Responses of N fluxes and pools to elevated atmospheric CO2 in model forest ecosystems with acidic and calcareous soils

The objectives of this study were to estimate how soil type, elevated N deposition (0.7 vs. 7 g N m^–2y^–1) and tree species influence the potential effects of elevated CO₂ (370 vs. 570 mol CO₂ mol^–1) on N pools and fluxes in forest soils. Model spruce-beech forest ecosystems were established on a nutrient-rich calcareous sand and on a nutrient-poor acidic loam in large open-top chambers. In the fourth year of treatment, we measured N concentrations in the soil solution at different depths, estimated N accumulation by ion exchange resin (IER) bags, and quantified N export in drainage water, denitrification, and net N uptake by trees. Under elevated CO₂, concentrations of N in the soil solution were significantly reduced. In the nutrient-rich calcareous sand, CO₂ enrichment decreased N concentrations in the soil solution at all depths (–45 to –100%). In the nutrient-poor acidic loam, the negative CO₂ effect was restricted to the uppermost 5 cm of the soil. Increasing the N deposition stimulated the negative impact of CO₂ enrichment on soil solution N in the acidic loam at 5 cm depth from –20% at low N inputs to –70% at high N inputs. In the nutrient-rich calcareous sand, N additions did not influence the CO₂ effect on soil solution N. Accumulation of N by IER bags, which were installed under individual trees, was decreased at high CO₂ levels under spruce in both soil types. Under beech, this decrease occurred only in the calcareous sand. N accumulation by IER bags was negatively correlated with current-years foliage biomass, suggesting that the reduction of soil N availability indices was related to a CO₂-induced growth enhancement. However, the net N uptake by trees was not significantly increased by elevated CO₂. Thus, we suppose that the reduced N concentrations in the soil solution at elevated CO₂ concentrations were rather caused by an increased N immobilisation in the soil. Denitrification was not influenced by atmospheric CO₂ concentrations. CO₂ enrichment decreased nitrate leaching in drainage by 65%, which suggests that rising atmospheric CO₂ potentially increases the N retention capacity of forest ecosystems.     

Mechanical properties of spruce and beech wood grown in elevated CO2

Biomechanical responses of stems of 6- to 7-year-old spruce [Picea abies (L.) Karst.] and beech (Fagus sylvatica L) trees were studied after 4 years of growth in elevated atmospheric CO₂ in combination with a nitrogen treatment and on two different soil types. At the end of the treatment, stems were harvested and tested in fresh and air-dried status. Bending characteristics of juvenile wood (modulus of elasticity, termed rigidity) were determined by bending tests. Fracture characteristics (termed toughness) were determined by stroke-pendulum tests. From the base disk of each stem densitometric data were obtained. In spruce, wood produced under elevated CO₂ was tougher on both soil types; enhanced N deposition made wood less rigid only on acidic soils. In contrast, beech wood samples showed no significant reaction to CO₂ but were significantly tougher under high nitrogen depositions on acidic soil. Effects on wood density of both CO₂ and N treatments were not significant, but wood density was higher on acidic soil and so were rigidity and toughness (soil effect). Different genotypes of spruce and beech reacted significantly differently to the treatments. Some genotypes reacted strongly to CO₂ or N, whereas others did not react or showed interactions between CO₂ and N. This underlines the importance of genetic diversity in tree communities.     

Four-year growth dynamics of beech-spruce model ecosystems under CO2 enrichment on two different forest soils

To elucidate how atmospheric CO₂ enrichment, enhanced nutrient supply and soil quality interact to affect regrowth of temperate forests, young Fagus sylvatica and Picea abies trees were grown together in large model ecosystems. Identical communities were established on a nutrient-poor acidic and on a more fertile calcareous soil and tree growth, leaf area index, fine root density and soil respiration monitored over four complete growing seasons. Biomass responses to CO₂ enrichment and enhanced N supply at the end of the experiment reflected compound interest effects of growth stimulation during the first two to three seasons rather than persistent stimulation over the whole duration of the experiment. Whereas biomass of Picea was enhanced in elevated CO₂ on both soils, Fagus responded negatively to CO₂ on acidic but positively on calcareous soil. Biomass of both species profited from enhanced N supply on the poor acidic soil only. Leaf area index on both soils was greater in high N supply as a consequence of a stimulation early in the experiment, but was unaffected by CO₂ enrichment. Fine root density on acidic soil was increased in high N supply, but this did not stimulate soil respiration rate. In contrast, elevated CO₂ stimulated both fine root density and soil CO₂ efflux on calcareous soil, especially towards the end of the experiment. Our experiment suggests that future species dominance in beech-spruce forests is likely to change in response to CO₂ enrichment, but this response is subject to complex interactions with environmental factors other than CO₂, particularly soil type.     

