A trade-off between plant and soil carbon storage under elevated CO2.

Terrestrial ecosystems remove about 30 per cent of the carbon dioxide (CO2) emitted by human activities each year1, yet the persistence of this carbon sink depends partly on how plant biomass and soil organic carbon (SOC) stocks respond to future increases in atmospheric CO2 (refs. 2,3). Although plant biomass often increases in elevated CO2 (eCO2) experiments4-6, SOC has been observed to increase, remain unchanged or even decline7. The mechanisms that drive this variation across experiments remain poorly understood, creating uncertainty in climate projections8,9. Here we synthesized data from 108 eCO2 experiments and found that the effect of eCO2 on SOC stocks is best explained by a negative relationship with plant biomass: when plant biomass is strongly stimulated by eCO2, SOC storage declines; conversely, when biomass is weakly stimulated, SOC storage increases. This trade-off appears to be related to plant nutrient acquisition, in which plants increase their biomass by mining the soil for nutrients, which decreases SOC storage. We found that, overall, SOC stocks increase with eCO2 in grasslands (8 ± 2 per cent) but not in forests (0 ± 2 per cent), even though plant biomass in grasslands increase less (9 ± 3 per cent) than in forests (23 ± 2 per cent). Ecosystem models do not reproduce this trade-off, which implies that projections of SOC may need to be revised.

Author Correction: Climatic controls of decomposition drive the global biogeography of forest-tree symbioses.

In this Letter, a middle initial and additional affiliation have been added for author G. J. Nabuurs; two statements have been added to the Supplementary Acknowledgements; and a citation to the French National Institute has been added to the Methods; see accompanying Author Correction for further details.

Climatic controls of decomposition drive the global biogeography of forest-tree symbioses.

The identity of the dominant root-associated microbial symbionts in a forest determines the ability of trees to access limiting nutrients from atmospheric or soil pools1,2, sequester carbon3,4 and withstand the effects of climate change5,6. Characterizing the global distribution of these symbioses and identifying the factors that control this distribution are thus integral to understanding the present and future functioning of forest ecosystems. Here we generate a spatially explicit global map of the symbiotic status of forests, using a database of over 1.1 million forest inventory plots that collectively contain over 28,000 tree species. Our analyses indicate that climate variables-in particular, climatically controlled variation in the rate of decomposition-are the primary drivers of the global distribution of major symbioses. We estimate that ectomycorrhizal trees, which represent only 2% of all plant species7, constitute approximately 60% of tree stems on Earth. Ectomycorrhizal symbiosis dominates forests in which seasonally cold and dry climates inhibit decomposition, and is the predominant form of symbiosis at high latitudes and elevation. By contrast, arbuscular mycorrhizal trees dominate in aseasonal, warm tropical forests, and occur with ectomycorrhizal trees in temperate biomes in which seasonally warm-and-wet climates enhance decomposition. Continental transitions between forests dominated by ectomycorrhizal or arbuscular mycorrhizal trees occur relatively abruptly along climate-driven decomposition gradients; these transitions are probably caused by positive feedback effects between plants and microorganisms. Symbiotic nitrogen fixers-which are insensitive to climatic controls on decomposition (compared with mycorrhizal fungi)-are most abundant in arid biomes with alkaline soils and high maximum temperatures. The climatically driven global symbiosis gradient that we document provides a spatially explicit quantitative understanding of microbial symbioses at the global scale, and demonstrates the critical role of microbial mutualisms in shaping the distribution of plant species.

Quantifying global soil carbon losses in response to warming.

