Drought legacies in mixed Mediterranean forests: Analysing the effects of structural overshoot, functional traits and site factors.

Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests.

Bell-shaped tree-ring responses to air temperature drive productivity trends in long-lived mountain Mediterranean pines.

We investigated the dendroclimatic response of a Pinus heldreichii metapopulation distributed over a wide elevation interval (from 882 to 2143 m a.s.l.), spanning from low mountain to upper subalpine vegetation belts in the southern Italian Apennines. The tested hypothesis is that wood growth along an elevational gradient is non-linearly related to air temperature. During three years of fieldwork (2012-2015) at 24 sites, we collected wood cores from a total of 214 pine trees with diameter at breast height from 19 to 180 cm (average 82.7 ± 32.9 cm). We used a combination of tree-ring and genetic methods to reveal factors involved in growth acclimation using a space-for-time approach. Scores from canonical correspondence analysis were used to combine individual tree-ring series into four composite chronologies related to air temperature along the elevation gradient. Overall, the June dendroclimatic response followed a bell-shaped thermal niche curve, increasing until a peak around 13-14 °C. A similarly bell-shaped response was found with previous autumn air temperature, and both dendroclimatic signals interacted with stem size and growth rates, generating a divergent growth response between the top and the bottom of the elevation gradient. Increased tree growth in the upper subalpine belt was consistent with the consequences of increasing air temperature under no drought stress. A positive link was uncovered between pine growth at all elevations and April mean temperature, with trees growing at the lowest elevations showing the strongest growth response. No elevational genetic differences were found, hence long-lived tree species with small geographical ranges may reverse their climatic response between the lower and upper bioclimatic zones of their environmental niche. Our study revealed a high resistance and acclimation capability of Mediterranean forest stands, and such low vulnerability to changing climatic conditions highlights the potential to store carbon in these ecosystems for the coming decades.

Tree-ring and remote sensing analyses uncover the role played by elevation on European beech sensitivity to late spring frost.

Extreme climate events such as late spring frosts (LSFs) negatively affect productivity and tree growth in temperate beech forests. However, detailed information on how these forests recover after such events are still missing. We investigated how LSFs affected forest cover and radial growth in European beech (Fagus sylvatica L.) populations located at different elevations at four sites in the Italian Apennines, where LSFs have been recorded. We combined tree-ring and remote-sensing data to analyse the sensitivity and recovery capacity of beech populations to LSFs. Using daily temperature records, we reconstructed LSF events and assessed legacy effects on growth. We also evaluated the role played by elevation and stand structure as modulators of LSFs impacts. Finally, using satellite images we computed Normalized Difference Vegetation Index (NDVI), Enhanced Vegetation Index (EVI) and LAI (Leaf Area Index) to evaluate the post-LSF canopy recovery. The growth reduction in LSF-affected trees ranged from 36 % to 84 %. We detected a negative impact of LSF on growth only during the LSF year, with growth recovery occurring within 1-2 years after the event. LSF-affected stands featured low vegetation indices until late June, i.e. on average 75 days after the frost events. We did not find a clear relationship between beech forest elevation and occurrence of LSFs defoliations. Our results indicate a high recovery capacity of common beech and no legacy effects of LSFs.

Towards sustainable management of forest residues in the southern Apennine Mediterranean mountain forests: a scenario-based approach.

Key message: Managing forest residues according to the carbon content of the soil helps to minimize the ecological footprint of their removal. Context: In Mediterranean mountain ecosystems, unsustainable harvesting of wood residues might contribute to land degradation, carbon, and nutrient depletion in forest soils. Aims: This study aimed to assess the amount of forest biomass residues that should be left on-site to minimize the depletion of soil fertility. Methods: We estimated the availability of biomass residues in the public forest land of the Basilicata region of Southern Italy by collecting stand-scale inventory attributes from forest management plans. Subsequently, we quantified the amount of forest biomass residue released by implementing a scenario-based approach. Results: Approximately 5800 m3 year-1 of forest residues could be potentially available for bio-based industries at the regional scale within the next 10 years. Such residues mainly belong to broadleaved forest types, having a high variability in their soil organic stock (228.5-705.8 Mg C ha-1) and altitudinally spanning from 400 to 1500 m a.s.l. In these forests, the simulated scenarios displayed a wide range of average harvestable residues from 2.5 to 5.5 m3 ha-1, containing approximately 1.1 to 2.1 Mg ha-1 of organic carbon. Conclusion: Our study suggests that forest management plans are a useful source of information to estimate the available forest biomass residues consistently. In southern Mediterranean mountain forests, the management of forest residues according to soil carbon content helps to minimize the environmental impact and increase their sustainability.

