Isotopic carbon discrimination and leaf nitrogen content of Erica arborea L. along a CO(2) concentration gradient in a CO(2) spring in Italy

We studied a Mediterranean species (Erica arborea L.) growing in a CO(2) spring in Italy that was naturally exposed for generations to a gradient of atmospheric CO(2) concentrations. The CO(2) concentration gradient to which different individual plants were exposed was determined by an indirect method based on radioisotope analysis. The stable carbon isotope ratio of sampled leaves was determined by mass spectrometry, and isotopic discrimination was then calculated. Leaf nitrogen, specific leaf area, total soil nitrogen, soil organic matter content and soil pH were also measured. In one group of plants, grown on a homogeneous soil and exposed to moderate CO(2) enrichment, isotopic discrimination was significantly reduced in response to increasing CO(2) concentrations, whereas the intercellular CO(2) concentration and leaf nitrogen content were almost unaffected. In a second group of plants, grown along a gradient of CO(2) concentration and soil nitrogen content, leaf nitrogen content was reduced when nitrogen availability was limiting. However, when soil nitrogen was available in excess, even very high CO(2) concentrations did not result in increased discrimination or reduced leaf nitrogen content in the long term. The results are discussed with respect to current theories about the long-term CO(2) response of plants based on several years of experimentation with elevated atmospheric CO(2) concentrations under controlled conditions.     

Kinetics of nitrogen uptake by Populus tremuloides in relation to atmospheric CO(2) and soil nitrogen availability

Sustained increases in plant production in response to elevated atmospheric carbon dioxide (CO(2)) concentration may be constrained by the availability of soil nitrogen (N). However, it is possible that plants will respond to N limitation at elevated CO(2) concentration by increasing the specific N uptake capacity of their roots. To explore this possibility, we examined the kinetics of (15)NH(4) (+) and (15)NO(3) (-) uptake by excised roots of Populus tremuloides Michx. grown in ambient and twice-ambient CO(2) concentrations, and in soils of low- and high-N availability. Elevated CO(2) concentration had no effect on either NH(4) (+) or NO(3) (-) uptake, whereas high-N availability decreased the capacity of roots to take up both NH(4) (+) and NO(3) (-). The maximal rate of NH(4) (+) uptake decreased from 12 to 8 &mgr;mol g(-1) h(-1), and K(m) increased from 49 to 162 &mgr;mol l(-1), from low to high soil N availability.Because NO(3) (-) uptake exhibited mixedkinetics over the concentration range we used (10-500 &mgr;mol l( -1)), it was not possible to calculate V(max) and K(m). Instead, we used an uptake rate of 100 &mgr;mol g(-1) h(-1) as our metric of NO(3) (-) uptake capacity, which averaged 0.45 and 0.23 &mgr;mol g(-1) h(-1) at low- and high-N availability, respectively. The proximal mechanisms for decreased N uptake capacity at high-N availability appeared to be an increase in fine-root carbohydrate status and a decrease in fine-root N concentration. Both NH(4) (+) and NO(3) (-) uptake were inversely related to fine-root N concentration, and positively related to fine-root total nonstructural carbohydrate concentration. We conclude that soil N availability, through its effects on fine-root N and carbohydrate status, has a much greater influence on the specific uptake capacity of P. tremuloides fine roots than elevated atmospheric CO(2). In elevated atmospheric CO(2), changes in N acquisition by P. tremuloides appeared to be driven by changes in root architecture and biomass, rather than by changes in the amount or activity of N-uptake enzymes.     

Ecophysiological responses of woody plants to past CO(2) concentrations

An approach is detailed for calculating historical rates of CO(2) uptake and water loss of leaves from measurements of leaf delta(13)C composition and climatic information. This approach was applied to investigate leaf gas exchange metabolism of woody taxa during the past 200 years of atmospheric CO(2) increase and in response to the longer-term atmospheric CO(2) increases plants experienced over the Pleistocene. Reconstructed net assimilation rates and water use efficiencies increased in response to increasing atmospheric CO(2) concentrations in both sets of material, whereas stomatal conductance, showing the combined responses of changes in stomatal density and leaf assimilation rates, was generally less responsive. Woody temperate taxa maintained a nearly constant c(i)/c(a) ratio in response to the increase in atmospheric CO(2) concentrations over both timescales, in part, as a result of changes in stomatal density. The reconstructed leaf-scale physiological responses to past global climatic and atmospheric change corroborated those anticipated from experimental work indicating the adequate capacity of experiments, at least at the scale of individual leaves, to predict plant responses to future environmental change.     