The input and fate of new C in two forest soils under elevated CO2

The aim of this study was to estimate (i) the influence of different soil types on the net input of new C into soils under CO₂ enrichment and (ii) the stability and fate of these new C inputs in soils. We exposed young beech–spruce model ecosystems on an acidic loam and calcareous sand for 4 years to elevated CO₂. The added CO₂ was depleted in ¹³C, allowing to trace new C inputs in the plant–soil system. We measured CO₂‐derived new C in soil C pools fractionated into particle sizes and monitored respiration as well as leaching of this new C during incubation for 1 year. Soil type played a crucial role in the partitioning of C. The net input of new C into soils under elevated CO₂ was about 75% greater in the acidic loam than in the calcareous sand, despite a 100% and a 45% greater above‐ and below‐ground biomass on the calcareous sand. This was most likely caused by a higher turnover of C in the calcareous sand as indicated by 30% higher losses of new C from the calcareous sand than from the acidic loam during incubation. Therefore, soil properties determining stabilization of soil C were apparently more important for the accumulation of C in soils than tree productivity. Soil fractionation revealed that about 60% of the CO₂‐derived new soil C was incorporated into sand fractions. Low natural ¹³C abundance and wide C/N ratios show that sand fractions comprise little decomposed organic matter. Consistently, incubation indicated that new soil C was preferentially respired as CO₂. During the first month, evolved CO₂ consisted to 40–55% of new C, whereas the fraction of new C in bulk soil C was 15–23% only. Leaching of DOC accounted for 8–23% of the total losses of new soil C. The overall effects of CO₂ enrichment on soil C were small in both soils, although tree growth increased significantly on the calcareous sand. Our results suggest that the potential of soils for C sequestration is limited, because only a small fraction of new C inputs into soils will become long‐term soil C.     

Forest carbon balance under elevated CO2

Free-air CO₂ enrichment (FACE) technology was used to expose a loblolly pine (Pinus taeda L.) forest to elevated atmospheric CO₂ (ambient + 200 µl l^-1). After 4 years, basal area of pine trees was 9.2% larger in elevated than in ambient CO₂ plots. During the first 3 years the growth rate of pine was stimulated by ~26%. In the fourth year this stimulation declined to 23%. The average net ecosystem production (NEP) in the ambient plots was 428 gC m^-2 year^-1, indicating that the forest was a net sink for atmospheric CO₂. Elevated atmospheric CO₂ stimulated NEP by 41%. This increase was primarily an increase in plant biomass increment (57%), and secondarily increased accumulation of carbon in the forest floor (35%) and fine root increment (8%). Net primary production (NPP) was stimulated by 27%, driven primarily by increases in the growth rate of the pines. Total heterotrophic respiration (R_h) increased by 165%, but total autotrophic respiration (R_a) was unaffected. Gross primary production was increased by 18%. The largest uncertainties in the carbon budget remain in separating belowground heterotrophic (soil microbes) and autotrophic (root) respiration. If applied to temperate forests globally, the increase in NEP that we measured would fix less than 10% of the anthropogenic CO₂ projected to be released into the atmosphere in the year 2050. This may represent an upper limit because rising global temperatures, land disturbance, and heterotrophic decomposition of woody tissues will ultimately cause an increased flux of carbon back to the atmosphere.     

