Effects of season,needle age and elevated atmospheric CO(2) on photosynthesis in Scots pine (Pinus sylvestris)

Five-year-old Scots pine (Pinus sylvestris L.) seedlings were grown in open-top chambers at ambient and elevated (ambient + 400 &mgr;mol mol(-1)) CO(2) concentrations. Net photosynthesis (A), specific leaf area (SLA) and concentrations of nitrogen (N), carbon (C), soluble sugars, starch and chlorophyll were measured in current-year and 1-year-old needles during the second year of CO(2) enrichment. The elevated CO(2) treatment stimulated photosynthetic rates when measured at the growth CO(2) concentration, but decreased photosynthetic capacity compared with the ambient CO(2) treatment. Acclimation to elevated CO(2) involved decreases in carboxylation efficiency and RuBP regeneration capacity. Compared with the ambient CO(2) treatment, elevated CO(2) reduced light-saturated photosynthesis (when measured at 350 &mgr;mol mol(-1) in both treatments) by 18 and 23% (averaged over the growing season) in current-year and 1-year-old needles, respectively. We observed significant interactive effects of CO(2) treatment, needle age and time during the growing season on photosynthesis. Large seasonal variations in photosynthetic parameters were attributed to changes in needle chemistry, needle structure and feedbacks governed by whole-plant growth dynamics. Down-regulation of photosynthesis was probably a result of reduced N concentration on an area basis, although a downward shift in the relationship between photosynthetic parameters and N was also observed.     

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).     

Effects of needle age, long-term temperature and CO(2) treatments on the photosynthesis of Scots pine

Naturally regenerated 20-25-year-old Scots pine (Pinus sylvestris L.) trees were grown in open-top chambers in the presence of an elevated temperature or CO(2) concentration, or both. The elevated temperature treatment was administered year-round for 3 years. The CO(2) treatment was applied between April 15 and September 15 for 2 years. The photosynthetic responses of 1- and 2-year-old needles to varying photon flux densities (0-1500 micro mol m(-2) s(-1)) and CO(2) concentrations (350, 700 and 1400 micro mol mol(-1)) during measurement were determined. The CO(2) treatment alone increased maximum photosynthetic rate and light-use efficiency, but decreased dark respiration rate, light compensation and light saturation regardless of needle age. In contrast, the temperature treatment decreased maximum photosynthetic rate and photosynthetic efficiency, but increased dark respiration rate, light compensation and light saturation. The aging of needles affected the photosynthetic performance of the shoots; values of all parameters except photosynthetic efficiency were less in 2- than in 1-year-old needles. The CO(2) treatment decreased and the temperature treatment enhanced the reduction in maximum photosynthesis due to needle aging.     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.     

Atmospheric carbon dioxide, irrigation, and fertilization effects on phenolic and nitrogen concentrations in loblolly pine (Pinus taeda) needles

Concentrations of total soluble phenolics, catechin, proanthocyanidins (PA), lignin and nitrogen (N) were measured in loblolly pine (Pinus taeda L.) needles exposed to either ambient CO(2) concentration ([CO(2)]), ambient plus 175 or ambient plus 350 micromol CO(2) mol(-1) in branch chambers for 2 years. The CO(2) treatments were superimposed on a 2 x 2 factorial combination of irrigation and fertilization treatments. In addition, we compared the effects of branch chambers and open-top chambers on needle chemistry. Proanthocyanidin and N concentrations were measured in needles from branch chambers and from trees in open-top chambers exposed concurrently for two years to either ambient [CO(2)] or ambient plus 200 micromol CO(2) mol(-1) in combination with a fertilization treatment. In the branch chambers, concentrations of total soluble phenolics in needles generally increased with needle age. Concentrations of total soluble phenolics, catechin and PA in needle extracts increased about 11% in response to the elevated [CO(2)] treatments. There were no significant treatment effects on foliar lignin concentrations. Nitrogen concentrations were about 10% lower in needles from the elevated [CO(2)] treatments than in needles from the ambient [CO(2)] treatments. Soluble phenolic and PA concentrations were higher in the control and irrigated soil treatments in about half of the comparisons; otherwise, differences were not statistically significant. Needle N concentrations increased 23% in response to fertilization. Treatment effects on PA and N concentrations were similar between branch and open-top chambers, although in this part of the study N concentrations were not significantly affected by the CO(2) treatments in either the branch or open-top chambers. We conclude that elevated [CO(2)] and low N availability affected foliar chemical composition, which could in turn affect plant-pathogen interactions, decomposition rates and mineral nutrient cycling.     