The majority of the Earth's terrestrial carbon is stored in the soil. If anthropogenic warming stimulates the loss of this carbon to the atmosphere, it could drive further planetary warming. Despite evidence that warming enhances carbon fluxes to and from the soil, the net global balance between these responses remains uncertain. Here we present a comprehensive analysis of warming-induced changes in soil carbon stocks by assembling data from 49 field experiments located across North America, Europe and Asia. We find that the effects of warming are contingent on the size of the initial soil carbon stock, with considerable losses occurring in high-latitude areas. By extrapolating this empirical relationship to the global scale, we provide estimates of soil carbon sensitivity to warming that may help to constrain Earth system model projections. Our empirical relationship suggests that global soil carbon stocks in the upper soil horizons will fall by 30 ± 30 petagrams of carbon to 203 ± 161 petagrams of carbon under one degree of warming, depending on the rate at which the effects of warming are realized. Under the conservative assumption that the response of soil carbon to warming occurs within a year, a business-as-usual climate scenario would drive the loss of 55 ± 50 petagrams of carbon from the upper soil horizons by 2050. This value is around 12-17 per cent of the expected anthropogenic emissions over this period. Despite the considerable uncertainty in our estimates, the direction of the global soil carbon response is consistent across all scenarios. This provides strong empirical support for the idea that rising temperatures will stimulate the net loss of soil carbon to the atmosphere, driving a positive land carbon-climate feedback that could accelerate climate change.

Hydraulic trade-offs and space filling enable better predictions of vascular structure and function in plants.

Plant vascular networks are central to botanical form, function, and diversity. Here, we develop a theory for plant network scaling that is based on optimal space filling by the vascular system along with trade-offs between hydraulic safety and efficiency. Including these evolutionary drivers leads to predictions for sap flow, the taper of the radii of xylem conduits from trunk to terminal twig, and how the frequency of xylem conduits varies with conduit radius. To test our predictions, we use comprehensive empirical measurements of maple, oak, and pine trees and complementary literature data that we obtained for a wide range of tree species. This robust intra- and interspecific assessment of our botanical network model indicates that the central tendency of observed scaling properties supports our predictions much better than the West, Brown, and Enquist (WBE) or pipe models. Consequently, our model is a more accurate description of vascular architecture than what is given by existing network models and should be used as a baseline to understand and to predict the scaling of individual plants to whole forests. In addition, our model is flexible enough to allow the quantification of species variation around rules for network design. These results suggest that the evolutionary drivers that we propose have been fundamental in determining how physiological processes scale within and across plant species.

Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species.

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (N(mass)) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (N(area)). In evergreen conifers, LMA and N(area) were less powerful predictors of area-based photosynthetic rate (A (max(area))) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, A (max(mass)) was related to N(mass) but not to LMA, whereas LMA became a significant codeterminant of A (max(mass)) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.

New tree-ring data from Canadian boreal and hemi-boreal forests provide insight for improving the climate sensitivity of terrestrial biosphere models.

Understanding boreal/hemi-boreal forest growth sensitivity to seasonal variations in temperature and water availability provides important basis for projecting the potential impacts of climate change on the productivity of these ecosystems. Our best available information currently comes from a limited number of field experiments and terrestrial biosphere model (TBM) simulations of varying predictive accuracy. Here, we assessed the sensitivity of annual boreal/hemi-boreal forest growth in Canada to yearly fluctuations in seasonal climate variables using a large tree-ring dataset and compared this to the climate sensitivity of annual net primary productivity (NPP) estimates obtained from fourteen TBMs. We found that boreal/hemi-boreal forest growth sensitivity to fluctuations in seasonal temperature and precipitation variables changed along a southwestern to northeastern gradient, with growth limited almost entirely by temperature in the northeast and west and by water availability in the southwest. We also found a lag in growth climate sensitivity, with growth largely determined by the climate during the summer prior to ring formation. Analyses of NPP sensitivity to the same climate variables produced a similar southwest to northeast gradient in growth climate sensitivity for NPP estimates from all but three TBMs. However, analyses of growth from tree-ring data and analyses of NPP from TBMs produced contrasting evidence concerning the key climate variables limiting growth. While analyses of NPP primarily indicated a positive relationship between growth and seasonal temperature, tree-ring analyses indicated negative growth relationships to temperature. Also, the positive effect of precipitation on NPP derived from most TBMs was weaker than the positive effect of precipitation on tree-ring based growth: temperature had a more important limiting effect on NPP than tree-ring data indicated. These mismatches regarding the key climate variables limiting growth suggested that characterization of tree growth in TBMs might need revision, particularly regarding the effects of stomatal conductance and carbohydrate reserve dynamics.

