Plant traits mediate foliar uptake of deposited nitrogen by mature woody plants.

Increased atmospheric nitrogen (N) deposition significantly disturbs ecosystem N cycle. Although foliar interception and uptake of N deposition can provide an important alternative N supply to forest ecosystems, the mechanisms regulating foliar N uptake from wet deposition are not fully understood. Here, we selected 19 woody species with a wide range of plant traits from different functional groups and conducted a 15N isotope labelling experiment through brushing 15NH4 + and 15NO3 - solution on canopy leaves. Our findings demonstrate that leaves can directly absorb N from wet deposition within a few hours. The average leaf 15N recoveries were 10% and 28% under 15NH4 + and 15NO3 - treatments across species, respectively, while twig N recoveries were only 1%-7% of leaf N recoveries. Differences in foliar N uptake efficiency among species were closely associated with leaf traits but were little influenced by meteorological conditions or soil nutrient status. Specifically, plants with higher leaf N concentration, larger specific leaf area and lower wax concentration exhibited higher leaf N recovery. Our results indicated that tree canopies could directly absorb N from atmospheric deposition. We highlight the critical role of leaf traits in determining canopy foliar N uptake, which may consequently influence plant competition under elevated N deposition.

Urbanization and fragmentation have opposing effects on soil nitrogen availability in temperate forest ecosystems.

Nitrogen (N) availability relative to plant demand has been declining in recent years in terrestrial ecosystems throughout the world, a phenomenon known as N oligotrophication. The temperate forests of the northeastern U.S. have experienced a particularly steep decline in bioavailable N, which is expected to be exacerbated by climate change. This region has also experienced rapid urban expansion in recent decades that leads to forest fragmentation, and it is unknown whether and how these changes affect N availability and uptake by forest trees. Many studies have examined the impact of either urbanization or forest fragmentation on nitrogen (N) cycling, but none to our knowledge have focused on the combined effects of these co-occurring environmental changes. We examined the effects of urbanization and fragmentation on oak-dominated (Quercus spp.) forests along an urban to rural gradient from Boston to central Massachusetts (MA). At eight study sites along the urbanization gradient, plant and soil measurements were made along a 90 m transect from a developed edge to an intact forest interior. Rates of net ammonification, net mineralization, and foliar N concentrations were significantly higher in urban than rural sites, while net nitrification and foliar C:N were not different between urban and rural forests. At urban sites, foliar N and net ammonification and mineralization were higher at forest interiors compared to edges, while net nitrification and foliar C:N were higher at rural forest edges than interiors. These results indicate that urban forests in the northeastern U.S. have greater soil N availability and N uptake by trees compared to rural forests, counteracting the trend for widespread N oligotrophication in temperate forests around the globe. Such increases in available N are diminished at forest edges, however, demonstrating that forest fragmentation has the opposite effect of urbanization on coupled N availability and demand by trees.

The energy-water limitation threshold explains divergent drought responses in tree growth, needle length, and stable isotope ratios.

Predicted increases in extreme droughts will likely cause major shifts in carbon sequestration and forest composition. Although growth declines during drought are widely documented, an increasing number of studies have reported both positive and negative responses to the same drought. These divergent growth patterns may reflect thresholds (i.e., nonlinear responses) promoted by changes in the dominant climatic constraints on tree growth. Here we tested whether stemwood growth exhibited linear or nonlinear responses to temperature and precipitation and whether stemwood growth thresholds co-occurred with multiple thresholds in source and sink processes that limit tree growth. We extracted 772 tree cores, 1398 needle length records, and 1075 stable isotope samples from 27 sites across whitebark pine's (Pinus albicaulis Engelm.) climatic niche in the Sierra Nevada. Our results indicated that a temperature threshold in stemwood growth occurred at 8.4°C (7.12-9.51°C; estimated using fall-spring maximum temperature). This threshold was significantly correlated with thresholds in foliar growth, as well as carbon (δ13 C) and nitrogen (δ15 N) stable isotope ratios, that emerged during drought. These co-occurring thresholds reflected the transition between energy- and water-limited tree growth (i.e., the E-W limitation threshold). This transition likely mediated carbon and nutrient cycling, as well as important differences in growth-defense trade-offs and drought adaptations. Furthermore, whitebark pine growing in energy-limited regions may continue to experience elevated growth in response to climate change. The positive effect of warming, however, may be offset by growth declines in water-limited regions, threatening the long-term sustainability of the recently listed whitebark pine species in the Sierra Nevada.

