Living Close to Your Neighbors: The Importance of Both Competition and Facilitation in Plant Communities

Recent work has demonstrated that competition and facilitation likely operate jointly in plant communities, but teasing out the relative role of each has proven difficult. Here we address how competition and facilitation vary with seasonal fluctuations in environmental conditions, and how the effects of these fluctuations change with plant ontogeny. We planted three sizes of pine seedlings (Pinus strobus) into an herbaceous diversity experiment and measured pine growth every two weeks for two growing seasons. Both competition and facilitation occurred at different times of year between pines and their neighbors. Facilitation was important for the smallest pines when environmental conditions were severe. This effect decreased as pines got larger. Competition was stronger than facilitation overall and outweighed facilitative effects at annual time scales. Our data suggest that both competition and the counter‐directional effects of facilitation may be more common and more intense than previously considered

SIMULATING OZONE EFFECTS ON FOREST PRODUCTIVITY: INTERACTIONS AMONG LEAF‐, CANOPY‐, AND STAND‐LEVEL PROCESSES

Ozone pollution in the lower atmosphere is known to have adverse effects on forest vegetation, but the degree to which mature forests are impacted has been very difficult to assess directly. In this study, we combined leaf‐level ozone response data from independent ozone fumigation studies with a forest ecosystem model in order simulate the effects of ambient ozone on mature hardwood forests. Reductions in leaf carbon gain were determined as a linear function of ozone flux to the leaf interior, calculated as the product of ozone concentration and leaf stomatal conductance. This relationship was applied to individual canopy layers within the model in order to allow interaction with stand‐ and canopy‐level factors such as light attenuation, leaf morphology, soil water limitations, and vertical ozone gradients. The resulting model was applied to 64 locations across the northeastern United States using ambient ozone data from 1987 to 1992. Predicted declines in annual net primary production ranged from 3 to 16% with greatest reductions in southern portions of the region where ozone levels were highest, and on soils with high water‐holding capacity where drought stress was absent. Reductions in predicted wood growth were slightly greater (3–22%) because wood is a lower carbon allocation priority in the model than leaf and root growth. Interannual variation in predicted ozone effects was small due to concurrent fluctuations in ozone and climate. Periods of high ozone often coincided with hot, dry weather conditions, causing reduced stomatal conductance and ozone uptake. Within‐canopy ozone concentration gradients had little effect on predicted growth reductions because concentrations remained high through upper canopy layers where net carbon assimilation and ozone uptake were greatest. Sensitivity analyses indicate a trade‐off between model sensitivity to available soil water and foliar nitrogen and demonstrate uncertainties regarding several assumptions used in the model. Uncertainties surrounding ozone effects on stomatal function and plant water use efficiency were found to have important implications on current predictions. Field measurements of ozone effects on mature forests will be needed before the accuracy of model predictions can be fully assessed

Nitrogen cycling, forest canopy reflectance, and emergent properties of ecosystems

In Ollinger et al. (1), we reported that mass-based concentrations of nitrogen in forest canopies (%N) are positively associated with whole-canopy photosynthetic capacity and canopy shortwave albedo in temperate and boreal forests, the latter result stemming from a positive correlation between %N and canopy near infrared (NIR) reflectance. This finding is intriguing because a functional link between %N and NIR reflectance could indicate an influence of nitrogen cycling on surface energy exchange, and could provide a means for estimating %N using broad-band satellite sensors

Predicting the effects of climate change on water yield and forest production in the northeastern United States

Rapid and simultaneous changes in temperature, precipitation and the atmospheric concentration of CO2 are predicted to occur over the next century. Simple, well-validated models of ecosystem function are required to predict the effects of these changes. This paper describes an improved version of a forest carbon and water balance model (PnET-II) and the application of the model to predict stand- and regional-level effects of changes in temperature, precipitation and atmospheric CO2 concentration. PnET-II is a simple, generalized, monthly time-step model of water and carbon balances (gross and net) driven by nitrogen availability as expressed through foliar N concentration. Improvements from the original model include a complete carbon balance and improvements in the prediction of canopy phenology, as well as in the computation of canopy structure and photosynthesis. The model was parameterized and run for 4 forest/site combinations and validated against available data for water yield, gross and net carbon exchange and biomass production. The validation exercise suggests that the determination of actual water availability to stands and the occurrence or non-occurrence of soil-based water stress are critical to accurate modeling of forest net primary production (NPP) and net ecosystem production (NEP). The model was then run for the entire NewEngland/New York (USA) region using a 1 km resolution geographic information system. Predicted long-term NEP ranged from -85 to +275 g C m-2 yr-1 for the 4 forest/site combinations, and from -150 to 350 g C m-2 yr-1 for the region, with a regional average of 76 g C m-2 yr-1. A combination of increased temperature (+6*C), decreased precipitation (-15%) and increased water use efficiency (2x, due to doubling of CO2) resulted generally in increases in NPP and decreases in water yield over the region

Divergent drivers of leaf trait variation within species, among species, and among functional groups.