Free-air CO2 enrichment (FACE) enhances biomass production in a short-rotation poplar plantation

This paper investigates the possible contribution of Short Rotation Cultures (SRC) to carbon sequestration in both current and elevated atmospheric CO2 concentrations ([CO2]). A dense poplar plantation (1 x 1 m) was exposed to a [CO2] of 550 ppm in Central Italy using the free-air CO2 enrichment (FACE) technique. Three species of Populus were examined, namely P. alba L., P. nigra L. and P. x euramericana Dode (Guinier). Aboveground woody biomass of trees exposed to elevated [CO2] for three growing seasons increased by 15 to 27%, depending on species. As a result, light-use efficiency increased. Aboveground biomass allocation was unaffected, and belowground biomass also increased under elevated [CO2] conditions, by 22 to 38%. Populus nigra, with total biomass equal to 62.02 and 72.03 Mg ha-1 in ambient and elevated [CO2], respectively, was the most productive species, although its productivity was stimulated least by atmospheric CO2 enrichment. There was greater depletion of inorganic nitrogen from the soil after three growing seasons in elevated [CO2], but no effect of [CO2] on stem wood density, which differed significantly only among species.     

Effects of predicted future and current atmospheric temperature and [CO2] and high and low soil moisture on gas exchange and growth of Pinus taeda seedlings at cool and warm sites in the species range

Predicted future changes in air temperature and atmospheric CO(2) concentration ([CO(2)]), coupled with altered precipitation, are expected to substantially affect tree growth. Effects on growth may vary considerably across a species range, as temperatures vary from sub-optimal to supra-optimal for growth. We performed an experiment simultaneously at two locations in the current range of loblolly pine, a cool site and a warm site, to examine the effect of future climate conditions on growth of loblolly pine seedlings in contrasting regions of the species range. At both sites 1-year-old loblolly pine seedlings were grown in current (local ambient temperature and [CO(2)]) and predicted future atmospheric conditions (ambient +2 °C temperature and 700 μmol mol(-1) [CO(2)]). Additionally, high and low soil moisture treatments were applied within each atmospheric treatment at each site by altering the amount of water provided to the seedlings. Averaged across water treatments, photosynthesis (A(net)) was 31% greater at the cool site and 34% greater at the warm site in elevated temperature and [CO(2)] compared with ambient temperature. Biomass accumulation was also stimulated by 38% at the cool site and by 24% at the warm site in that treatment. These results suggest that a temperature increase of 2 °C coupled with an increase in [CO(2)] (predicted future climate) will create conditions favorable for growth of this species. Reduced soil moisture decreased growth in both current and predicted atmospheric conditions. Biomass accumulation and A(net) were reduced by ～39 and 17%, respectively, in the low water treatment. These results suggest that any benefit of future atmospheric conditions may be negated if soil moisture is reduced by altered precipitation patterns.     

Leaf physiology, production, water use, and nitrogen dynamics of the grassland invader Acacia smallii at elevated CO(2) concentrations