Declines in canopy greenness and tree growth are caused by combined climate extremes during drought-induced dieback.

Several dieback episodes triggered by droughts are revealing the high vulnerability of Mediterranean forests, manifested as declines in growth, increased defoliation, and rising mortality rates. Understanding forest responses to such climate extreme events is of high priority for predicting their future vegetation dynamics. We examined how remotely sensed measures of vegetation activity (NDVI, Normalized Difference Vegetation Index) and radial growth (BAI, basal area increment) responded to climate extreme events. We considered tree (Pinus sylvestris, Quercus pubescens, Quercus frainetto) and shrub (Juniperus phoenicea) populations from Italy and Spain showing recent dieback phenomena. Two components of drought, namely elevated atmospheric demand (VPD, vapor pressure deficit) and low soil moisture were analyzed in nearby stands showing or not showing dieback symptoms. Dieback stands exhibited lower NDVI values than non-dieback stands. NDVI and BAI were positively related in all sites except for the dieback stand of Q. frainetto that was negatively related. Such NDVI-BAI linkages were related to specific time windows, which could be useful for identifying when climatic conditions have the greatest influence on vegetation. Growth decline occurred in response to increasing VPD, but responses differed among species. J. phoenicea was the most negatively impacted by higher VPD, whereas oaks responded to soil moisture. A high VPD was related to stronger growth reduction in dieback P. sylvestris trees regardless of soil moisture changes. We highlighted that coupling between proxies of forest productivity (NDVI, BAI) allows better understanding and forecasting of drought-induced dieback phenomena in forests and shrublands. Scaling up from tree to stand levels might be feasible when using the maximum growing season NDVI, which can be applied for retrospective modeling of the impact of drought stress on forest productivity and tree growth.

Mediterranean old-growth forests exhibit resistance to climate warming.

Old-growth mountain forests represent an ideal setting for studying long-term impacts of climate change. We studied the few remnants of old-growth forests located within the Pollino massif (southern Italy) to evaluate how the growth of conspecific young and old trees responded to climate change. We investigated two conifer species (Abies alba and Pinus leucodermis) and two hardwood species (Fagus sylvatica and Quercus cerris). We sampled one stand per species along an altitudinal gradient, ranging from a drought-limited low-elevation hardwood forest to a cold-limited subalpine pine forest. We used a dendrochronological approach to characterize the long-term growth dynamics of old (age > 120 years) versus young (age < 120 years) trees. Younger trees grew faster than their older conspecifics during their juvenile stage, regardless of species. Linear mixed effect models were used to quantify recent growth trends (1950-2015) and responses to climate for old and young trees. Climate sensitivity, expressed as radial growth responses to climate during the last three decades, partially differed between species because high spring temperatures enhanced conifer growth, whereas F. sylvatica growth was negatively affected by warmer spring conditions. Furthermore, tree growth was negatively impacted by summer drought in all species. Climate sensitivity differed between young and old trees, with younger trees tending to be more sensitive in P. leucodermis and A. alba, whereas older F. sylvatica trees were more sensitive. In low-elevation Q. cerris stands, limitation of growth due to drought was not related to tree age, suggesting symmetric water competition. We found evidence for a fast-growth trend in young individuals compared with that in their older conspecifics. Notably, old trees tended to have relatively stable growth rates, showing remarkable resistance to climate warming. These responses to climate change should be recognized when forecasting the future dynamics of old-growth forests for their sustainable management.

A meta-analysis of mesophyll conductance to CO2 in relation to major abiotic stresses in poplar species.