Atmospheric carbon dioxide concentration, nitrogen availability, temperature and the photosynthetic capacity of current-year Norway spruce shoots

Branches of field-grown Norway spruce (Picea abies (L.) Karst.) trees were exposed to either long-term ambient or to elevated CO2 concentrations ([CO2]) using the branch bag technique. The light-saturated photosynthetic rates (A(max)) of current-year shoots differing in nitrogen (N) status were measured at various temperatures and at either ambient (360 micromol mol(-1), AMB) or elevated (ambient + 350 micromol mol(-1), EL) [CO2]. The value of A(max) was determined at various intercellular [CO2]s (A/Ci curves) and used to normalize photosynthetic rates to the mean treatment C(i) values, which were 200 micromol mol(-1) (AMB) and 450 micromol mol(-1) (EL), respectively. Needle N status and temperature strongly affected A(max). The response to N increased with temperature, and the photosynthetic temperature optimum increased with N status. This was assumed to be a result of reduced mesophyll CO2 conductance. The relative increase of Amax in the EL treatment compared to the AMB treatment varied from 15 to 90%, and increased with temperature, but decreased with N status. Nevertheless, the absolute photosynthetic response to EL increased with shoot N status. The relative increase in the instantaneous response of A(max) to elevated [CO2] was about 20% higher than the long-term response, i.e., there was downward acclimation in Amax in response to elevated [CO2]. The photosynthetic temperature optimum increased 4 degrees C with either a short- or a long-term increase in [CO2]. The bag treatment itself increased A(max) by approximately 16% and the temperature optimum of A(max) by approximately 3 degrees C.     

Canopy position and needle age affect photosynthetic response in field-grown Pinus radiata after five years of exposure to elevated carbon dioxide partial pressure

Photosynthesis of tree seedlings is generally enhanced during short-term exposure to elevated atmospheric CO2 partial pressure, but longer-term studies often indicate some degree of photosynthetic adjustment. We present physiological and biochemical evidence to explain observed long-term photosynthetic responses to elevated CO2 partial pressure as influenced by needle age and canopy position. We grew Pinus radiata D. Don. trees in open-top chambers for 5 years in sandy soil at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressures. The trees were well watered and exposed to natural light and ambient temperature. In the fourth year of CO2 exposure (fall 1997), when foliage growth had ceased for the year, photosynthetic down-regulation was observed in 1-year-old needles, but not in current-year needles, suggesting a reduction in carbohydrate sink strength as a result of increasing needle age (Turnbull et al. 1998). In 5-year-old trees (spring 1997), when foliage expansion was occurring, photosynthetic down-regulation was not observed, reflecting significantly large sinks for carbohydrates throughout the tree. Net photosynthesis was stimulated by 79% in trees growing in elevated CO2 partial pressure, but there was no significant effect on photosynthetic capacity or Rubisco activity and concentration. Current-year needles were more responsive to elevated CO2 partial pressure than 1-year-old needles, exhibiting larger relative increases in net photosynthesis to elevated CO2 partial pressure (98 versus 64%). Lower canopy and upper canopy leaves exhibited similar relative responses to growth in elevated CO2 partial pressure. However, needles in the upper canopy exhibited higher net photosynthesis, photosynthetic capacity, and Rubisco activity and concentration than needles in the lower canopy. Given that the ratio of mature to juvenile foliage mass in the canopy will increase as trees mature, we suggest that trees may become less responsive to elevated CO2 partial pressure with increasing age. We conclude that tree response to elevated CO2 partial pressure is based primarily on sink strength and not on the duration of exposure.     

Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO(2) and temperature

Biochemical models of photosynthesis suggest that rising temperatures will increase rates of net carbon dioxide assimilation and enhance plant responses to increasing atmospheric concentrations of CO(2). We tested this hypothesis by evaluating acclimation and ontogenetic drift in net photosynthesis in seedlings of five boreal tree species grown at 370 and 580 &mgr;mol mol(-1) CO(2) in combination with day/night temperatures of 18/12, 21/15, 24/18, 27/21, and 30/24 degrees C. Leaf-area-based rates of net photosynthesis increased between 13 and 36% among species in plants grown and measured in elevated CO(2) compared to ambient CO(2). These CO(2)-induced increases in net photosynthesis were greater for slower-growing Picea mariana (Mill.) B.S.P., Pinus banksiana Lamb., and Larix laricina (Du Roi) K. Koch than for faster-growing Populus tremuloides Michx. and Betula papyrifera Marsh., paralleling longer-term growth differences between CO(2) treatments. Measures at common CO(2) concentrations revealed that net photosynthesis was down-regulated in plants grown at elevated CO(2). In situ leaf gas exchange rates varied minimally across temperature treatments and, contrary to predictions, increasing growth temperatures did not enhance the response of net photosynthesis to elevated CO(2) in four of the five species. Overall, the species exhibited declines in specific leaf area and leaf nitrogen concentration, and increases in total nonstructural carbohydrates in response to CO(2) enrichment. Consequently, the elevated CO(2) treatment enhanced rates of net photosynthesis much more when expressed on a leaf area basis (25%) than when expressed on a leaf mass basis (10%). In all species, rates of leaf net CO(2) exchange exhibited modest declines with increasing plant size through ontogeny. Among the conifers, enhancements of photosynthetic rates in elevated CO(2) were sustained through time across a wide range of plant sizes. In contrast, for Populus tremuloides and B. papyrifera, mass-based photosynthetic rates did not differ between CO(2) treatments. Overall, net photosynthetic rates were highly correlated with relative growth rate as it varied among species and treatment combinations through time. We conclude that interspecific variation may be a more important determinant of photosynthetic response to CO(2) than temperature.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