Biogeography and variability of eleven mineral elements in plant leaves across gradients of climate, soil and plant functional type in China.

Understanding variation of plant nutrients is largely limited to nitrogen and to a lesser extent phosphorus. Here we analyse patterns of variation in 11 elements (nitrogen/phosphorus/potassium/calcium/magnesium/sulphur/silicon/iron/sodium/manganese/aluminium) in leaves of 1900 plant species across China. The concentrations of these elements show significant latitudinal and longitudinal trends, driven by significant influences of climate, soil and plant functional type. Precipitation explains more variation than temperature for all elements except phosphorus and aluminium, and the 11 elements differentiate in relation to climate, soil and functional type. Variability (assessed as the coefficient of variation) and environmental sensitivity (slope of responses to environmental gradients) are lowest for elements that are required in the highest concentrations, most abundant and most often limiting in nature (the Stability of Limiting Elements Hypothesis). Our findings can help initiate a more holistic approach to ecological plant nutrition and lay the groundwork for the eventual development of multiple element biogeochemical models.

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's 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 CO(2) 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 CO(2) 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.

Acclimation of respiratory temperature responses in northern and southern populations of Pinus banksiana.

Temperature acclimation of respiration may contribute to climatic adaptation and thus differ among populations from contrasting climates. Short-term temperature responses of foliar dark respiration were measured in 33-yr-old trees of jack pine (Pinus banksiana) in eight populations of wide-ranging origin (44-55 degrees N) grown in a common garden at 46.7 degrees N. It was tested whether seasonal adjustments in respiration and population differences in this regard resulted from changes in base respiration rate at 5 degrees C (R(5)) or Q(10) (temperature sensitivity) and covaried with nitrogen and soluble sugars. In all populations, acclimation was manifest primarily through shifts in R(5) rather than altered Q(10). R(5) was higher in cooler periods in late autumn and winter and lower in spring and summer, inversely tracking variation in ambient air temperature. Overall, R(5) covaried with sugars and not with nitrogen. Although acclimation was comparable among all populations, the observed seasonal ranges in R(5) and Q(10) were greater in populations originating from warmer than from colder sites. Population differences in respiratory traits appeared associated with autumnal cold hardening. Common patterns of respiratory temperature acclimation among biogeographically diverse populations provide a basis for predicting respiratory carbon fluxes in a wide-ranging species.

CO2, nitrogen, and diversity differentially affect seed production of prairie plants.

Plant species composition and diversity is often influenced by early life history stages; thus, global change could dramatically affect plant community structure by altering seed production. Unfortunately, plant reproductive responses to global change are rarely studied in field settings, making it difficult to assess this possibility. To address this issue, we quantified the effects of elevated CO2, nitrogen deposition, and declining diversity on inflorescence production and inflorescence mass of 11 perennial grassland species in central Minnesota, U.S.A. We analyzed these data to ask whether (1) global change differentially affects seed production of co-occurring species; (2) seed production responses to global change are similar for species within the same functional group (defined by ecophysiology and growth form); and (3) seed production responses to global change match productivity responses: We found that, on average, allocation to seed production decreased under elevated CO2, although individual species responses were rarely significant due to low power (CO2 treatment df = 2). The effects of nitrogen deposition on seed production were similar within functional groups: C4 grasses tended to increase while C3 grasses tended to decrease allocation to seed production. Responses to nitrogen deposition were negatively correlated to productivity responses, suggesting a trade-off. Allocation to seed production of some species responded to a diversity gradient, but responses were uncorrelated to productivity responses and not similar within functional groups. Presumably, species richness has complex effects on the biotic and abiotic variables that influence seed production. In total, our results suggest that seed production of co-occurring species will be altered by global change, which may affect plant communities in unpredictable ways. Although functional groups could be used to generalize seed production responses to nitrogen deposition in Minnesota prairies, we caution against relying on them for predictive purposes without a mechanistic understanding of how resource availability and biotic interactions affect seed production.

Ambient levels of ozone reduce net photosynthesis in tree and crop species.