Invasion by woody legumes can alter hydrology, nutrient accumulation and cycling, and carbon sequestration on grasslands. The rate and magnitude of these changes are likely to be sensitive to the effects of atmospheric CO(2) enrichment on growth and water and nitrogen dynamics of leguminous shrubs. To assess potential effects of increased atmospheric CO(2) concentrations on plant growth and acquisition and utilization of water and nitrogen, seedlings of Acacia smallii Isely (huisache) were grown for 13 months at CO(2) concentrations of 385 (ambient), 690, and 980 micro mol mol(-1). Seedlings grown at elevated CO(2) concentrations exhibited parallel declines in leaf N concentration and photosynthetic capacity; however, at the highest CO(2) concentration, biomass production increased more than 2.5-fold as a result of increased leaf photosynthetic rates, leaf area, and N(2) fixation. Measurements of leaf gas exchange and aboveground biomass production and soil water balance indicated that water use efficiency increased in proportion to the increase in atmospheric CO(2) concentration. The effects on transpiration of an accompanying decline in leaf conductance were offset by an increase in leaf area, and total water loss was similar across CO(2) treatments. Plants grown at elevated CO(2) fixed three to four times as much N as plants grown at ambient CO(2) concentration. The increase in N(2) fixation resulted from an increase in fixation per unit of nodule mass in the 690 micro mol mol(-1) CO(2) treatment and from a large increase in the number and mass of nodules in plants in the 980 micro mol mol(-1) CO(2) treatment. Increased symbiotic N(2) fixation by woody invaders in response to CO(2) enrichment may result in increased N deposition in litterfall, and thus increased productivity on many grasslands.     

Elevated atmospheric CO(2) concentration changes ectomycorrhizal morphotype assemblages in Betula papyrifera

Ectomycorrhizae are extremely diverse, with different species of fungi having very different physiologies and morphologies that, in turn, confer a range of different benefits to the host plant. To test the hypothesis that elevated CO(2) leads to changes in the assemblage of ectomycorrhizae associated with trees, we examined the number and frequency of ectomycorrhizal morphotypes colonizing roots of Betula papyrifera Marsh. saplings grown at an ambient or elevated (700 ppm) atmospheric CO(2) concentration for 24 weeks. Elevated CO(2) resulted in significant changes in the composition of the ectomycorrhizal assemblage toward morphotypes with a higher incidence of emanating hyphae and rhizomorphs. We conclude that B. papyrifera saplings will be able to support a more costly mycorrhization in future elevated-CO(2) atmospheres.     

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.     

Elevated CO(2) concentration affects leaf photosynthesis-nitrogen relationships in Pinus taeda over nine years in FACE

To investigate whether long-term elevated carbon dioxide concentration ([CO(2)]) causes declines in photosynthetic enhancement and leaf nitrogen (N) owing to limited soil fertility, we measured photosynthesis, carboxylation capacity and area-based leaf nitrogen concentration (N(a)) in Pinus taeda L. growing in a long-term free-air CO(2) enrichment (FACE) facility at an N-limited site. We also determined how maximum rates of carboxylation (V(cmax)) and electron transport (J(max)) varied with N(a) under elevated [CO(2)]. In trees exposed to elevated [CO(2)] for 5 to 9 years, the slope of the relationship between leaf photosynthetic capacity (A(net-Ca)) and N(a) was significantly reduced by 37% in 1-year-old needles, whereas it was unaffected in current-year needles. The slope of the relationships of both V(cmax) and J(max) with N(a) decreased in 1-year-old needles after up to 9 years of growth in elevated [CO(2)], which was accompanied by a 15% reduction in N allocation to the carboxylating enzyme. Nitrogen fertilization (110 kg N ha(-1)) in the ninth year of exposure to elevated [CO(2)] restored the slopes of the relationships of V(cmax) and J(max) with N(a) to those of control trees (i.e., in ambient [CO(2)]). The J(max):V(cmax) ratio was unaffected by either [CO(2)] or N fertilization. Changes in the apparent allocation of N to photosynthetic components may be an important adjustment in pines exposed to elevated [CO(2)] on low-fertility sites. We conclude that fundamental relationships between photosynthesis or its component processes with N(a) may be altered in aging pine needles after more than 5 years of exposure to elevated atmospheric [CO(2)].     

Growth,water relations,and survival of drought-exposed seedlings from six maternal families of honey mesquite (Prosopis glandulosa): responses to CO(2) enrichment

