = 0.88). Additional analysis of the eddy-covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

Sustained carbon uptake and storage following moderate disturbance in a Great Lakes forest.

Carbon (C) uptake rates in many forests are sustained, or decline only briefly, following disturbances that partially defoliate the canopy. The mechanisms supporting such functional resistance to moderate forest disturbance are largely unknown. We used a large-scale experiment, in which > 6700 Populus (aspen) and Betula (birch) trees were stem-girdled within a 39-ha area, to identify mechanisms sustaining C uptake through partial canopy defoliation. The Forest Accelerated Succession Experiment in northern Michigan, USA, employs a suite of C-cycling measurements within paired treatment and control meteorological flux tower footprints. We found that enhancement of canopy light-use efficiency and maintenance of light absorption maintained net ecosystem production (NEP) and aboveground wood net primary production (NPP) when leaf-area index (LAI) of the treatment forest temporarily declined by nearly half its maximum value. In the year following peak defoliation, redistribution of nitrogen (N) in the treatment forest from senescent early successional aspen and birch to non-girdled later successional species facilitated the recovery of total LAI to pre-disturbance levels. Sustained canopy physiological competency following disturbance coincided with a downward shift in maximum canopy height, indicating that compensatory photosynthetic C uptake by undisturbed, later successional subdominant and subcanopy vegetation supported C-uptake resistance to disturbance. These findings have implications for ecosystem management and modeling, demonstrating that forests may tolerate considerable leaf-area losses without diminishing rates of C uptake. We conclude that the resistance of C uptake to moderate disturbance depends not only on replacement of lost leaf area, but also on rapid compensatory photosynthetic C uptake during defoliation by emerging later successional species.

Herbarium specimens reveal the footprint of climate change on flowering trends across north-central North America.

Shifting flowering phenology with rising temperatures is occurring worldwide, but the rarity of co-occurring long-term observational and temperature records has hindered the evaluation of phenological responsiveness in many species and across large spatial scales. We used herbarium specimens combined with historic temperature data to examine the impact of climate change on flowering trends in 141 species collected across 116,000 km(2) in north-central North America. On average, date of maximum flowering advanced 2.4 days °C(-1), although species-specific responses varied from - 13.5 to + 7.3 days °C(-1). Plant functional types exhibited distinct patterns of phenological responsiveness with significant differences between native and introduced species, among flowering seasons, and between wind- and biotically pollinated species. This study is the first to assess large-scale patterns of phenological responsiveness with broad species representation and is an important step towards understanding current and future impacts of climate change on species performance and biodiversity.

Optimizing wind power generation while minimizing wildlife impacts in an urban area.

The location of a wind turbine is critical to its power output, which is strongly affected by the local wind field. Turbine operators typically seek locations with the best wind at the lowest level above ground since turbine height affects installation costs. In many urban applications, such as small-scale turbines owned by local communities or organizations, turbine placement is challenging because of limited available space and because the turbine often must be added without removing existing infrastructure, including buildings and trees. The need to minimize turbine hazard to wildlife compounds the challenge. We used an exclusion zone approach for turbine-placement optimization that incorporates spatially detailed maps of wind distribution and wildlife densities with power output predictions for the Ohio State University campus. We processed public GIS records and airborne lidar point-cloud data to develop a 3D map of all campus buildings and trees. High resolution large-eddy simulations and long-term wind climatology were combined to provide land-surface-affected 3D wind fields and the corresponding wind-power generation potential. This power prediction map was then combined with bird survey data. Our assessment predicts that exclusion of areas where bird numbers are highest will have modest effects on the availability of locations for power generation. The exclusion zone approach allows the incorporation of wildlife hazard in wind turbine siting and power output considerations in complex urban environments even when the quantitative interaction between wildlife behavior and turbine activity is unknown.

Fire effects on temperate forest soil C and N storage.