Effects of elevated CO<Subscript>2</Subscript> on foliar chemistry of saplings of nine species of tropical tree

This study examined the effects of elevated CO₂ on secondary metabolites for saplings of tropical trees. In the first experiment, nine species of trees were grown in the ground in open-top chambers in central Panama at ambient and elevated CO₂ (about twice ambient). On average, leaf phenolic contents were 48% higher under elevated CO₂. Biomass accumulation was not affected by CO₂, but starch, total non-structural carbohydrates and C/N ratios all increased. In a second experiment with Ficus, an early successional species, and Virola, a late successional species, treatments were enriched for both CO₂ and nutrients. For both species, nutrient fertilization increased plant growth and decreased leaf carbohydrates, C/N ratios and phenolic contents, as predicted by the carbon/nutrient balance hypothesis. Changes in leaf C/N levels were correlated with changes in phenolic contents for Virola (r=0.95, P<0.05), but not for Ficus. Thus, elevated CO₂, particularly under conditions of low soil fertility, significantly increased phenolic content as well as the C/N ratio of leaves. The magnitude of the changes is sufficient to negatively affect herbivore growth, survival and fecundity, which should have impacts on plant/herbivore interactions.     

Microbial community composition and function beneath temperate trees exposed to elevated atmospheric carbon dioxide and ozone

We hypothesized that changes in plant growth resulting from atmospheric CO₂ and O₃ enrichment would alter the flow of C through soil food webs and that this effect would vary with tree species. To test this idea, we traced the course of C through the soil microbial community using soils from the free-air CO₂ and O₃ enrichment site in Rhinelander, Wisconsin. We added either ¹³C-labeled cellobiose or ¹³C-labeled N-acetylglucosamine to soils collected beneath ecologically distinct temperate trees exposed for 3 years to factorial CO₂ (ambient and 200 µl l^-1 above ambient) and O₃ (ambient and 20 µl l^-1 above ambient) treatments. For both labeled substrates, recovery of ¹³C in microbial respiration increased beneath plants grown under elevated CO₂ by 29% compared to ambient; elevated O₃ eliminated this effect. Production of ¹³C-CO₂ from soils beneath aspen (Populus tremuloides Michx.) and aspen-birch (Betula papyrifera Marsh.) was greater than that beneath aspen-maple (Acer saccharum Marsh.). Phospholipid fatty acid analyses (¹³C-PLFAs) indicated that the microbial community beneath plants exposed to elevated CO₂ metabolized more ¹³C-cellobiose, compared to the microbial community beneath plants exposed to the ambient condition. Recovery of ¹³C in PLFAs was an order of magnitude greater for N-acetylglucosamine-amended soil compared to cellobiose-amended soil, indicating that substrate type influenced microbial metabolism and soil C cycling. We found that elevated CO₂ increased fungal activity and microbial metabolism of cellobiose, and that microbial processes under early-successional aspen and birch species were more strongly affected by CO₂ and O₃ enrichment than those under late-successional maple.     

A link between plant diversity, elevated CO2 and soil nitrate

Interactive effects of reductions in plant species diversity and increases in atmospheric CO₂ were investigated in a long-term study in nutrient-poor calcareous grassland. Throughout the experiment, soil nitrate was persistently increased at low plant species diversity, and CO₂ enrichment reduced soil [NO₃^-] at all levels of plant species diversity. In our study, soil [NO₃^-] was unrelated to root length density, microbial biomass N, community legume contents, and experimental plant communities differed only little in total N pools. However, potential nitrification revealed exactly the same treatment effects as soil [NO₃^-], providing circumstantial evidence that nitrification rates drove the observed changes in [NO₃^-]. One possible explanation for plant diversity effects on nitrification lies in spatial and temporal interspecific differences in plant N uptake, which would more often allow accumulation of NH₄⁺ in part of the soil profile at low diversity than in more species-rich plant communities. Consequently, nitrification rates and soil [NO₃^-] would increase. Elevated CO₂ increased soil water contents, which may have improved NO₃^- diffusion to the root surface thereby reducing soil [NO₃^-]. Higher soil moisture at elevated CO₂ might also reduce nitrification rates due to less aerobic conditions. The accordance of the diversity effect on soil [NO₃^-] with previous experiments suggests that increased soil [NO₃^-] at low species diversity is a fairly general phenomenon, although the mechanisms causing high [NO₃^-] may vary. In contrast, experimental evidence for effects of CO₂ enrichment on soil [NO₃^-] is ambiguous, and the antagonistic interaction of plant species reductions and elevated CO₂ we have observed is thus probably less universal.     