Low water availability is a leading contributor to mortality of woody seedlings on grasslands, including those of the invasive shrub Prosopis. Increasing atmospheric CO(2) concentration could favor some genotypes of this species over others if there exists intraspecific variation in the responsiveness of survivorship to CO(2). To investigate such variation, we studied effects of CO(2) enrichment on seedling survival in response to uniform rates of soil water depletion in six maternal families of honey mesquite (P. glandulosa Torr. var. glandulosa). Three families each from the arid and mesic extremes of the species' distribution in the southwestern United States were studied in environmentally controlled glasshouses. Relative water content at turgor loss and osmotic potential were not affected by CO(2) treatment. Increased atmospheric CO(2) concentration, however, increased growth, leaf production and area, and midday xylem pressure potential, and apparently reduced transpiration per unit leaf area of seedlings as soil dried. Consequently, CO(2) enrichment about doubled the fraction of seedlings that survived soil water depletion. Maternal families of honey mesquite differed in percentage survival of drought and in several other characteristics, but differences were of similar or of smaller magnitude compared with differences between CO(2) treatments. There was no evidence for genetic variation in the responsiveness of survivorship to CO(2). By increasing seedling survival of drought, increasing atmospheric CO(2) concentration could increase the abundance of honey mesquite where establishment is limited by water availability. Genetic types with superior ability to survive drought today, however, apparently will maintain that advantage in the future.     

Long-term photosynthetic acclimation to increased atmospheric CO(2) concentration in young birch (Betula pendula) trees

To study the long-term response of photosynthesis to elevated atmospheric CO(2) concentration in silver birch (Betula pendula Roth.), 18 trees were grown in the field in open-top chambers supplied with 350 or 700 &mgr;mol mol(-1) CO(2) for four consecutive growing seasons. Maximum photosynthetic rates, stomatal conductance and CO(2) response curves were measured over the fourth growing season with a portable photosynthesis system. The photosynthesis model developed by Farquhar et al. (1980) was fitted to the CO(2) response curves. Chlorophyll, soluble proteins, total nonstructural carbohydrates, nitrogen and Rubisco activity were determined monthly. Elevated CO(2) concentration stimulated photosynthesis by 33% on average over the fourth growing season. However, comparison of maximum photosynthetic rates at the same CO(2) concentration (350 or 700 &mgr;mol mol(-1)) revealed that the photosynthetic capacity of trees grown in an elevated CO(2) concentration was reduced. Analysis of the response curves showed that acclimation to elevated CO(2) concentration involved decreases in carboxylation efficiency and RuBP regeneration capacity. No clear evidence for a redistribution of nitrogen within the leaf was observed. Down-regulation of photosynthesis increased as the growing season progressed and appeared to be related to the source-sink balance of the trees. Analysis of the main leaf components revealed that the reduction in photosynthetic capacity was accompanied by an accumulation of starch in leaves (100%), which was probably responsible for the reduction in Rubisco activity (27%) and to a lesser extent for reductions in other photosynthetic components: chlorophyll (10%), soluble protein (9%), and N concentrations (12%) expressed on an area basis. Despite a 21% reduction in stomatal conductance in response to the elevated CO(2) treatment, stomatal limitation was significantly less in the elevated, than in the ambient, CO(2) treatment. Thus, after four growing seasons exposed to an elevated CO(2) concentration in the field, the trees maintained increased photosynthetic rates, although their photosynthetic capacity was reduced compared with trees grown in ambient CO(2).     

Photosynthetic responses of Scots pine to elevated CO(2) and nitrogen supply: results of a branch-in-bag experiment

Naturally seeded Scots pine (Pinus sylvestris L.) trees, age 25-30 years, were subjected to two soil-nitrogen-supply regimes and to elevated atmospheric CO(2) concentrations by the branch-in-bag method from April 15 to September 15 for two or three years. Gas exchange in detached shoots was measured in a diffuse radiation field. Seven parameters associated with photosynthetic performance and two describing stomatal conductance were determined to assess the effects of treatments on photosynthetic components. An elevated concentration of CO(2) did not lead to a significant downward regulation in maximum carboxylation rate (V(cmax)) or maximum electron transport rate (J(max)), but it significantly decreased light-saturated stomatal conductance (g(sat)) and increased minimum stomatal conductance (g(min)). Light-saturated rates of CO(2) assimilation were higher (24-31%) in shoots grown and measured at elevated CO(2) concentration than in shoots grown and measured at ambient CO(2) concentration, regardless of treatment time or nitrogen-supply regime. High soil-nitrogen supply significantly increased photosynthetic capacity, corresponding to significant increases in V(cmax) and J(max). However, the combined elevated CO(2) + high nitrogen-supply treatment did not enhance the photosynthetic response above that observed in the elevated CO(2) treatment alone.     