Mesophyll conductance (gm) determines the diffusion of CO2 from the substomatal cavities to the site of carboxylation in the chloroplasts and represents a critical component of the diffusive limitation of photosynthesis. In this study, we evaluated the average effect sizes of different environmental constraints on gm in Populus spp., a forest tree model. We collected raw data of 815 A-Ci response curves from 26 datasets to estimate gm, using a single curve-fitting method to alleviate method-related bias. We performed a meta-analysis to assess the effects of different abiotic stresses on gm. We found a significant increase in gm from the bottom to the top of the canopy that was concomitant with the increase of maximum rate of carboxylation and light-saturated photosynthetic rate (Amax). gm was positively associated with increases in soil moisture and nutrient availability, but was insensitive to increasing soil copper concentration and did not vary with atmospheric CO2 concentration. Our results showed that gm was strongly related to Amax and to a lesser extent to stomatal conductance (gs). Moreover, a negative exponential relationship was obtained between gm and specific leaf area, which may be used to scale-up gm within the canopy.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Variation in the access to deep soil water pools explains tree-to-tree differences in drought-triggered dieback of Mediterranean oaks.

Individual differences in the access to deep soil water pools may explain the differential damage among coexisting, conspecific trees as a consequence of drought-induced dieback. We addressed this issue by comparing the responses to a severe drought of three Mediterranean oak species with different drought tolerance, Quercus pubescens L. and Quercus frainetto Ten., mainly thriving at xeric and mesic sites, respectively, and Quercus cerris L., which dominates at intermediate sites. For each species, we compared coexisting declining (D) and non-declining (ND) trees. The stable isotope composition (δ2H, δ18O) of xylem and soil water was used to infer a differential use of soil water sources. We also measured tree size and radial growth to quantify the long-term divergence of wood production between D and ND trees and non-structural carbohydrates (NSCs) in sapwood to evaluate if D trees presented lower NSC values. The ND trees had access to deeper soil water than D trees except in Q. frainetto, as indicated by significantly more depleted xylem water values. However, a strong δ2H offset between soil and xylem water isotopes observed in peak summer could suggest that both tree types were not physiologically active under extreme drought conditions. Alternative processes causing deuterium fractionation, however, could not be ruled out. Tree height and recent (last 15-25 years) growth rates in all species studied were lower in D than in ND trees by 22 and 44%, respectively. Lastly, there was not a consistent pattern of NSC sapwood concentration; in Q. pubescens, it was higher in ND trees while in Q. frainetto, the D trees were the ones exhibiting the higher NSC concentration. We conclude that the vulnerability to drought among conspecific Mediterranean oaks depends on the differential access to deep soil water pools, which may be related to differences in rooting depth, tree size and growth rate.

The impact of drought spells on forests depends on site conditions: The case of 2017 summer heat wave in southern Europe.

A major component of climate change is an increase in temperature and precipitation variability. Over the last few decades, an increase in the frequency of extremely warm temperatures and drought severity has been observed across Europe. These warmer and drier conditions may reduce productivity and trigger compositional shifts in forest communities. However, we still lack a robust, biogeographical characterization of the negative impacts of climate extremes, such as droughts on forests. In this context, we investigated the impact of the 2017 summer drought on European forests. The normalized difference vegetation index (NDVI) was used as a proxy of forest productivity and was related to the standardized precipitation evapotranspiration index, which accounts for the temperature effects of the climate water balance. The spatial pattern of NDVI reduction in 2017 was largely driven by the extremely warm summer for parts of the central and eastern Mediterranean Basin (Italian and Balkan Peninsulas). The vulnerability to the 2017 summer drought was heterogeneously distributed over Europe, and topographic factors buffered some of the negative impacts. Mediterranean forests dominated by oak species were the most negatively impacted, whereas Pinus pinaster was the most resilient species. The impact of drought on the NDVI decreased at high elevations and mainly on east and north-east facing slopes. We illustrate how an adequate characterization of the coupling between climate conditions and forest productivity (NDVI) allows the determination of the most vulnerable areas to drought. This approach could be widely used for other extreme climate events and when considering other spatially resolved proxies of forest growth and health.