Downward adjustment of carbon fluxes at the biochemical, leaf, and ecosystem scale in beech-spruce model communities exposed to long-term atmospheric CO2 enrichment

Young beech (Fagus sylvatica) and spruce (Picea abies) trees from different provenances or genotypes were grown in competition in large model ecosystems and were exposed to two concentrations of atmospheric CO₂ (370 vs 570 μmol mol^?1), two levels of wet nitrogen deposition (7 vs 70 kg N ha^?1 yr^?1), and two native forest soils (acidic vs calcareous) for four years in open‐top chambers. The 2×2×2 factorial experimental design was fully replicated (n=4) with each CO₂×N combination applied to each soil type. Exposure to atmospheric CO₂ enrichment stimulated daytime net ecosystem CO₂ flux (NEC) as measured during sunny days in the middle of the third growing season. Nevertheless, we observed substantial down‐regulation of NEC, with larger adjustments on acidic than on calcareous soil. NEC adjustment was associated with slightly reduced leaf area index (LAI) on the acidic soil (no response on calcareous soil), enhanced soil CO₂ efflux from both substrate types, and, most importantly, with down‐regulation of CO₂ uptake at the leaf scale. Downward adjustment of light‐saturated single‐leaf photosynthesis (A) and of Rubisco was more pronounced in beech than in spruce and these species‐specific differences increased over time. By year four, A adjustment (except in one specific treatment combination in each species) had become complete in beech but had disappeared in spruce. At no time did we observe a genotype or provenance effect on the downward adjustment of carbon fluxes, and nitrogen deposition rate generally had little effect as well. Overall, our results suggest that tree species and soil quality will have profound effects on ecosystem CO₂ fluxes under continued atmospheric CO₂ enrichment.     

The effects of elevated CO2 on seed production and seedling recruitment in a sheep-grazed pasture

Seed production and seedling recruitment were measured over 2 years under ambient (360 ppm) and elevated (475 ppm) atmospheric CO₂ in a free air carbon dioxide enrichment (FACE) experiment, carried out in a sheep-grazed pasture on dry, sandy soil in New Zealand. In both years elevated CO₂ led to more dispersed seeds of the grasses Anthoxanthum odoratum, Lolium perenne and Poa pratensis, the legumes Trifolium repens and T. subterraneum and the herbs Hypochaeris radicata and Leontodon saxatilis. The increased seed dispersal in A. odoratum, H. radicata, Leontodon saxatilis and T. repens reflected both more inflorescences per unit area and more seeds per inflorescence under elevated CO₂. The increased seed dispersal in Lolium perenne, P. pratensis and T. subterraneum was due solely to more inflorescences per unit area. The number of seedlings that emerged and survived to at least 7 months of age was increased by elevated CO₂ for H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in both years and for A. odoratum and Lolium perenne in the first year. For species where increased seedling recruitment was noted, there was a significant positive correlation between seed production in summer and seedling emergence in the following autumn and winter, and sowing 200 extra seeds per species m^-2 resulted in more seedlings compared to unsown controls. Elevated CO₂ did not affect seedling survival in any species. There was no measurable effect of elevated CO₂ on canopy and soil surface conditions or soil moisture at the time of seedling emergence. The results suggest the dominant effect of elevated CO₂ on seedling recruitment in this pasture was an indirect one, reflecting effects on the number of seeds produced. The biomass of H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in the above-ground vegetation was greater under elevated than ambient CO₂. However, the size of individual seedlings and mature plants of these four species was unaffected by elevated CO₂. The results indicate an important way elevated CO₂ influenced plant species composition in this pasture was through changes in the pattern of seedling recruitment.     

Photosynthetic characteristics and growth responses of dwarf apple (Malus domestica Borkh. cv. Fuji) saplings after 3 years of exposure to elevated atmospheric carbon dioxide concentration and temperature