Carbon allocation, root exudation and mycorrhizal colonization of Pinus echinata seedlings grown under CO(2) enrichment

Increased exudation of carbon compounds from roots may provide a mechanism for enhancement of nutrient availability to plants growing in a CO(2)-enriched atmosphere. Therefore, the effect of atmospheric CO(2) concentration on carbon allocation and root exudation was investigated in Pinus echinata Mill. (shortleaf pine) seedlings. After 34 and 41 weeks, seedlings growing in 695 microl l(-1) CO(2) allocated proportionately more (14)C-labeled photosynthate to fine roots than did seedlings growing in ambient air. This was associated with greater fine root mass and mycorrhizal density in CO(2)-enriched plants after 34 weeks. Exudation of soluble, (14)C-labeled compounds from roots also was greater in these plants at 34 weeks, but the effect of CO(2) concentration on exudation did not persist at 41 weeks.     

Responses of deciduous broadleaf trees to defoliation in a CO2 enriched atmosphere

Relatively little is known about the implications of atmospheric CO2 enrichment for tree responses to biotic disturbances such as folivory. We examined the combined effects of elevated CO2 concentration ([CO2]) and defoliation on growth and physiology of sugar maple (Acer saccharum Marsh.) and trembling aspen (Populus tremuloides Michx.). Seedlings were planted in the ground in eight open-top chambers. Four chambers were ventilated with CO2-enriched air (ambient + 283 micromol mol-1) and four chambers were supplied with ambient air. After 6 weeks of growth, half of the leaf area was removed on a subset of seedlings of each species in each CO2 treatment. We monitored subsequent biomass gain and allocation, along with leaf gas exchange and chemistry. Defoliation did not significantly affect final seedling biomass in either species or CO2 treatment. Growth recovery following defoliation was associated with increased allocation to leaf mass in maple and a slight enhancement of mean photosynthesis in aspen. Elevated [CO2] did not significantly affect aspen growth, and the observed stimulation of maple growth was significant only in mid-season. Correspondingly, simulated responses of whole-tree photosynthesis to elevated [CO2] were constrained by a decrease in photosynthetic capacity in maple, and were partially offset by reductions in specific leaf area and biomass allocation to foliage in aspen. There was a significant interaction between [CO2] and defoliation on only a few of the measured traits. Thus, the data do not support the hypothesis that atmospheric CO2 enrichment will substantially alter tree responses to folivory. However, our findings do provide further indication that regeneration-stage growth rates of certain temperate tree species may respond only moderately to a near doubling of atmospheric [CO2].     

Sustainable Management of Forests for Atmospheric CO2 Depletion

Abstract

The topic of forest sector carbon balance in connection with climate changes currently has both great scientific and political importance for ecological sustainability on a global scale. The concentration of CO2 in the atmosphere has increased by 1-2 ppm per year in the last few decades. The present paper examines the actual and potential role of forest management to deplete atmospheric CO2 concentration, with specific reference to the Italian situation as a case study. Italian carbon emission derived from fossil fuels amounts to 432 Mt/year, of which 65% comes from burning oil. Annual CO2 absorption, estimated by combining the 1985 National Forest Inventory data with selected biomass data, proves to be quite relevant, amounting to more than 10% of the annual Italian CO2 emission. This is a prudent estimate, since no account is taken of the contributions of non-tree vegetation and soil. Current forest management standards are mainly oriented to conservation. Optimisation of forest management specifically purposed also to carbon storage and non-wood products substitution may further enhance the role of Italian forest sector for atmospheric CO2 depletion: in such a view, practical issues (amelioration of existing forest stands; adjustment of harvesting yield to the actual production capacity of forest stands and adjustment of production standards towards high durable wood products; afforestation and tree cropping by means of changes in land use) are addressed within a sustainability framework.     