Drivers of foliar 15 N trends in southern China over the last century.

Foliar stable nitrogen (N) isotopes (δ15 N) generally reflect N availability to plants and have been used to infer about changes thereof. However, previous studies of temporal trends in foliar δ15 N have ignored the influence of confounding factors, leading to uncertainties on its indication to N availability. In this study, we measured foliar δ15 N of 1811 herbarium specimens from 12 plant species collected in southern China forests from 1920 to 2010. We explored how changes in atmospheric CO2 , N deposition and global warming have affected foliar δ15 N and N concentrations ([N]) and identified whether N availability decreased in southern China. Across all species, foliar δ15 N significantly decreased by 0.82‰ over the study period. However, foliar [N] did not decrease significantly, implying N homeostasis in forest trees in the region. The spatiotemporal patterns of foliar δ15 N were explained by mean annual temperature (MAT), atmospheric CO2 ( P CO 2 ), atmospheric N deposition, and foliar [N]. The spatiotemporal trends of foliar [N] were explained by MAT, temperature seasonality, P CO 2 , and N deposition. N deposition within the rates from 5.3 to 12.6 kg N ha-1  year-1 substantially contributed to the temporal decline in foliar δ15 N. The decline in foliar δ15 N was not accompanied by changes in foliar [N] and therefore does not necessarily reflect a decline in N availability. This is important to understand changes in N availability, which is essential to validate and parameterize biogeochemical cycles of N.

Enhancement of anthocyanin and chromatic profiles in 'Cabernet Sauvignon' (Vitis vinifera L.) by foliar nitrogen fertilizer during veraison.

BACKGROUND: The influence of foliar nitrogen fertilizer during veraison (FNFV) on anthocyanin accumulation and chromatic characteristics of 'Cabernet Sauvignon' grapes over two seasons was investigated. RESULTS: Urea and phenylalanine fertilizers (TU and TP, respectively) and a control were sprayed three times at veraison. In 2018, TU displayed a significant enhancement in total individual anthocyanin content and a* and Cab * profiles. In 2019, FNAV significantly improved the content of total non-acylated, acylated anthocyanin and total individual anthocyanin, and the profiles of L*, a* and Cab *, except a* in TU. The whole process from phenylalanine variation to anthocyanin accumulation in grape skins was analyzed. On the whole, after the first FNFV to harvest, the increase in phenylalanine metabolism, abscisic acid content, effects of PAL (Phenylalanine ammonia lyase), UFGT (UDP glucose-flavonoid 3-O-glucosyltransferase) and transcript concentrations of VvPAL and VvUFGT involved in anthocyanin biosynthesis were also strong evidence explaining the increased anthocyanin and chromatic profiles in 2019. CONCLUSION: Overall, FNFV for nitrogen-deficient grapevines could significantly improve grape color, especially in the 2019 veraison with a proper climate. © 2021 Society of Chemical Industry.

Precipitation is the main factor affecting the variation of foliar nitrogen isotope composition in two leguminous shrub species of northwestern China.

An increase in foliar nitrogen isotope composition (δ15N) with decreasing precipitation has been shown to occur widely in non-N2-fixing plant species. However, similar patterns have not been identified in N2-fixing species. Here, we tested the relationships of foliar δ15N with local environmental factors and leaf properties in two leguminous shrub species (Caragana korshinskii and Caragana liouana) sampled from 30 populations. Results indicated that the mean annual precipitation (MAP) primarily accounted for the variation of foliar δ15N in the two species. Further analysis revealed strong negative correlations between foliar δ15N and MAP within and across species. These results suggest that the foliar δ15N of leguminous shrub species also shift along precipitation gradients, which augments our understanding of the relationships between foliar δ15N and climatic factors.