Growth and photosynthetic responses of dwarf apple saplings (Malus domestica Borkh. cv. Fuji) acclimated to 3 years of exposure to contrasting atmospheric CO₂ concentrations (360 and 650 µmol mol^-1) in combination with current ambient or elevated (ambient +5°C) temperature patterns were determined. Four 1-year-old apple saplings grafted onto M.9 rootstocks were each enclosed in late fall 1997 in a controlled environment unit in nutrient-optimal soil. Soil moisture regimes were automatically controlled by drip irrigation scheduled at 50 kPa of soil moisture tension. For the elevated CO₂ concentration alone, overall tree growth was suppressed. However, tree growth was slightly enhanced when warmer temperatures were combined with the elevated CO₂ concentration. Neither temperature nor CO₂ concentration affected leaf chlorophyll content and stomatal density. The elevated CO₂ concentration decreased mean leaf area, but increased starch accumulation, thus resulting in a higher specific dry mass of leaves. An elevated temperature reduced starch accumulation. Light-saturated rates of leaf photosynthesis were suppressed due to the elevated CO₂ concentration, but this effect was removed or enhanced with warmer temperatures. The elevated CO₂ concentration increased the optimum temperature for photosynthesis by ca. 4°C, while the warmer temperature did not. The results of this study suggested that the long-term adaptation of apple saplings to growth at an elevated CO₂ concentration may be associated with a potential for increased growth and productivity, if a doubling of the CO₂ concentration also leads to elevated temperatures.     

Nitrogen mineralization and phenol accumulation along a fire chronosequence in northern Sweden

Scots pine (Pinus sylvestris L.) forests of northern Sweden are often considered to be N limited. This limitation may have been exacerbated by the elimination of wildfire as a natural disturbance factor in these boreal forests. Phenolic inhibition of N mineralization and nitrification (due to litter and exudates of ericaceous shrubs) has been proposed as a mechanism for N limitation of these forests, but this hypothesis remains largely untested. N mineralization rates, nitrification rates, and sorption of free phenolic compounds were assessed along a fire-induced chronosequence in northern Sweden. A total of 34 forest stands varying in age since the last fire were identified and characterized. Overstorey and understorey vegetative composition and depth of humus were analysed in replicated plots at all 34 sites. Eight of the forest stands aged 3-352 years since the last fire were selected for intensive investigation in which ten replicate ionic resin capsules (used to assess net N mineralization and nitrification) and non-ionic carbonaceous resin capsules (used to assess free phenolic compounds) were installed at the interface of humus and mineral soil. A highly significant correlation was observed between site age and net sorption of inorganic N to resin capsules. Net accumulation of NH₄⁺ and NO₃^- on resin capsules followed a linear decrease (R²=0.61, P<0.01) with time perhaps as a result of increased N immobilization with successional C loading. NO₃^- sorption to resin capsules followed a logarithmic decrease (R²=0.80, P<0.01) that may be related to a logarithmic increase in dwarf shrub cover and decreased soil charcoal sorption potential along this chronosequence. A replicated field study was conducted at one of the late successional field sites to assess the influence of charcoal and an added labile N source on N turnover. Three rates of charcoal (0, 100, and 1,000 g M^-2) and two rates of glycine (0 and 50 g N as glycine M^-2) were applied in a factorial design to microplots in a randomized complete block pattern. Net ammonification (as assessed by NH₄⁺ sorption to resins) was readily increased by the addition of a labile N source, but this increase in NH₄⁺ did not stimulate nitrification. Nitrification was stimulated slightly by the addition of charcoal resulting in similar levels of resin-sorbed NO₃^- as those found in early successional sites. Resin-sorbed polyphenol concentrations were decreased with charcoal amendments, but were actually increased with N amendments (likely due to decomposition of polyphenols). Net N mineralization appears to be limited by rapid NH₄⁺ immobilization whereas nitrification is limited by the lack of an appropriate environment or by the presence of inhibitory compounds in late successional forests of northern Sweden.     

Differential impacts of long-term (CO<Subscript>2</Subscript>) and O<Subscript>3</Subscript> exposure on growth of northern conifer and deciduous tree species

The long-term effects of elevated CO₂ and CO₂+O₃ concentrations on the growth allocation in northern provenances of Norway spruce [Picea abies (L.) Karst.], Scots pine [Pinus sylvestris (L.)] and pubescent birch clones (Betula pubescens Ehrh.) were examined in open-top chambers after a 4-year-long experiment. The total biomass responses of the tree seedlings to increased CO₂ and CO₂+O₃ concentrations were not statistically significant and varied between the provenances and species. The seedlings of northern origin were the least sensitive in their response to treatments. The total biomass of the Norway spruce seedlings slightly decreased in response to CO₂ in three provenances. Scots pine from the local provenance had a slight biomass increase after elevated CO₂+O₃ treatment. The slower-growing birch clone seemed to benefit from elevated CO₂, whereas in the faster-growing clone, reductions in biomass accumulation were seen. The combined CO₂+O₃ treatment reduced the positive effects of elevated CO₂, especially in the slower-growing birches. Observations of significant effects were limited to a few parameters. Carbon dioxide treatment decreased needle dry weight of Norway spruce in one northern provenance. The needle and wood dry weight increased (CO₂ + O₃) in local Scots pine. Significant birch response was limited to increased fine root density (O₃ + CO₂) in the inland clone. The diverse effects of elevated CO₂ and CO₂ +O₃ on seedling growth and biomass provide evidence that exposure of northern trees to the enhanced variable CO₂ and O₃ concentrations of the future will have varied effects on the growth of these species. The direction and magnitude of those effects will differ depending on species and origins.     