Experiments were conducted to measure the photosynthetic response of three crop and four tree species to realistic concentrations of ozone and (for tree species only) simulated acidic rain. The ozone concentrations were representative of those found in clean ambient air, in mildly to moderately polluted air such as occurs in much of the United States during the summer, and in more heavily polluted air. However, the highest concentrations of ozone used were lower than those found regularly in the Los Angeles area. The mean pH of the simulated acid rain treatments ranged from more alkaline to much more acidic than the mean pH of precipitation in the United States. Exposure to any increase in ozone reduced net photosynthesis in all species tested. In contrast, acidic rain had no negative effect on photosynthesis in tree species, and no interaction between ozone and acidic rain was observed. Ozone-induced reductions in photosynthesis were related to declines in growth or yield. Species with higher stomatal conductances and thus higher potential for pollutant uptake exhibited greater negative responses to similar ozone treatments. Since exposure to ozone concentrations typical of levels of the pollutant observed in the eastern half of the United States reduced the rates of net photosynthesis of all species tested, reductions in net photosynthesis may be occurring over much of the eastern United States.

Diversity and productivity in a long-term grassland experiment.

Plant diversity and niche complementarity had progressively stronger effects on ecosystem functioning during a 7-year experiment, with 16-species plots attaining 2.7 times greater biomass than monocultures. Diversity effects were neither transients nor explained solely by a few productive or unviable species. Rather, many higher-diversity plots outperformed the best monoculture. These results help resolve debate over biodiversity and ecosystem functioning, show effects at higher than expected diversity levels, and demonstrate, for these ecosystems, that even the best-chosen monocultures cannot achieve greater productivity or carbon stores than higher-diversity sites.

Ectomycorrhizal fungal communities of oak savanna are distinct from forest communities.

Oak savanna is one of the most endangered ecosystems of North America, with less than 0.02% of its original area remaining. Here we test whether oak savanna supports a unique community of ectomycorrhizal fungi, a higher diversity of ectomycorrhizal fungi or a greater proportional abundance of ascomycete fungi compared with adjacent areas where the absence of fire has resulted in oak savanna conversion to oak forest. The overall fungal community was highly diverse and dominated by Cenococcum geophilum and other ascomycetes, Cortinarius, Russula, Lactarius and Thelephoraceae. Oak savanna mycorrhizal communities were distinct from oak forest communities both aboveground (sporocarp surveys) and belowground (RFLP identification of ectomycorrhizal root tips); however total diversity was not higher in oak savanna than oak forests and there was no evidence of a greater abundance of ascomycetes. Despite not having a higher local diversity than oak forests, the presence of a unique fungal community indicates that oak savanna plays an important role in maintaining regional ectomycorrhizal diversity.

Climate warming will reduce growth and survival of Scots pine except in the far north.

Tree growth and survival were assessed in 283 populations of Scots pine (Pinus sylvestris L.) originating from a broad geographic range and grown at 90 common-garden experimental sites across Europe, and in 101 populations grown at 14 sites in North America. Growth and survival were analysed in response to climatic transfer distance, the difference in mean annual temperature (MAT) between the site and the population origin. Differences among populations at each site, and across sites for regional groups of populations, were related to climate transfer distance, but in opposite ways in the northern vs. southern parts of the species range. Climate transfers equivalent to warming by 1-4 degrees C markedly increased the survival of populations in northern Europe (>or= 62 degrees N, < 2 degrees C MAT) and modestly increased height growth >or= 57 degrees N but decreased survival at < 62 degrees N and modestly decreased height growth at < 54 degrees N latitude in Europe. Thus, even modest climate warming will likely influence Scots pine survival and growth, but in distinct ways in different parts of the species range.

Leaf physiological versus morphological acclimation to high-light exposure at different stages of foliar development in oak.