Landscape variation in canopy nitrogen and carbon assimilation in a temperate mixed forest.

Canopy nitrogen (N) is a key factor regulating carbon cycling in forest ecosystems through linkages among foliar N and photosynthesis, decomposition, and N cycling. This analysis examined landscape variation in canopy nitrogen and carbon assimilation in a temperate mixed forest surrounding Harvard Forest in central Massachusetts, USA by integration of canopy nitrogen mapping with ecosystem modeling, and spatial data from soils, stand characteristics and disturbance history. Canopy %N was mapped using high spectral resolution remote sensing from NASA's AVIRIS (Airborne Visible/Infrared Imaging Spectrometer) instrument and linked to an ecosystem model, PnET-II, to estimate gross primary productivity (GPP). Predicted GPP was validated with estimates derived from eddy covariance towers. Estimated canopy %N ranged from 0.5 to 2.9% with a mean of 1.75% across the study region. Predicted GPP ranged from 797 to 1622 g C m-2 year-1 with a mean of 1324 g C m-2 year-1. The prediction that spatial patterns in forest growth are associated with spatial patterns in estimated canopy %N was supported by a strong, positive relationship between field-measured canopy %N and aboveground net primary production. Estimated canopy %N and GPP were related to forest composition, land-use history, and soil drainage. At the landscape scale, PnET-II GPP was compared with predicted GPP from the BigFoot project and from NASA's MODIS (Moderate Resolution Imaging Spectroradiometer) data products. Estimated canopy %N explained much of the difference between MODIS GPP and PnET-II GPP, suggesting that global MODIS GPP estimates may be improved if broad-scale estimates of foliar N were available.

Variations in yield and gluten proteins in durum wheat varieties under late-season foliar versus soil application of nitrogen fertilizer in a northern Mediterranean environment.

BACKGROUND: With the increasing demand for high-quality foodstuffs and concern for environmental sustainability, late-season nitrogen (N) foliar fertilization of common wheat is now an important and widespread practice. This study investigated the effects of late-season foliar versus soil N fertilization on yield and protein content of four varieties of durum wheat, Aureo, Ariosto, Biensur and Liberdur, in a three-year field trial in northern Italy. RESULTS: Variations in low-molecular-weight glutenins (LMW-GS), high-molecular-weight glutenins (HMW-GS) and gliadins were assessed by sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE). It was found that N applied to the canopy did not improve protein rate compared with N application to the soil (general mean 138 mg g-1 ), but moderately increased productivity in the high-yielding varieties Liberdur and Biensur (three-year means 7.23 vs 7.13 and 7.53 vs 7.09 t ha-1 respectively). Technological quality was mainly related to variety choice, Aureo and Ariosto having higher protein rates and glutenin/gliadin ratios. Also found was a strong 'variety × N application method' interaction in the proportions of protein subunits within each class, particularly LMW-GS and gliadins. A promising result was the higher N uptake efficiency, although as apparent balance, combined with higher HMW/LMW-GS ratio in var. Biensur. CONCLUSION: Late-season foliar N fertilization allows N fertilizer saving, potentially providing environmental benefits in the rainy climate of the northern Mediterranean area, and also leads to variety-dependent up-regulation of essential LMW-GS and gliadins. Variety choice is a key factor in obtaining high technological quality, although it is currently associated with modest grain yield. This study provides evidence of high quality in the specific high-yielding variety Biensur, suggesting its potential as a mono-varietal semolina for pasta production. © 2017 Society of Chemical Industry.

Native arbuscular mycorrhizal symbiosis alters foliar bacterial community composition.