Feeding rates of the woodlouse Armadillidium vulgare on herb litters produced at two levels of atmospheric CO2

The consumption and assimilation rates of the woodlouse Armadillidium vulgare were measured on leaf litters from five herb species grown and naturally senesced at 350 and 700 µl l^-1 CO₂. Each type of litter was tested separately after 12, 30 and 45 days of decomposition at 18°C. The effects of elevated CO₂ differed depending on the plant species. In Medicago minima (Fabaceae), the CO₂ treatment had no significant effect on consumption and assimilation. In Tyrimnus leucographus (Asteraceae), the CO₂ treatment had no significant effect on consumption, but the elevated CO₂ litter was assimilated at a lower rate than the ambient CO₂ litter after 30 days of decomposition. In the three other species, Galactites tomentosa (Asteraceae), Trifolium angustifolium (Fabaceae) and Lolium rigidum (Poaceae), the elevated CO₂ litter was consumed and/or assimilated at a higher rate than the ambient CO₂ litter. Examination of the nitrogen contents in these three species of litter did not support the hypothesis of compensatory feeding, i.e. an increase in woodlouse consumption to compensate for low nitrogen content of the food. Rather, the results suggest that in herbs that were unpalatable at the start of the experiment (Galactites, Trifolium and Lolium), more of the the litter produced at 700 µl l^-1 CO₂ was consumed than of that produced at 350 µl l^-1 because inhibitory factors were eliminated faster during decomposition.     

Physiology and Growth of Wheat Across a Subambient Carbon Dioxide Gradient

Two cultivars of wheat (Triticum aestivum L.), 'Yaqui 54' and'Seri M82', were grown along a gradient of daytime carbon dioxideconcentrations ([CO₂]) from near 350-200 µmol CO₂ mol^-1air in a 38 m long controlled environment chamber. Carbon dioxidefluxes and evapotranspiration were measured for stands (plantsand soil) in five consecutive 7·6-m lengths of the chamberto determined potential effects of the glacial/interglacialincrease in atmospheric [CO₂] on C₃ plants. Growth rates andleaf areas of individual plants and net assimilation per unitleaf area and daily (24-h) net CO₂ accumulation of wheat standsrose with increasing [CO₂]. Daytime net assimilation (P_D, mmolCO₂ m^-2 soil surface area) and water use efficiency of wheatstands increased and the daily total of photosynthetic photonflux density required by stands for positive CO₂ accumulation(light compensation point) declined at higher [CO₂]. Nighttimerespiration (R_N, mmol CO₂ m^-2 soil surface) of wheat, measuredat 369-397 µmol mol^-1 CO₂, apparently was not alteredby growth at different daytime [CO₂], but R_N /P_D of stands declinedlinearly as daytime [CO₂] and P_D increased. The responses ofwheat to [CO₂], if representative of other C₃ species, suggestthat the 75-100% increase in [CO₂] since glaciation and the30% increase since 1800 reduced the minimum light and waterrequirements for growth and increased the productivity of C₃plants.Copyright 1993, 1999 Academic Press Atmospheric carbon dioxide, carbon accumulation, evapotranspiration, light compensation point, net assimilation, respiration, Triticum aestivum, water use efficiency, wheat     