Understanding variation in leaf functional traits-including rates of photosynthesis and respiration and concentrations of nitrogen and phosphorus-is a fundamental challenge in plant ecophysiology. When expressed per unit leaf area, these traits typically increase with leaf mass per area (LMA) within species but are roughly independent of LMA across the global flora. LMA is determined by mass components with different biological functions, including photosynthetic mass that largely determines metabolic rates and contains most nitrogen and phosphorus, and structural mass that affects toughness and leaf lifespan (LL). A possible explanation for the contrasting trait relationships is that most LMA variation within species is associated with variation in photosynthetic mass, whereas most LMA variation across the global flora is associated with variation in structural mass. This hypothesis leads to the predictions that (i) gas exchange rates and nutrient concentrations per unit leaf area should increase strongly with LMA across species assemblages with low LL variance but should increase weakly with LMA across species assemblages with high LL variance and that (ii) controlling for LL variation should increase the strength of the above LMA relationships. We present analyses of intra- and interspecific trait variation from three tropical forest sites and interspecific analyses within functional groups in a global dataset that are consistent with the above predictions. Our analysis suggests that the qualitatively different trait relationships exhibited by different leaf assemblages can be understood by considering the degree to which photosynthetic and structural mass components contribute to LMA variation in a given assemblage

Improved representation of plant functional types and physiology in the Joint UK Land Environment Simulator (JULES v4.2) using plant trait information

Dynamic global vegetation models are used to predict the response of vegetation to climate change. They are essential for planning ecosystem management, understanding carbon cycle–climate feedbacks, and evaluating the potential impacts of climate change on global ecosystems. JULES (the Joint UK Land Environment Simulator) represents terrestrial processes in the UK Hadley Centre family of models and in the first generation UK Earth System Model. Previously, JULES represented five plant functional types (PFTs): broadleaf trees, needle-leaf trees, C3 and C4 grasses, and shrubs. This study addresses three developments in JULES. First, trees and shrubs were split into deciduous and evergreen PFTs to better represent the range of leaf life spans and metabolic capacities that exists in nature. Second, we distinguished between temperate and tropical broadleaf evergreen trees. These first two changes result in a new set of nine PFTs: tropical and temperate broadleaf evergreen trees, broadleaf deciduous trees, needle-leaf evergreen and deciduous trees, C3 and C4 grasses, and evergreen and deciduous shrubs. Third, using data from the TRY database, we updated the relationship between leaf nitrogen and the maximum rate of carboxylation of Rubisco (Vcmax), and updated the leaf turnover and growth rates to include a trade-off between leaf life span and leaf mass per unit area. Overall, the simulation of gross and net primary productivity (GPP and NPP, respectively) is improved with the nine PFTs when compared to FLUXNET sites, a global GPP data set based on FLUXNET, and MODIS NPP. Compared to the standard five PFTs, the new nine PFTs simulate a higher GPP and NPP, with the exception of C3 grasses in cold environments and C4 grasses that were previously over-productive. On a biome scale, GPP is improved for all eight biomes evaluated and NPP is improved for most biomes – the exceptions being the tropical forests, savannahs, and extratropical mixed forests where simulated NPP is too high. With the new PFTs, the global present-day GPP and NPP are 128 and 62 Pg C year−1, respectively. We conclude that the inclusion of trait-based data and the evergreen/deciduous distinction has substantially improved productivity fluxes in JULES, in particular the representation of GPP. These developments increase the realism of JULES, enabling higher confidence in simulations of vegetation dynamics and carbon storage

Canopy nitrogen, carbon assimilation, and albedo in temperate and boreal forests: Functional relations and potential climate feedbacks

The availability of nitrogen represents a key constraint on carbon cycling in terrestrial ecosystems, and it is largely in this capacity that the role of N in the Earth\u27s climate system has been considered. Despite this, few studies have included continuous variation in plant N status as a driver of broad-scale carbon cycle analyses. This is partly because of uncertainties in how leaf-level physiological relationships scale to whole ecosystems and because methods for regional to continental detection of plant N concentrations have yet to be developed. Here, we show that ecosystem CO2 uptake capacity in temperate and boreal forests scales directly with whole-canopy N concentrations, mirroring a leaf-level trend that has been observed for woody plants worldwide. We further show that both CO2 uptake capacity and canopy N concentration are strongly and positively correlated with shortwave surface albedo. These results suggest that N plays an additional, and overlooked, role in the climate system via its influence on vegetation reflectivity and shortwave surface energy exchange. We also demonstrate that much of the spatial variation in canopy N can be detected by using broad-band satellite sensors, offering a means through which these findings can be applied toward improved application of coupled carbon cycle–climate models

Computing Nash Equilibrium in Wireless Ad Hoc Networks: A Simulation-Based Approach

This paper studies the problem of computing Nash equilibrium in wireless networks modeled by Weighted Timed Automata. Such formalism comes together with a logic that can be used to describe complex features such as timed energy constraints. Our contribution is a method for solving this problem using Statistical Model Checking. The method has been implemented in UPPAAL model checker and has been applied to the analysis of Aloha CSMA/CD and IEEE 802.15.4 CSMA/CA protocols.Comment: In Proceedings IWIGP 2012, arXiv:1202.422