The effects of arbuscular mycorrhizal (AM) fungi on plant-associated microbes are poorly known. We tested the hypothesis that colonization by an AM fungus affects microbial species richness and microbial community composition of host plant tissues. We grew the grass, Deschampsia flexuosa in a greenhouse with or without the native AM fungus, Claroideoglomus etunicatum. We divided clonally produced tillers into two parts: one inoculated with AM fungus spores and one without AM fungus inoculation (non-mycorrhizal, NM). We characterized bacterial (16S rRNA gene) and fungal communities (internal transcribed spacer region) in surface-sterilized leaf and root plant compartments. AM fungus inoculation did not affect microbial species richness or diversity indices in leaves or roots, but the AM fungus inoculation significantly affected bacterial community composition in leaves. A total of three OTUs in leaves belonging to the phylum Firmicutes positively responded to the presence of the AM fungus in roots. Another six OTUs belonging to the Proteobacteria (Alpha, Beta, and Gamma) and Bacteroidetes were significantly more abundant in NM plants when compared to AM fungus-inoculated plants. Further, there was a significant correlation between plant dry weight and leaf microbial community compositional shift. Also, there was a significant correlation between leaf bacterial community compositional shift and foliar nitrogen content changes due to AM fungus inoculation. The results suggest that AM fungus colonization in roots has a profound effect on plant physiology that is reflected in leaf bacterial community composition.

Interspecific vs intraspecific patterns in leaf nitrogen of forest trees across nitrogen availability gradients.

Leaf nitrogen content (δ) coordinates with total canopy N and leaf area index (LAI) to maximize whole-crown carbon (C) gain, but the constraints and contributions of within-species plasticity to this phenomenon are poorly understood. Here, we introduce a game theoretic, physiologically based community model of height-structured competition between late-successional tree species. Species are constrained by an increasing, but saturating, relationship between photosynthesis and leaf N per unit leaf area. Higher saturating rates carry higher fixed costs. For a given whole-crown N content, a C gain-maximizing compromise exists between δ and LAI. With greater whole-crown N, both δ and LAI increase within species. However, a shift in community composition caused by reduced understory light at high soil N availability (which competitively favors species with low leaf costs and consequent low optimal δ) counteracts the within-species response, such that community-level δ changes little with soil N availability. These model predictions provide a new explanation for the changes in leaf N per mass observed in data from three dominant broadleaf species in temperate deciduous forests of New England. Attempts to understand large-scale patterns in vegetation often omit competitive interactions and intraspecific plasticity, but here both are essential to an understanding of ecosystem-level patterns.

Legacy Effects in Buds and Leaves of European Beech Saplings (Fagus sylvatica) after Severe Drought.

Against the background of climate change, we studied the effects of a severe summer drought on buds of European beech (Fagus sylvatica L.) saplings and on leaves formed during the subsequent spring in trees attributed to different drought-damage classes. For the first time, we combined assessments of the vitality (assessed through histochemical staining), mass and stable carbon isotope ratios (δ13C) of buds from drought-stressed woody plants with morphological and physiological variables of leaves that have emerged from the same plants and crown parts. The number, individual mass and vitality of the buds decreased and δ13C increased with increasing drought-induced damage. Bud mass, vitality and δ13C were significantly intercorrelated. The δ13C of the buds was imprinted on the leaves formed in the subsequent spring, but individual leaf mass, leaf size and specific leaf area were not significantly different among damage classes. Vitality and δ13C of the buds are suitable indicators of the extent of preceding drought impact. Bud vitality may be used as a simple means of screening saplings for the flushing capability in the subsequent spring. European beech saplings are susceptible, but-due to interindividual differences-are resilient, to a certain extent, to a singular severe drought stress.

Non-invasive Estimation of Foliar Nitrogen Concentration Using Spectral Characteristics of Menthol Mint (Mentha arvensis L.).