Temperate forest soils store globally significant amounts of carbon (C) and nitrogen (N). Understanding how soil pools of these two elements change in response to disturbance and management is critical to maintaining ecosystem services such as forest productivity, greenhouse gas mitigation, and water resource protection. Fire is one of the principal disturbances acting on forest soil C and N storage and is also the subject of enormous management efforts. In the present article, we use meta-analysis to quantify fire effects on temperate forest soil C and N storage. Across a combined total of 468 soil C and N response ratios from 57 publications (concentrations and pool sizes), fire had significant overall effects on soil C (-26%) and soil N (-22%). The impacts of fire on forest floors were significantly different from its effects on mineral soils. Fires reduced forest floor C and N storage (pool sizes only) by an average of 59% and 50%, respectively, but the concentrations of these two elements did not change. Prescribed fires caused smaller reductions in forest floor C and N storage (-46% and -35%) than wildfires (-67% and -69%), and the presence of hardwoods also mitigated fire impacts. Burned forest floors recovered their C and N pools in an average of 128 and 103 years, respectively. Among mineral soils, there were no significant changes in C or N storage, but C and N concentrations declined significantly (-11% and -12%, respectively). Mineral soil C and N concentrations were significantly affected by fire type, with no change following prescribed burns, but significant reductions in response to wildfires. Geographic variation in fire effects on mineral soil C and N storage underscores the need for region-specific fire management plans, and the role of fire type in mediating C and N shifts (especially in the forest floor) indicates that averting wildfires through prescribed burning is desirable from a soils perspective.

15N discrimination and the sensitivity of nitrogen fixation to changes in dietary nitrogen in Reticulitermes flavipes (Isoptera: Rhinotermitidae).

Xylophagous termites possess symbiotic bacteria that fix atmospheric nitrogen (N(2)). Although symbiotic N(2) fixation is central to termite nutrition and ecologically important, it is energetically costly. Using stable isotopes, we tested the hypothesis that symbiotic N(2) fixation would decrease in workers of the eastern subterranean termite, Reticulitermes flavipes Kollar, which were exposed to high nitrogen diets. To calculate the isotope discrimination factor occurring as a result of digestion, Δ(dig), and which occurs as the result of N(2) fixation, Δ(fix), symbiotic N(2) fixation was inhibited via force feeding termites the antibiotic kanamycin. Antibiotic-treated termites and control (N(2)-fixing) termites were exposed to different concentrations of dietary N (0, 0.21, and 0.94% N), their (15)N signatures were obtained, and the percent nitrogen derived from the atmosphere within termite samples was calculated. As we hypothesized, symbiotic N(2) fixation rates were negatively correlated with dietary N, suggesting that high concentrations of dietary N suppressed symbiotic N(2) fixation in R. flavipes. A comparison of the (15)N isotope signatures of antibiotic-treated termites with their food sources demonstrated that Δ(dig) = 2.284‰, and a comparison of the (15)N isotope signatures of antibiotic-treated termites with control termites indicated that Δ(fix) = -1.238‰. These are the first estimates of Δ(dig) for R. flavipes, and the first estimate of Δ(fix) for any N(2)-fixing termite species.

Ecosystem CO2 fluxes of arbuscular and ectomycorrhizal dominated vegetation types are differentially influenced by precipitation and temperature.

Here, we explore how interannual variations in environmental factors (i.e. temperature, precipitation and light) influence CO(2) fluxes (gross primary production and ecosystem respiration) in terrestrial ecosystems classified by vegetation type and the mycorrhizal type of dominant plants (arbuscular mycorrhizal (AM) or ectomycorrhizal (EM)). We combined 236 site-year measurements of terrestrial ecosystem CO(2) fluxes and environmental factors from 50 eddy-covariance flux tower sites with information about climate, vegetation type and dominant plant species. Across large geographical distances, interannual variations in ecosystem CO(2) fluxes for EM-dominated sites were primarily controlled by interannual variations in mean annual temperature. By contrast, interannual variations in ecosystem CO(2) fluxes at AM-dominated sites were primarily controlled by interannual variations in precipitation. This study represents the first large-scale assessment of terrestrial CO(2) fluxes in multiple vegetation types classified according to dominant mycorrhizal association. Our results support and complement the hypothesis that bioclimatic conditions influence the distribution of AM and EM systems across large geographical distances, which leads to important differences in the major climatic factors controlling ecosystem CO(2) fluxes.

Neither mycorrhizal inoculation nor atmospheric CO2 concentration has strong effects on pea root production and root loss.