Carbon isotope discrimination in photosynthetic bark

We developed and tested a theoretical model describing carbon isotope discrimination during photosynthesis in tree bark. Bark photosynthesis reduces losses of respired CO₂ from the underlying stem. As a consequence, the isotopic composition of source CO₂ and the CO₂ concentration around the chloroplasts are quite different from those of photosynthesizing leaves. We found three lines of evidence that bark photosynthesis discriminates against ¹³C. First, in bark of Populus tremuloides, the '¹³C of CO₂ efflux increased from -24.2‰ in darkness to -15.8‰ in the light. In Pinus monticola, the '¹³C of CO₂ efflux increased from -27.7‰ in darkness to -10.2‰ in the light. Observed increases in '¹³C were generally in good agreement with predictions from the theoretical model. Second, we found that '¹³C of dark-respired CO₂ decreased following 2-3 h of illumination (P<0.01 for Populus tremuloides, P<0.001 for Pinus monticola). These decreases suggest that refixed photosynthate rapidly mixes into the respiratory substrate pool. Third, a field experiment demonstrated that bark photosynthesis influenced whole-tissue '¹³C. Long-term light exclusion caused a localized increase in the '¹³C of whole bark and current-year wood in branches of P. monticola (P<0.001 and P<0.0001, respectively). Thus bark photosynthesis was shown to discriminate against ¹³C and create a pool of photosynthate isotopically lighter than the dark respiratory pool in all three experiments. Failure to account for discrimination during bark photosynthesis could interfere with interpretation of the '¹³C in woody tissues or in woody-tissue respiration.     

Consequences of elevated CO2, augmented nitrogen-deposition and soil type on the soluble nitrogen and sulphur in the phloem of beech (Fagus sylvatica) and spruce (Picea abies) in a competitive situation

Mixed spruce-beech plantations grown in large open-top chambers (OTC) were used to study consequences of elevated CO2, nitrogen-deposition and soil type on plant internal nitrogen and sulphur cycling of juvenile beech (Fagus sylvatica L.) and spruce (Picea abies Karst.) in a competitive situation. Processes of re-cycling as a consequence of protein turnover during leaf senescence in autumn were of further interest. For this purpose, phloem sap was collected in September 1998 and analysed for the composition and concentrations of organic and inorganic nitrogen and sulphur compounds. The phloem exudate of spruce showed higher total soluble non-protein nitrogen (TSNN) concentration on calcareous soil than on acidic soil, independent of the treatment. N-fertilization increased the N-concentration of phloem exudate significantly on both soil types, mainly by an increase of Arg and Gln concentrations. Elevated CO2 slightly increased TSNN on calcareous, but not on acidic soil. The combination of elevated CO2 and augmented N-deposition induced a further increase of TSNN on calcareous soil, but caused a lower N-effect on TSNN on acidic soil. Arg, the main TSNN component in phloem exudate, mediated this effect. Since Arg is considered to be a major nitrogen storage compound, it is concluded that in autumn elevated CO2 and augmented N-deposition, influence storage of N rather than N-supply of spruce. An effect of elevated CO2 and augmented N-deposition on GSH and sulphate concentrations in phloem exudate of spruce was not observed on acidic soil. On calcareous soil augmented N-deposition enhanced, elevated CO2 decreased phloem exudate GSH contents. In combination, elevated CO2 compensated the positive effect of N-deposition. The effects of elevated CO2 and augmented N-deposition on phloem sap N- and S-contents described above were not observed for beech trees. Apparently, elevated CO2 and augmented N-deposition did not affect plants internal S and N cycling of beech grown in spruce-beech plantations.     

Variations of stomatal density and carbon isotope values of<Emphasis Type="Italic"> Picea crassifolia</Emphasis> at different altitudes in the Qilian Mountains

The stomatal characteristics, length and dry weight as well as stable carbon isotope composition (i¹³C) of needles and tree rings of Qinghai spruce (Picea crassifolia) at different altitudes in the Qilian mountains were investigated. The results showed that stomatal density, distribution pattern on epidermis, and length and dry weight of needles all increased with altitude below 3,000 m. In contrast, these parameters all decreased with increasing altitude above 3,000 m. Furthermore an assay of tree rings showed that near 3,000 m in altitude was the optimum zone for growth and development of Qinghai spruce. Although atmospheric CO₂ concentration influences stomatal density, the effects of many environmental factors, such as temperature, light and rainfall, could obscure the effects of changes in atmospheric CO₂ concentration on stomatal density at altitudes higher than the optimum for growth. The correlation of stomatal density with atmospheric CO₂ concentration and i¹³C of Qinghai spruce needles was significant below 3,000 m, but was insignificant above 3,000 m altitude. The i¹³C value of needles, however, was negatively correlated with atmospheric CO₂ concentrations. Therefore, the influence of altitude should be considered in research on plant physiological ecology and the relationship of stomatal density with i¹³C value or atmospheric CO₂ concentration.     