Drought and Phytophthora Are Associated With the Decline of Oak Species in Southern Italy.

Forest decline induced by climate change is a global phenomenon that affects many tree species, mainly in drought-prone areas as the Mediterranean region. In southern Italy, several oak species have shown decline symptoms and elevated mortality since the 2000s due to drought stress. However, it remains to be answered whether decline occurred alone or whether a pathogen was also involved. To this aim, we compared two coexisting oak species in a forest located in southern Italy which are assumed to be less (Quercus cerris) and more tolerant to drought (Quercus pubescens). We sampled fifteen couples of neighboring declining (D) and non-declining (ND) trees of both species. Wood cores were taken from all trees to perform dendrochronological analyses to detect the decline onset and link it to potential climatic drivers. Carbon isotope ratios (d13C) were analyzed in wood of the two vigor classes to compare their water-use efficiency. Phytophthora presence was also assessed in soil samples from ten D-ND couples of trees per species. The oak species most affected by drought-induced decline in terms of leaf shedding and mortality was Q. cerris, i.e., the least tolerant to drought. In both species, the D trees showed a reduced growth rate compared with ND trees from 2000 onward when drought and warming intensified. Q. pubescens showed higher growth sensitivity to precipitation, temperature and drought than Q. cerris. This sensitivity to climate was magnified in D trees whose growth decreased in response to warm and dry conditions during the prior winter and the late summer. The Q. pubescens D trees were more efficient in their water use than ND trees before the growth divergence between D and ND trees amplified. In the studied area, Phytophthora quercina was isolated from 40% of the sampled trees, and tended to be more frequent amongst ND than amongst D trees. Our data suggests that droughts and warm summer conditions triggered oak decline. The high prevalence of P. quercina in the studied area warrants further study as a potential predisposing factor.

Size Matters a Lot: Drought-Affected Italian Oaks Are Smaller and Show Lower Growth Prior to Tree Death.

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

A multi-proxy assessment of dieback causes in a Mediterranean oak species.

Drought stress causes forest dieback that is often explained by two interrelated mechanisms, namely hydraulic failure and carbon starvation. However, it is still unclear which functional and structural alterations, related to these mechanisms, predispose to dieback. Here we apply a multi-proxy approach for the characterization of tree structure (radial growth, wood anatomy) and functioning (δ13C, δ18O and non-structural carbohydrates (NSCs)) in tree rings before and after drought-induced dieback. We aim to discriminate which is the main mechanism and to assess which variables can act as early-warning proxies of drought-triggered damage. The study was tailored in southern Italy in two forests (i.e., San Paolo (SP) and Oriolo (OR)) where declining and non-declining trees of a ring-porous tree species (Quercus frainetto Ten.) showing anisohydric behavior coexist. Both stands showed growth decline in response to warm and dry spring conditions, although the onset of dieback was shifted between them (2002 in SP and 2009 in OR). Declining trees displayed a sharp growth drop after this onset with reductions of 49% and 44% at SP and OR sites, respectively. Further, contrary to what we expected, declining trees showed a lower intrinsic water-use efficiency compared with non-declining trees after the dieback onset (with reductions of 9.7% and 5.6% at sites SP and OR, respectively), due to enhanced water loss through transpiration, as indicated by the lower δ18O values. This was more noticeable at the most drought-affected SP stand. Sapwood NSCs did not differ between declining and non-declining trees, indicating no carbon starvation in affected trees. Thus, the characterized structural and functional alterations partially support the hydraulic failure mechanism of dieback. Finally, we show that growth data are reliable early-warning proxies of drought-triggered dieback.

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.

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.

Environmental effects on oxygen isotope enrichment of leaf water in cotton leaves.