• Root responses to elevated CO2 concentrations, where nutrient demand was expected to be higher than at ambient CO2 , and possible interactions with mycorrhizal symbionts are reported for pea (Pisum sativum). These are important below-ground components affecting carbon flow into the soil. • A video-minirhizotron system was used to study root growth in pot-grown mycorrhizal (inoculated with Glomus caledonium) and nonmycorrhizal pea plants at ambient or elevated CO2 concentrations over 9 wk. Analyses were made of root length changes, cohort size and survivorship. • Root length production at ambient, but not at elevated CO2 , was higher in nonmycorrhizal than in mycorrhizal plants from week 4-7. Root loss began at week 5, peaking 2 wk later with 40-50% loss of the root length produced by week 8. The decline in root production and increase in root loss coincided with the onset of flowering. • Neither mycorrhizal inoculation nor CO2 concentration has a strong effect on pea root production and root loss, although mycorrhizal infection has a greater effect than CO2 .

Kinetics of nitrogen uptake by Populus tremuloides in relation to atmospheric CO(2) and soil nitrogen availability.

Sustained increases in plant production in response to elevated atmospheric carbon dioxide (CO(2)) concentration may be constrained by the availability of soil nitrogen (N). However, it is possible that plants will respond to N limitation at elevated CO(2) concentration by increasing the specific N uptake capacity of their roots. To explore this possibility, we examined the kinetics of (15)NH(4) (+) and (15)NO(3) (-) uptake by excised roots of Populus tremuloides Michx. grown in ambient and twice-ambient CO(2) concentrations, and in soils of low- and high-N availability. Elevated CO(2) concentration had no effect on either NH(4) (+) or NO(3) (-) uptake, whereas high-N availability decreased the capacity of roots to take up both NH(4) (+) and NO(3) (-). The maximal rate of NH(4) (+) uptake decreased from 12 to 8 &mgr;mol g(-1) h(-1), and K(m) increased from 49 to 162 &mgr;mol l(-1), from low to high soil N availability.Because NO(3) (-) uptake exhibited mixedkinetics over the concentration range we used (10-500 &mgr;mol l( -1)), it was not possible to calculate V(max) and K(m). Instead, we used an uptake rate of 100 &mgr;mol g(-1) h(-1) as our metric of NO(3) (-) uptake capacity, which averaged 0.45 and 0.23 &mgr;mol g(-1) h(-1) at low- and high-N availability, respectively. The proximal mechanisms for decreased N uptake capacity at high-N availability appeared to be an increase in fine-root carbohydrate status and a decrease in fine-root N concentration. Both NH(4) (+) and NO(3) (-) uptake were inversely related to fine-root N concentration, and positively related to fine-root total nonstructural carbohydrate concentration. We conclude that soil N availability, through its effects on fine-root N and carbohydrate status, has a much greater influence on the specific uptake capacity of P. tremuloides fine roots than elevated atmospheric CO(2). In elevated atmospheric CO(2), changes in N acquisition by P. tremuloides appeared to be driven by changes in root architecture and biomass, rather than by changes in the amount or activity of N-uptake enzymes.

A meta-analysis of elevated CO2 effects on woody plant mass, form, and physiology.

Quantitative integration of the literature on the effect of elevated CO2 on woody plants is important to aid our understanding of forest health in coming decades and to better predict terrestrial feedbacks on the global carbon cycle. We used meta-analytic methods to summarize and interpret more than 500 reports of effects of elevated CO2 on woody plant biomass accumulation and partitioning, gas exchange, and leaf nitrogen and starch content. The CO2 effect size metric we used was the log-transformed ratio of elevated compared to ambient response means weighted by the inverse of the variance of the log ratio. Variation in effect size among studies was partitioned according to the presence of interacting stress factors, length of CO2 exposure, functional group status, pot size, and type of CO2 exposure facility. Both total biomass (W T) and net CO2 assimilation (A) increased significantly at about twice ambient CO2, regardless of growth conditions. Low soil nutrient availability reduced the CO2 stimulation of W T by half, from +31% under optimal conditions to +16%, while low light increased the response to +52%. We found no significant shifts in biomass allocation under high CO2. Interacting stress factors had no effect on the magnitude of responses of A to CO2, although plants grown in growth chambers had significantly lower responses (+19%) than those grown in greenhouses or in open-top chambers (+54%). We found no consistent evidence for photosynthetic acclimation to CO2 enrichment except in trees grown in pots <0.5 l (-36%) and no significant CO2 effect on stomatal conductance. Both leaf dark respiration and leaf nitrogen were significantly reduced under elevated CO2 (-18% and -16% respectively, data expressed on a leaf mass basis), while leaf starch content increased significantly except in low nutrient grown gymnosperms. Our results provide robust, statistically defensible estimates of elevated CO2 effect sizes against which new results may be compared or for use in forest and climate model parameterization.