Rossella Guerrieri

Assessing the effects of nitrogen deposition and climate on carbon isotope discrimination and intrinsic water-use efficiency of angiosperm and conifer trees under rising CO2 conditions.

The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO2 , on the variability of carbon isotope discrimination (Δ(13) C), and intrinsic water-use efficiency (iWUE) of angiosperm and conifer tree species. Eighty-nine long-term isotope tree-ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long-term time series of Δ(13) C and iWUE. Δ(13) C and iWUE were related to the increasing concentration of atmospheric CO2 over the industrial period (1850-2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950-2000. We applied generalized additive models and linear mixed-effects models to predict the effects of climatic variables and nitrogen deposition on Δ(13) C and iWUE. Results showed a declining Δ(13) C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO2 values. The temporal variation in Δ(13) C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO2 concentrations. We defined linear mixed-effects models that were effective to describe the variation of Δ(13) C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (Nrate ) and long-term cumulative (Ncum ) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ(13) C and iWUE showed complex dependent interactions between different covariates. A significant association of Nrate with iWUE and Δ(13) C was observed in conifers and in the angiosperms, and Ncum was the only independent term with a significant positive association with iWUE, although a multi-factorial control was evident in conifers.

Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru.

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (Vcmax ), and the maximum rate of electron transport (Jmax )), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (Ma , Na and Pa , respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO2 -fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf Pa were key explanatory factors for models of area-based Vcmax and Jmax but did not account for variations in photosynthetic N-use efficiency. At any given Na and Pa , the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests.

Nutritional regulation in mixotrophic plants: new insights from Limodorum abortivum

Partially mycoheterotrophic (mixotrophic) plants gain carbon from both photosynthesis and their mycorrhizal fungi. This is considered an ancestral state in the evolution of full mycoheterotrophy, but little is known about this nutrition, and especially about the physiological balance between photosynthesis and fungal C gain. To investigate possible compensation between photosynthesis and mycoheterotrophy in the Mediterranean mixotrophic orchid Limodorum abortivum, fungal colonization was experimentally reduced in situ by fungicide treatment. We measured photosynthetic pigments of leaves, stems, and ovaries, as well as the stable C isotope compositions (a proxy for photosynthetic C gain) of seeds and the sizes of ovaries and seeds. We demonstrate that (1) in natural conditions, photosynthetic pigments are most concentrated in ovaries; (2) pigments and photosynthetic C increase in ovaries when fungal C supply is impaired, buffering C limitations and allowing the same development of ovaries and seeds as in natural conditions; and (3) responses to light of pigment and 13C contents in ovaries shift from null responses in natural conditions to responses typical of autotrophic plants in treated L. abortivum, demonstrating photoadaptation and enhanced use of light in the latter. L. abortivum thus preferentially feeds on fungi in natural conditions, but employs compensatory photosynthesis to buffer fungal C limitations and allow seed development.

Anthropogenic NOx emissions alter the intrinsic water-use efficiency (WUEi) for Quercus cerris stands under Mediterranean climate conditions.

We investigated the effect of N deposition (Ndep) on intrinsic water-use efficiency (WUEi), the ratio of photosynthesis (A) to stomatal conductance (gs), for two Quercus cerris stands at different distances to an oil refinery in Southern Italy. We used delta13C in tree rings for assessing changes in WUEi; while the influence of climate and NOx emission was explored through delta18O and delta15N, respectively. Differences in WUEi between the two sites were significant, with trees exposed to different degrees of NOx emissions showing an abrupt increase with the onset of pollution. Assuming similar gs at the two sites, as inferred through delta18O, the higher N availability at the polluted site caused the shift of the A/gs ratio in favour of A. Overall, our result suggests that an increase of Ndep may enhance tree WUE under a scenario of reduction of precipitation predicted for Mediterranean area.

Response of Quercus velutina growth and water use efficiency to climate variability and nitrogen fertilization in a temperate deciduous forest in the northeastern USA

Nitrogen (N) deposition and changing climate patterns in the northeastern USA can influence forest productivity through effects on plant nutrient relations and water use. This study evaluates the combined effects of N fertilization, climate and rising atmospheric CO2on tree growth and ecophysiology in a temperate deciduous forest. Tree ring widths and stable carbon (δ(13)C) and oxygen (δ(18)O) isotopes were used to assess tree growth (basal area increment, BAI) and intrinsic water use efficiency (iWUE) ofQuercus velutinaLamb., the dominant tree species in a 20+ year N fertilization experiment at Harvard Forest (MA, USA). We found that fertilized trees exhibited a pronounced and sustained growth enhancement relative to control trees, with the low- and high-N treatments responding similarly. All treatments exhibited improved iWUE over the study period (1984-2011). Intrinsic water use efficiency trends in the control trees were primarily driven by changes in stomatal conductance, while a stimulation in photosynthesis, supported by an increase in foliar %N, contributed to enhancing iWUE in fertilized trees. All treatments were predominantly influenced by growing season vapor pressure deficit (VPD), with BAI responding most strongly to early season VPD and iWUE responding most strongly to late season VPD. Nitrogen fertilization increasedQ. velutinasensitivity to July temperature and precipitation. Combined, these results suggest that ambient N deposition in N-limited northeastern US forests has enhanced tree growth over the past 30 years, while rising ambient CO2has improved iWUE, with N fertilization and CO2having synergistic effects on iWUE.

Evapotranspiration and water use efficiency in relation to climate and canopy nitrogen in U.S. forests

Understanding relations among forest carbon (C) uptake and water use is critical for predicting forest-climate interactions. Although the basic properties of tree-water relations have long been known, our understanding of broader-scale patterns is limited by several factors including: 1) incomplete understanding of drivers of change in coupled C and water fluxes, and water use efficiency (WUE); 2) difficulty in reconciling WUE estimates obtained at different scales; and 3) uncertainty in how evapotranspiration (ET) and WUE vary with other important resources such as nitrogen (N). To address these issues, we examined ET, gross primary production (GPP) and WUE at eleven AmeriFlux sites across North America. Our analysis spanned leaf and ecosystem scales, and included foliar δ13C, δ18O and %N measurements, eddy covariance estimates of GPP and ET, and remotely sensed estimates of canopy %N. We used flux data to derive ecosystem WUE (WUEe) and foliar δ13C to infer intrinsic WUE (iWUE). We found that GPP, ET and WUEe scaled with canopy %N, even when environmental variables were considered, and discuss the implications of these relationships for forest-atmosphere-climate interactions. We observed opposing patterns of WUE at leaf and ecosystem scales, and examined uncertainties to help explain these opposing patterns. Nevertheless, significant relationship between C isotope-derived ci/ca and GPP indicates that δ13C can be an effective predictor of forest GPP. Finally, we show that incorporating species functional traits - wood anatomy, hydraulic strategy and foliar %N -into a conceptual model improved the interpretation of Δ13C and δ18O vis-a-vis leaf to canopy water-carbon fluxes.