Responses of foliar gas exchange to long-term elevated CO(2) concentrations in mature loblolly pine trees

Branches of field-grown mature loblolly pine (Pinus taeda L.) trees were exposed for 2 years (1992 and 1993) to ambient or elevated CO(2) concentrations (ambient + 165 micro mol mol(-1) or ambient + 330 micro mol mol(-1) CO(2)). Exposure to elevated CO(2) concentrations enhanced rates of net photosynthesis (P(n)) by 53-111% compared to P(n) of foliage exposed to ambient CO(2). At the same CO(2) measurement concentration, the ratio of intercellular to atmospheric CO(2) concentration (C(i)/C(a)) and stomatal conductance to water vapor did not differ among foliage grown in an ambient or enriched CO(2) concentration. Analysis of the relationship between P(n) and C(i) indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase during growth in elevated CO(2) concentrations. Based on estimates derived from P(n)/C(i) curves, there were no apparent treatment differences in dark respiration, CO(2) compensation point or P(n) at the mean C(i). In 1992, foliage in the three CO(2) treatments yielded similar estimates of CO(2)-saturated P(n) (P(max)), whereas in 1993, estimates of P(max) were higher for branches grown in elevated CO(2) than in ambient CO(2). We conclude that field-grown loblolly pine trees do not exhibit downward acclimation of leaf-level photosynthesis in their long-term response to elevated CO(2) concentrations.     

Growth, shoot phenology and physiology of diverse seed sources of black spruce: I. Seedling responses to varied atmospheric CO(2) concentrations and photoperiods

We conducted a greenhouse experiment to determine: (1) if diverse provenances of black spruce (Picea mariana (Mill.) B.S.P.) respond similarly in growth, phenology and physiology to an approximately 300 ppm increase in atmospheric CO(2) concentration, and (2) the influence of photoperiod on both provenance and provenance x CO(2) interaction effects. Seedlings from provenances that originated from the Yukon (63 degrees 34' N, 135 degrees 55' W), British Columbia (58 degrees 47' N, 123 degrees 38' W), Alberta (52 degrees 22' N, 115 degrees 15' W), Newfoundland (50 degrees 54' N, 56 degrees 06' W) and Ontario (48 degrees 59' N, 80 degrees 38' W and 45 degrees 10' N, 77 degrees 10' W) were subjected to growth analysis in greenhouse growth chambers supplied with 712 +/- 93 (SD) ppm CO(2) (elevated) or 394 +/- 59 ppm CO(2) (ambient). Seedlings from Provenances 7000 and 6901 were also subjected to an extended photoperiod treatment and periodically measured for shoot and root gas exchange. In response to a natural photoperiod, southern provenances grew more, broke and set bud later, and partitioned more biomass to shoot versus root than northern provenances. These differences among provenances were influenced by the extended photoperiod treatment but not by the elevated CO(2) treatment. Averaged across all provenances, elevated CO(2) increased seedling final weights by 55%; however, the elevated CO(2) treatment had no effect on the provenance differences in any measured trait. We conclude that the large differences in physiology, phenology and growth among these diverse provenances of black spruce were expressed similarly in both ambient and elevated atmospheric CO(2) concentrations.     

Canopy position affects photosynthetic adjustments to long-term elevated CO2 concentration (FACE) in aging needles in a mature Pinus taeda forest

Few studies have examined the effects of elevated CO2 concentration ([CO2]) on the physiology of intact forest canopies, despite the need to understand how leaf-level responses can be aggregated to assess effects on whole-canopy functioning. We examined the long-term effects of elevated [CO2] (ambient + 200 ppm CO2) on two age classes of needles in the upper and lower canopy of Pinus taeda L. during the second through sixth year of exposure to elevated [CO2] in free-air (free-air CO2 enrichment (FACE)) in North Carolina, USA. Strong photosynthetic enhancement in response to elevated [CO2] (e.g., +60% across age classes and canopy locations) was observed across the years. This stimulation was 33% greater for current-year needles than for 1-year-old needles in the fifth and sixth years of treatment. Although photosynthetic stimulation in response to elevated [CO2] was maintained through the sixth year of exposure, we found evidence of concurrent down-regulation of Rubisco and electron transport capacity in the upper-canopy sunlit leaves. The lower canopy showed no evidence of down-regulation. The upper canopy down-regulated carboxylation capacity (Vcmax) and electron transport capacity (Jmax) by about 17-20% in 1-year-old needles; however, this response was significant across sampling years only for Jmax in 1-year-old needles (P < 0.02). A reduction in leaf photosynthetic capacity in aging conifer needles at the canopy top could have important consequences for canopy carbon balance and global carbon sinks because 1-year-old sunlit needles contribute a major proportion of the annual carbon balance of these conifers. Our finding of a significant interaction between canopy position and CO2 treatment on the biochemical capacity for CO2 assimilation suggests that it is important to take canopy position and needle aging into account because morphologically and physiologically distinct leaves could respond differently to elevated [CO2].     