Environmental controls on carbon dioxide flux from black spruce coarse woody debris

Carbon dioxide flux from coarse woody debris (CWD) is an important source of CO₂ in forests with moderate to large amounts of CWD. A process-based understanding of environmental controls on CWD CO₂ flux (R_CWD) is needed to accurately model carbon exchange between forests and the atmosphere. The objectives of this study were to: (1) use a laboratory incubation factorial experiment to quantify the effect of temperature (T_CWD), water content (W_C), decay status, and their interactions on R_CWD for black spruce [Picea mariana (Mill.) BSP] CWD; (2) measure and model spatial and temporal dynamics in T_CWD for a boreal black spruce fire chronosequence; and (3) validate the R_CWD model with field measurements, and quantify potential errors in estimating annual R_CWD from this model on various time steps. The R_CWD was positively correlated to T_CWD (R²=0.37, P<0.001) and W_C (R²=0.18, P<0.001), and an empirical R_CWD polynomial model that included T_CWD and W_C interactions explained 74% of the observed variation of R_CWD. The R_CWD estimates from the R_CWD model excellently matched the field measurements. Decay status of CWD significantly (P<0.001) affected R_CWD. The temperature coefficient (Q₁₀) averaged 2.5, but varied by 141% across the 5-42°C temperature range, illustrating the potential shortcomings of using a constant Q₁₀. The CWD temperature was positively correlated to air temperature (R²=0.79, P<0.001), with a hysteresis effect that was correlated to CWD decay status and stand leaf area index . Ignoring this temperature hysteresis introduced errors of -1% to +32% in annual R_CWD estimates. Increasing T_CWD modeling time step from hourly to daily or monthly introduced a 5-11% underestimate in annual R_CWD. The annual R_CWD values in this study were more than two-fold greater than those in a previous study, illustrating the need to incorporate spatial and temporal responses of R_CWD to temperature and water content into models for long-term R_CWD estimation in boreal forest ecosystems.     

Water availability and carbon isotope discrimination in conifers

The stable C isotope composition ('¹³C) of leaf and wood tissue has been used as an index of water availability at both the species and landscape level. However, the generality of this relationship across species has received little attention. We compiled literature data for a range of conifers and examined relationships among landscape and environmental variables (altitude, precipitation, evaporation) and '¹³C. A significant component of the variation in '¹³C was related to altitude (discrimination decreased with altitude in stemwood, 2.53‰ km^-1 altitude, r²=0.49, and in foliage, 1.91‰ km^-1, r²=0.42), as has been noted previously. The decrease in discrimination with altitude was such that the gradient in CO₂ partial pressure into the leaf (P_a-P_i) and altitude were generally unrelated. The ratio of precipitation to evaporation (P/E) explained significant variation in P_a-P_i of stemwood (r²=0.45) and foliage (r²=0.27), but only at low (<0.8) P/E. At greater P/E there was little or no relationship, and other influences on '¹³C probably dominated the effect of water availability. We also examined the relationship between plant drought stress (O) and '¹³C within annual rings of stemwood from Pinus radiata and Pinus pinaster in south-western Australia. Differential thinning and fertiliser application produced large differences in the availability of water, nutrients and light to individual trees. At a density of 750 stems ha^-1, O and '¹³C were less (more negative) than at 250 stems ha^-1 indicating greater drought stress and less efficient water use, contrary to what was expected in light of the general relationship between discrimination and P/E. The greater '¹³C of trees from heavily thinned plots may well be related to an increased interception of radiation by individual trees and greater concentrations of nutrients in foliage - attributes that increase rates of photosynthesis, reduce P_i and increase '¹³C. '¹³C was thus modified to a greater extent by interception of radiation and by nutrient concentrations than by water availability and the '¹³C-O relationship varied between thinning treatments. Within treatments, the relationship between '¹³C and O was strong (0.38<r²<0.58). We conclude that '¹³C may well be a useful indicator of water availability or drought stress, but only in seasonally dry climates (P/E<1) and where variation in other environmental factors can be accounted for.