Menthol mint (Mentha arvensis L., Family: Lamiaceae), popularly known as corn mint or Japanese mint, is an important industrial crop that is widely grown for its valued essential oil. Nitrogen (N) is an important macro-nutrient and an essential factor for optimizing the yield and quality of crops. Hence, rapid and accurate estimation of the N content is crucial for nutrient diagnosis in plants and to make precise N fertilizer recommendations. Generally, N concentration is estimated by destructive sampling methods; however, an indirect assessment may be possible based on spectral characteristics. This study aimed to compare the foliar N concentration based on non-destructive (reflectance) and destructive (laboratory analyses) methods in menthol mint. Foliar N concentration was measured through the Kjeldahl method and reflectance by Miniature Leaf Spectrometer C-710 (CID Bio-Science). Using reflectance data, several vegetation indices (VIs), that is, normalized difference red edge (NDRE), red edge normalized difference vegetation index (reNDVI), simple ratio (SR), green-red vegetation index (GRVI), canopy chlorophyll content index (CCCI), photochemical reflectance index (PRI), green chlorophyll index (CI Green ), red edge chlorophyll index (CI Red Edge ), canopy chlorophyll index (CCI), normalized pigment chlorophyll ratio index (NPCI), and structure insensitive pigment index (SIPI), were developed to determine the foliar N concentration. The highest correlation (r) between VIs and foliar N concentrations was achieved by NDRE (0.89), followed by reNDVI (0.84), SR (0.83), GRVI (0.78), and CCCI (0.76). Among the VIs, the NDRE index has been found to be the most accurate index that can precisely predict the foliar N concentration (R 2 = 0.79, RMSE = 0.18). In summary, the N deficiencies faced by the crop during its growth period can be detected effectively by calculating NDRE and reNDVI, which can be used as indicators for recommending precise management strategies for the application of nitrogenous fertilizers.

Long-term effects of artificial nighttime lighting and trophic complexity on plant biomass and foliar carbon and nitrogen in a grassland community.

The introduction of artificial nighttime lighting due to human settlements and transport networks is increasingly altering the timing, intensity, and spectra of natural light regimes worldwide. Much of the research on the impacts of nighttime light pollution on organisms has focused on animal species. Little is known about the impacts of daylength extension due to outdoor lighting technologies on wild plant communities, despite the fact that plant growth and development are under photoperiodic control. In a five-year field experiment, artificial ecosystems ("mesocosms") of grassland communities both alone or in combination with invertebrate herbivores and predators were exposed to light treatments that simulated street lighting technologies (low-pressure sodium, and light-emitting diode [LED]-based white lighting), at ground-level illuminance. Most of the plant species in the mesocosms did not exhibit changes in biomass accumulation after 5 years of exposure to the light treatments. However, the white LED treatment had a significant negative effect on biomass production in the herbaceous species Lotus pedunculatus. Likewise, the interaction between the white LED treatment and the presence of herbivores significantly reduced the mean shoot/root ratio of the grass species Holcus lanatus. Artificial nighttime lighting had no effect on the foliar carbon or nitrogen in most of the grassland species. Nevertheless, the white LED treatment significantly increased the leaf nitrogen content in Lotus corniculatus in the presence of herbivores. Long-term exposure to artificial light at night had no general effects on plant biomass responses in experimental grassland communities. However, species-specific and negative effects of cool white LED lighting at ground-level illuminance on biomass production and allocation in mixed plant communities are suggested by our findings. Further studies on the impacts of light pollution on biomass accumulation in plant communities are required as these effects could be mediated by different factors, including herbivory, competition, and soil nutrient availability.

Accounting for the effect of temperature in clarifying the response of foliar nitrogen isotope ratios to atmospheric nitrogen deposition.

Atmospheric nitrogen deposition affects nitrogen isotope composition (δ15N) in plants. However, both negative effect and positive effect have been reported. The effects of climate on plant δ15N have not been corrected for in previous studies, this has impeded discovery of a true effect of atmospheric N deposition on plant δ15N. To obtain a more reliable result, it is necessary to correct for the effects of climatic factors. Here, we measured δ15N and N contents of plants and soils in Baiwangshan and Mount Dongling, north China. Atmospheric N deposition in Baiwangshan was much higher than Mount Dongling. Generally, however, foliar N contents showed no difference between the two regions and foliar δ15N was significantly lower in Baiwangshan than Mount Dongling. The corrected foliar δ15N after accounting for a predicted value assumed to vary with temperature was obviously more negative in Baiwangshan than Mount Dongling. Thus, this suggested the necessity of temperature correction in revealing the effect of N deposition on foliar δ15N. Temperature, soil N sources and mycorrhizal fungi could not explain the difference in foliar δ15N between the two regions, this indicated that atmospheric N deposition had a negative effect on plant δ15N. Additionally, this study also showed that the corrected foliar δ15N of bulk data set increased with altitude above 1300m in Mount Dongling, this provided an another evidence for the conclusion that atmospheric N deposition could cause 15N-depletion in plants.