Growth and carbon accumulation in root systems of Pinus taeda and Pinus ponderosa seedlings as affected by varying CO(2), temperature and nitrogen

It has been hypothesized that increasing atmospheric CO(2) concentration enhances accumulation of carbon in fine roots, thereby altering soil carbon dynamics and nutrient cycling. To evaluate possible changes to belowground pools of carbon and nitrogen in response to elevated CO(2), an early and a late successional species of pine (Pinus taeda L. and Pinus ponderosa Dougl. ex Laws, respectively) were grown from seed for 160 days in a 35 or 70 Pa CO(2) partial pressure at low or high temperature (30-year weekly mean and 30-year weekly mean + 5 degrees C) and a soil solution nitrogen concentration of 1 or 5 mM NH(4)NO(3) at the Duke University Phytotron. Seedlings were harvested at monthly intervals and growth parameters of the primary root, secondary root and tap root fractions evaluated. Total root biomass of P. ponderosa showed a positive CO(2) response (105% increase) (P = 0.0001) as a result of significant increases in all root fractions in the elevated CO(2) treatment, but all other main effects and interactions were insignificant. In P. taeda, there were significant interactions between CO(2) and temperature (P = 0.04) and CO(2) and nitrogen (P = 0.04) for total root biomass. An allometric analysis indicated that modulation of the secondary root fraction was the main response of the trees to altered environmental conditions. In P. ponderosa, there was an increase in the secondary root fraction relative to the primary and tap root fractions under conditions of low temperature. In P. taeda, there was a shift in carbon accumulation to the secondary roots relative to the primary roots under low temperature and low nitrogen. Neither species exhibited shifts in carbon accumulation in response to elevated CO(2). We conclude that both species have the potential to increase belowground biomass substantially in response to rising atmospheric CO(2) concentration, and this response is sensitive to temperature and nitrogen in P. taeda. Both species displayed small shifts in belowground carbon accumulation in response to altered temperature and nitrogen that may have substantial ecosystem consequences over time.     

Gas exchange and dry matter allocation responses to elevation of atmospheric CO(2) concentration in seedlings of three tree species

Photosynthetic rates of 13-month-old Pinus radiata D. Don, Nothofagus fusca (Hook f.) ?rst. and Pseudotsuga menziesii (Mirb.) Franco seedlings grown and measured at elevated atmospheric concentrations of CO(2) (~620 microl l(-1)) were 32 to 55% greater than those of seedlings grown and measured at ambient (~310 microl l(-1)) concentrations of CO(2). Seedlings grown in ambient and elevated concentrations of CO(2) had similar rates of photosynthesis when measured at ~620 microl l(-1) CO(2), but when measured at ~310 microl l(-1) CO(2), the P. radiata and N. fusca seedlings which were grown at elevated CO(2) had lower rates of photosynthesis than the seedlings grown at an ambient concentration of CO(2). Stomatal conductances in general were lower when measured at ~620 microl l(-1) CO(2) than at ~310 microl l(-1) CO(2). Stomatal conductances declined in all species grown at both CO(2) concentrations when the leaf-air water vapor concentration gradient (DeltaW) was increased from 10 to 20 mmol H(2)O mol(-1) air. The percent enhancement in photosynthesis for P. radiata and P. menziesii at elevated CO(2) was greater at 20 mmol than at 10 mmol DeltaW, suggesting that elevated CO(2) may moderate the effects of atmospheric water stress. Dry matter allocation patterns were not significantly different for plants grown in ambient or high CO(2) air.