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)].     

Effect of elevated carbon dioxide concentration and root restriction on net photosynthesis, water relations and foliar carbohydrate status of loblolly pine seedlings

To determine the effects of CO(2)-enriched air and root restriction on photosynthetic capacity, we measured net photosynthetic rates of 1-year-old loblolly pine seedlings grown in 0.6-, 3.8- or 18.9-liter pots in ambient (360 micro mol mol(-1)) or 2x ambient CO(2) (720 micro mol mol(-1)) concentration for 23 weeks. We also measured needle carbohydrate concentration and water relations to determine whether feedback inhibition or water stress was responsible for any decreases in net photosynthesis. Across all treatments, carbon dioxide enrichment increased net photosynthesis by approximately 60 to 70%. Net photosynthetic rates of seedlings in the smallest pots decreased over time with the reduction occurring first in the ambient CO(2) treatment and then in the 2x ambient CO(2) treatment. Needle starch concentrations of seedlings grown in the smallest pots were two to three times greater in the 2x ambient CO(2) treatment than in the ambient CO(2) treatment, but decreased net photosynthesis was not associated with increased starch or sugar concentrations. The reduction in net photosynthesis of seedlings in small pots was correlated with decreased needle water potentials, indicating that seedlings in the small pots had restricted root systems and were unable to supply sufficient water to the shoots. We conclude that the decrease in net photosynthesis of seedlings in small pots was not the result of CO(2) enrichment or an accumulation of carbohydrates causing feedback inhibition, but was caused by water stress.     

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.     

Response of photosynthesis in second-generation Pinus radiata trees to long-term exposure to elevated carbon dioxide partial pressure

Second-generation Pinus radiata D. Don trees, propagated from cuttings of 4-year-old trees previously grown at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressure (Ca) were grown under the same conditions in open-top chambers for a further year. As cuttings of the original trees, these second-generation trees were physiologically the same age as the first-generation trees with the only difference between the two being size. This allowed us to test the effects of tree size independently of age or duration of exposure. Total non-structural carbohydrate concentration, area-based nitrogen concentration, leaf mass per unit area and chlorophyll concentration measured in three foliage age cohorts were unaffected by either age or Ca. There were no signs of photosynthetic down-regulation in trees grown at elevated Ca. When measured at the growth Ca, photosynthetic rate in young needles during summer, autumn and spring was 34, 43 and 38% higher, respectively, in trees grown at elevated Ca than in trees grown at ambient Ca. In older needles, the corresponding photosythetic rate increases were 26, 47 and 49%. Water-use efficiency, determined by stable carbon isotope analysis, was 49% higher in foliage in the elevated Ca treatment than in foliage in the ambient Ca treatment. This increase was entirely due to photosynthetic enhancement, because stomatal conductance did not differ between treatments. We conclude that down-regulation of photosynthesis at elevated Ca is related to tree size rather than tree age or duration of exposure, and that enhanced photosynthetic rates can be maintained while sink strength is high enough to use the excess photosynthates.elevated CO2, needle age, photosynthetic down-regulation, photosynthetic enhancement, sink strength, water-use efficiency.     