We investigated light acclimation in seedlings of the temperate oak Quercus petraea (Matt.) Liebl. and the co-occurring sub-Mediterranean oak Quercus pyrenaica Willd. Seedlings were raised in a greenhouse for 1 year in either 70 (HL) or 5.3% (LL) of ambient irradiance of full sunlight, and, in the following year, subsets of the LL-grown seedlings were transferred to HL either before leaf flushing (LL-HLBF plants) or after full leaf expansion (LL-HLAF plants). Gas exchange, chlorophyll a fluorescence, nitrogen fractions in photosynthetic components and leaf anatomy were examined in leaves of all seedlings 5 months after plants were moved from LL to HL. Differences between species in the acclimation of LL-grown plants to HL were minor. For LL-grown plants in HL, area-based photosynthetic capacity, maximum rate of carboxylation, maximum rate of electron transport and the effective photochemical quantum yield of photosystem II were comparable to those for plants grown solely in HL. A rapid change in nitrogen distribution among photosynthetic components was observed in LL-HLAF plants, which had the highest photosynthetic nitrogen-use efficiency. Increases in mesophyll thickness and dry mass per unit area governed leaf acclimation in LL-HLBF plants, which tended to have less nitrogen in photosynthetic components and a lower assimilation potential per unit of leaf mass or nitrogen than LL-HLAF plants. The data indicate that the phenological state of seedlings modified the acclimatory response of leaf attributes to increased irradiance. Morphological adaptation of leaves of LL-HLBF plants enhanced photosynthetic capacity per unit leaf area, but not per unit leaf dry mass, whereas substantial redistribution of nitrogen among photosynthetic components in leaves of LL-HLAF plants enhanced both mass- and area-based photosynthetic capacity.

Effects of plant species richness on invasion dynamics, disease outbreaks, insect abundances and diversity.

Declining biodiversity represents one of the most dramatic and irreversible aspects of anthropogenic global change, yet the ecological implications of this change are poorly understood. Recent studies have shown that biodiversity loss of basal species, such as autotrophs or plants, affects fundamental ecosystem processes such as nutrient dynamics and autotrophic production. Ecological theory predicts that changes induced by the loss of biodiversity at the base of an ecosystem should impact the entire system. Here we show that experimental reductions in grassland plant richness increase ecosystem vulnerability to invasions by plant species, enhance the spread of plant fungal diseases, and alter the richness and structure of insect communities. These results suggest that the loss of basal species may have profound effects on the integrity and functioning of ecosystems.

Temperature and ontogeny mediate growth response to elevated CO2 in seedlings of five boreal tree species.

We tested the extent to which growth responses to elevated carbon dioxide (CO2 ) are temperature-dependent and change through early seedling ontogeny among boreal tree species of contrasting relative growth rates (rgr). Populus tremuloides Michx, Betula papyrifera Marsh, Larix laricina (Du Roi) K. Koch, Pinus banksiana Lamb., and Picea mariana (Mill.) B.S.P. were grown from seeds for 3 months in controlled-environment chambers at two CO2 concentrations (370 and 580 µmol mol-1 ) and five temperature regimes of 18/12, 21/15, 24/18, 27/21 and 30/24°C (light/dark). Growth increases in response to CO2 enrichment were minimal at the lowest temperature and maximal at 21/15°C for the three conifers and at 24/18°C or higher for the two broadleaved species, corresponding with differences in optimal temperatures for growth. In both CO2 treatments, rgr among species and temperatures correlated positively with leaf area ratio (lar) (r⩾0·90, P<0·0001). However, at a given lar, rgr was higher in elevated CO2 , owing to enhanced whole-plant net assimilation rate. On average in all species and temperatures at a common plant mass, CO2 enrichment increased rgr (9%) through higher whole-plant net assimilation rate (22%), despite declines in lar in high CO2 (11%). Reductions in lar are thus an important feedback mechanism reducing positive plant growth responses to CO2 . Proportional allocation of dry mass to roots did not vary between CO2 treatments. Early in the experiment, proportional increases in plant dry mass in elevated CO2 were larger in faster-growing Populus tremuloides and B. papyrifera than in the slower-growing conifers. However, growth increases in response to CO2 enrichment fell with time for broadleaved species and increased for the conifers. With increasing plant size over time, compensatory adjustments to CO2 enrichment in the factors that determine rgr, such as lar, were much larger in broadleaves than in conifers. Thus, the hypothesis that faster-growing species are more responsive to elevated CO2 was not supported, given contrasting patterns of growth response to CO2 with increasing plant size and age.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: I. In situ net photosynthesis, dark respiration and growth.