A coastal and an interior Douglas fir provenance exhibit different metabolic strategies to deal with drought stress.

Drought is a major environmental stress affecting growth and vitality of forest ecosystems. In the present study, foliar nitrogen (N) and carbon (C) metabolism of two Douglas fir (Pseudotsuga menziesii) provenances with assumed different drought tolerance were investigated. We worked with 1-year-old seedlings of the interior provenance Fehr Lake (FEHR) originating from a dry environment and the coastal provenance Snoqualmie (SNO) from a more humid origin. Total C and N, structural N and the concentrations of soluble protein, total amino acids (TAAs) and individual amino acids as well as the relative abundance of polar, low-molecular-weight metabolites including antioxidants were determined in current-year needles exposed either to 42 days of drought or to 42 days drought plus 14 days of rewatering. The seedlings reacted in a provenance-specific manner to drought stress. Coastal provenance SNO showed considerably increased contents of TAAs, which were caused by increased abundance of the quantitatively most important amino acids arginine, ornithine and lysine. Additionally, the polyamine putrescine accumulated exclusively in drought-stressed trees of this provenance. In contrast, the interior provenance FEHR showed the opposite response, i.e., drastically reduced concentrations of these amino acids. However, FEHR showed considerably increased contents of pyruvate-derived and aromatic amino acids, and also higher drought-induced levels of the antioxidants ascorbate and α-tocopherol. In response to drought, both provenances produced large amounts of carbohydrates, such as glucose and fructose, most likely as osmolytes that can readily be metabolized for protection against osmotic stress. We conclude that FEHR and SNO cope with drought stress in a provenance-specific manner: the coastal provenance SNO was mainly synthesizing N-based osmolytes, a reaction not observed in the interior provenance FEHR; instead, the latter increased the levels of scavengers of reactive oxygen species. Our results underline the importance of provenance-specific reactions to abiotic stress.

Patterns in hydraulic architecture from roots to branches in six tropical tree species from cacao agroforestry and their relation to wood density and stem growth.

For decades it has been assumed that the largest vessels are generally found in roots and that vessel size and corresponding sapwood area-specific hydraulic conductivity are acropetally decreasing toward the distal twigs. However, recent studies from the perhumid tropics revealed a hump-shaped vessel size distribution. Worldwide tropical perhumid forests are extensively replaced by agroforestry systems often using introduced species of various biogeographical and climatic origins. Nonetheless, it is unknown so far what kind of hydraulic architectural patterns are developed in those agroforestry tree species and which impact this exerts regarding important tree functional traits, such as stem growth, hydraulic efficiency and wood density (WD). We investigated wood anatomical and hydraulic properties of the root, stem and branch wood in Theobroma cacao and five common shade tree species in agroforestry systems on Sulawesi (Indonesia); three of these were strictly perhumid tree species, and the other three tree species are tolerating seasonal drought. The overall goal of our study was to relate these properties to stem growth and other tree functional traits such as foliar nitrogen content and sapwood to leaf area ratio. Our results confirmed a hump-shaped vessel size distribution in nearly all species. Drought-adapted species showed divergent patterns of hydraulic conductivity, vessel density, and relative vessel lumen area between root, stem and branch wood compared to wet forest species. Confirming findings from natural old-growth forests in the same region, WD showed no relationship to specific conductivity. Overall, aboveground growth performance was better predicted by specific hydraulic conductivity than by foliar traits and WD. Our study results suggest that future research on conceptual trade-offs of tree hydraulic architecture should consider biogeographical patterns underlining the importance of anatomical adaptation mechanisms to environment.

A generalized, lumped-parameter model of photosynthesis, evapotranspiration and net primary production in temperate and boreal forest ecosystems.