The oxygen isotope enrichment of bulk leaf water (Delta(b)) was measured in cotton (Gossypium hirsutum) leaves to test the Craig-Gordon and Farquhar-Gan models under different environmental conditions. Delta(b) increased with increasing leaf-to-air vapor pressure difference (VPd) as an overall result of the responses to the ratio of ambient to intercellular vapor pressures (e(a)/e(i)) and to stomatal conductance (g(s)). The oxygen isotope enrichment of lamina water relative to source water (Delta(1)), which increased with increasing VPd, was estimated by mass balance between less enriched water in primary veins and enriched water in the leaf. The Craig-Gordon model overestimated Delta(b) (and Delta(1)), as expected. Such discrepancies increased with increase in transpiration rate (E), supporting the Farquhar-Gan model, which gave reasonable predictions of Delta(b) and Delta(1) with an L of 7.9 mm, much less than the total radial effective length L(r) of 43 mm. The fitted values of L for Delta(1) of individual leaves showed little dependence on VPd and temperature, supporting the assumption that the Farquhar-Gan formulation is relevant and useful in describing leaf water isotopic enrichment.

Variation in nitrogen supply changes water-use efficiency of Pseudotsuga menziesii and Populus x euroamericana; a comparison of three approaches to determine water-use efficiency.

We studied the effects of three nitrogen (N) supply rates (low, intermediate and high) on Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings and poplar clone "I-214" (Populus x euroamericana (Dole) Guinier) cuttings growing in mini-stands. Our specific objectives were to: (1) evaluate the effects of N supply on water-use efficiency (WUE) and biomass production; (2) determine if N affects WUE through control of carbon assimilation rates or through stomatal control of water loss; and (3) compare three methods of estimating WUE: one short-term method (WUE(i), based on gas exchange measurements) and two long-term methods (WUE(T), based on the ratio between biomass production and transpired water, and Delta, based on leaf carbon isotope discrimination tested as a proxy of WUE). In both species, biomass production, WUE(i) and WUE(T) increased with increasing N supply, but there was no effect of N supply on either transpiration or stomatal conductance and Delta was negatively related to leaf N concentration. Plots of Delta versus both WUE(i) and WUE(T) revealed negative trends, but the regression between WUE(i) and Delta was significant only for Douglas-fir, and the regression between WUE(T) and Delta was significant only for poplar. Thus, the mechanisms underlying the response of WUE to N supply were mainly related to a positive effect of N supply on photosynthetic rates. The data confirm that carbon isotope discrimination may be a useful proxy of WUE. The finding that N availability enhances both biomass production and WUE may have practical implications in regions where these factors impose constraints on forest productivity.

Photosynthesis-nitrogen relationships: interpretation of different patterns between Pseudotsuga menziesii and Populus x euroamericana in a mini-stand experiment.

We compared photosynthesis-nitrogen relationships of one broad-leaved (poplar; Populus x euroamericana (Dole) Guinier) and one conifer (Douglas-fir; Pseudotsuga menziesii (Mirb.) Franco) species. Plants were grown in large pots to allow free root development and were kept well watered. We determined effects of low, intermediate and high nitrogen supply rates on area-based leaf nitrogen (Na) and chlorophyll concentrations, leaf mass per area (LMA), light-saturated photosynthesis (Amax), maximum carboxylation (Vcmax) and electron transport rate (Jmax), photosynthetic nitrogen-use efficiency (PNUE), and proportions of leaf N in active Rubisco (PR), bioenergetic pools (PB) and the light-harvesting complex (PLH). Nitrogen supply significantly affected leaf Na. Leaf mass per area did not differ between species and was unaffected by the N treatments. In both species, there was a positive correlation between leaf Na and chlorophyll concentration, and between leaf Na and the photosynthetic parameters Amax, Jmax and Vcmax. At comparable leaf Na, however, poplar showed twofold higher PNUE and a threefold steeper slope of the Amax- nitrogen relationship than Douglas-fir. Leaf Na was negatively correlated with PNUE in Douglas-fir but not in poplar. Leaf Na was also negatively correlated with PR, PB and PLH in Douglas-fir, whereas in poplar, a negative correlation was found only for PLH. Parameter PR was significantly higher in poplar than in Douglas-fir. The ratio of CO2 concentration in the intercellular space to that in ambient air was higher in poplar than in Douglas-fir. Overall, our data suggest that differences in the photosynthesis-nitrogen relationship and PNUE between Douglas-fir and poplar primarily reflect a different investment of N to active Rubisco, and possibly a different constraint to CO2 diffusion.