Hybrid poplar (Populus tristis Fisch. ×P. balsamifera L., cv. Tristis) and sugar maple (Acer saccharum Marsh.) seedlings were grown under contrasting light and ozone treatments to investigate the role of the light environment in their response to chronic ozone stress. In consecutive growth chamber experiments, cuttings of shade-intolerant poplar and 3-yr-old seedlings of shade-tolerant sugar maple were grown in pots for 6 and 10 wk, respectively, under shaded, low light irradiance (c. 2.5 mol m-2 d-1 PPFD or 7% of full sunlight) and six-fold greater irradiance (c. 16.6 mol m-2 d-1 PPFD or 45% of full sunlight) in combination with low (< 10 nl 1-1 ) and elevated levels of ozone (c. 99-115 nl 1-1 ). In unshaded poplar plants, ozone exposure reduced root dry mass by 33% at final harvest, while shaded plants had no such response. By comparison, sugar maple root dry mass was reduced by ozone in shaded plants by 10%, but was unaffected by ozone in unshaded plants. In poplar, leaf area: plant dry mass ratios were unaffected by ozone, whereas in sugar maple ozone-exposed plants had a 24% lower leaf area: plant dry mass ratio in the shaded treatment. In shade-grown sugar maple, ozone doubled dark respiration rates of leaves, but in unshaded seedlings ozone had no effect on respiration. In comparison, in poplar plants ozone exposure resulted in greater increases in dark respiration under unshaded than shaded conditions. In unshaded plants, ozone treatment resulted in lower in situ net photosynthesis in poplar, but not in sugar maple. Overall, shade-grown sugar maple appeared more sensitive to ozone stress than unshaded plants in terms of lower leaf area: plant dry mass ratio and root growth and higher leaf respiration. In poplar on the other hand, root growth, leaf respiration and photosynthesis were more affected by ozone in unshaded than in shaded plants. These findings suggest that shade-grown sugar maple and unshaded poplar may experience greater reductions in carbon gain and growth under elevated levels of ozone than plants under the opposite light conditions.

Light environment alters response to ozone stress in seedlings of Acer saccharum Marsh, and hybrid Populus L.: II. Diagnostic gas exchange and leaf chemistry.

Diagnostic gas exchange measurements and foliar chemical assays were conducted on hybrid poplar (Populus tristis Fisch. ×P. balsamifera L. cv. Tristis) and sugar maple (Acer saccharum Marsh.) seedlings grown under contrasting light and ozone treatments. Seedlings were grown in low irradiance (c. 2.5 mol m-2 d-1 ) and six-fold greater irradiance (c. 16.6 mol m-2 d-1 ) in combination with low (< 10 nl I -l ) and elevated (99-115 nl 1-1 ) ozone. Analysis of light response curves showed ozone-induced reductions in photosynthetic capacity and quantum yield for unshaded poplar and shaded sugar maple, but not the contrasting light treatments. Photosynthesis at saturating CO2 concentrations was decreased in the elevated ozone treatment in both the unshaded and shaded poplar and in shaded sugar maple. Poplar had significant reductions in chlorophyll concentration due to ozone exposure in both unshaded and shaded treatments. Older leaves of unshaded poplar plants had significantly greater reductions in chlorophyll levels due to ozone than older leaves of shaded plants. In maple, only shade-grown leaves had significant decreases in chlorophyll concentration due to ozone exposure. The diagnostic gas exchange measurements in conjunction with chlorophyll measurements indicate that in hybrid poplar, unshaded leaves may be more sensitive to ozone than shade leaves, while in sugar maple, shade leaves are more sensitive to ozone. For hybrid poplar a decrease in photosynthetic capacity, quantum yield and chlorophyll concentration in the unshaded, moderately high light environment due to elevated ozone is consistent with prior studies. The results indicating that sugar maple seedlings may be more detrimentally affected by elevated ozone in the lower light environment may have serious implications for this and other shade-adapted species with respect to their performance in an understorey environment.