PnET is a simple, lumped-parameter, monthlytime-step model of carbon and water balances of forests built on two principal relationships: 1) maximum photosynthetic rate is a function of foliar nitrogen concentration, and 2) stomatal conductance is a function of realized photosynthetic rate. Monthyly leaf area display and carbon and water balances are predicted by combining these with standard equations describing light attenuation in canopies and photosynthetic response to diminishing radiation intensity, along with effects of soil water stress and vapor pressure deficit (VPD). PnET has been validated against field data from 10 well-studied temperate and boreal forest ecosystems, supporting our central hypothesis that aggregation of climatic data to the monthly scale and biological data such as foliar characteristics to the ecosystem level does not cause a significant loss of information relative to long-term, mean ecosystem responses. Sensitivity analyses reveal a diversity of responses among systems to identical alterations in climatic drivers. This suggests that great care should be used in developing generalizations as to how forests will respond to a changing climate. Also critical is the degree to which the temperature responses of photosynthesis and respiration might acclimate to changes in mean temperatures at decadal time scales. An extreme climate change simulation (+3° C maximum temperature, -25% precipitation with no change in minimum temperature or radiation, direct effects of increased atmospheric CO2 ignored) suggests that major increases in water stress, and reductions in biomass production (net carbon gain) and water yield would follow such a change.

Extrapolating leaf CO2 exchange to the canopy: a generalized model of forest photosynthesis compared with measurements by eddy correlation.

Over the last 4 years, two data sets have emerged which allow increased accuracy and resolution in the definition and validation of a photosynthesis model for whole forest canopies. The first is a greatly expanded set of data on the nitrogen-photosynthesis relationship for temperate and tropical woody species. The second is a unique set of long-term (4 year) daily carbon balance measurements at the Harvard Forest, Petersham, Massachusetts, collected by the eddy-correlation technique. A model (PhET-Day) is presented which is derived directly from, and validated against, these data sets. The PnET-Day model uses foliar nitrogen concentration to calculate maximum instantaneous rates of gross and net photosynthesis which are then reduced for suboptimal temperature, photosynthetically active radiation (PAR), and vapor pressure deficit (VPD). Predicted daily gross photosynthesis is closely related to gross carbon exchange at the Harvard Forest as determined by eddy-correlation measurements. Predictions made by the full canopy model were significantly better than those produced by a multiple linear regression model. Sensitivity analyses for this model for a deciduous broad-leaved forest showed results to be much more sensitive to parameters related to maximum leaf-level photosynthetic rate (A max) than to those related to light, temperature, VPD or total foliar mass. Aggregation analyses suggest that using monthly mean climatic data to drive the canopy model will give results similar to those achieved by averaging daily eddy correlation measurements of gross carbon exchange (GCE).

Foliar carbon isotope discrimination in Larix species and sympatric evergreen conifers: a global comparison.

Larches (Larix spp.), deciduous conifers, occur in the northern hemisphere in cold-temperate and boreal climates - an environment normally thought to favor evergreen tree species. We compare foliar carbon isotope discrimination (Δ), instantaneous water use efficiency, total foliar nitrogen concentration, and specific leaf area (for a subset of sites) between Larix spp. and co-occurring evergreen conifers at 20 sites throughout the natural range of larches. Except for Larix occidentalis in the xeric Intermountain West, USA, Δ is significantly (P < 0.05) greater for larches than co-occurring evergreen conifers at 77% of the sites, suggesting that larches use water less efficiently. At elevations greater than 3000 m, the Δ of Larix spp. and co-occurring conifers converge, suggesting that water is not the limiting resource. Foliar nitrogen concentration and specific leaf area are two ecophysiological characteristics that are positively correlated with high photosynthetic capacity. Foliar nitrogen concentration is significantly greater for larches than evergreen conifers at 88% of the sites and specific leaf area is approximately three times greater for larches than co-occurring conifers. Future studies should examine the potential effect that global warming may have on the distribution of larch forests because the water use efficiency of larches is commonly less than co-occurring evergreen conifers and the boreal and high-latitude environments are likely to experience the greatest climate warming.