Effect of elevated [CO(2)] and varying nutrient application rates on physiology and biomass accumulation of Sitka spruce (Picea sitchensis)

Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings were supplied with solutions containing nitrogen (N) at 0.1 x or 2 x the optimum rate (low-N and high-N supply, respectively) and grown either outside in a control plot or inside open-top chambers and exposed to ambient (355 &mgr;mol mol(-1)) or elevated (700 &mgr;mol mol(-1)) CO(2) concentration ([CO(2)]). Gas exchange measurements, chlorophyll determinations and nutrient analysis were made on current-year (< 1-year-old) shoots of the upper whorl after the seedlings had been growing in the [CO(2)] treatments for 17 months and the nutrient treatments for 6 months. Total seedling biomass and biomass allocation were assessed at the end of the experiment. Nutrient treatment had a significant effect on the light response curves, irrespective of [CO(2)] or chamber treatment; seedlings supplied with high-N rates had higher net photosynthetic rates than seedlings supplied with low-N rates. The degree of photosynthetic stimulation in response to elevated [CO(2)] was larger in seedlings receiving high-N rates than in seedlings receiving low-N rates. Light-saturated net photosynthesis of seedlings grown and measured in elevated [CO(2)] was 26% higher than that of seedlings grown and measured in ambient [CO(2)]. There was no significant effect of [CO(2)] or chamber treatment on the CO(2) response curves of seedlings receiving High-N supply rates. In contrast, analysis of the CO(2) response curves of seedlings receiving Low-N supply rates showed acclimation to elevated [CO(2)]. Both maximum rate of carboxylation (V(cmax)) and maximum electron transport capacity (J(max)) were lower and J(max)/V(cmax) higher in seedlings in the elevated [CO(2)] treatment. There was no effect of elevated [CO(2)] on stomatal conductance, although it was highly dependent on foliar [N], ranging from ~60 mmol m(-2) s(-1) at ~1.5 g N m(-2) to 200 mmol m(-2) s(-1) at ~5 g N m(-2). In the high-N and low-N treatments, foliar N concentration was 10 and 28% lower in seedlings grown in elevated [CO(2)] than in seedlings grown in ambient [CO(2)], respectively. There was no [CO(2)] effect on foliar phosphorus concentration ([P]). Chlorophyll concentration increased with increasing N supply in all treatments. There was no significant effect of elevated [CO(2)] on specific leaf area. Chlorophyll concentration expressed either on an area or dry mass basis for a given foliar [N] was higher in seedlings grown in elevated [CO(2)] than in seedings grown in ambient [CO(2)]. Elevated [CO(2)] increased total biomass accumulation by 37% in seedlings in the high-N treatment but had no effect in seedlings in the low-N treatment. There was a proportionally bigger allocation of biomass to roots of seedlings in the elevated [CO(2)] + low-N supply rate treatment compared with seedlings in other treatments. This resulted in a reduction in aboveground biomass compared with corresponding seedlings grown in ambient [CO(2)].     

Growth and photosynthetic traits of hybrid larch F1 (Larix gmelinii var. japonica x L. kaempferi) under elevated CO2 concentration with low nutrient availability

The hybrid larch F(1) (Larix gmelinii var. japonica × Larix kaempferi) is considered one of the most important tree species not only for timber production but also as an afforestation material for severe conditions such as infertile soil. To predict the ability of hybrid larch F(1) as an afforestation material under potential climates in the future, it is important to understand the response of hybrid larch F(1) to elevated CO(2) concentration ([CO(2)]) under low nutrient availability. Three-year-old seedlings of hybrid larch F(1) were grown under two different levels of [CO(2)], 360 (ambient) and 720 μmol mol(-1) (elevated), in combination with two different levels of nitrogen (N) supply (0 and 30 kg ha(-1)) for one growing season. Elevated [CO(2)] reduced the maximum rates of carboxylation and electron transport in the needles. Net photosynthetic rates at growth [CO(2)] (i.e., 360 and 720 μmol mol(-1) for ambient and elevated treatment, respectively) did not differ between the two CO(2) treatments. Reductions in N content and N use efficiency to perform photosynthetic functions owing to the deficiency of nutrients other than N, such as P and K, and/or increase in cell wall mass were considered factors of photosynthetic down-regulation under elevated [CO(2)], whereas stomatal closure little affected the photosynthetic down-regulation. Although we observed strong down-regulation of photosynthesis, the dry matter increase of hybrid larch F(1) seedlings was enhanced under elevated [CO(2)]. This is mainly attributable to the increase in the amount of needles on increasing the number of sylleptic branches. These results suggest that elevated CO(2) may increase the growth of hybrid larch F(1) even under low nutrient availability, and that this increase may be regulated by changes in both crown architecture and needle photosynthesis, which is mainly affected not by stomatal limitation but by biochemical limitation.     

Effects of foliar nitrogen concentration on photosynthesis and water use efficiency in Douglas-fir

Leaf-level physiological processes were studied in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) to determine whether apparent increases in stand-level water use efficiency (WUE) observed in response to nitrogen (N) fertilization were attributable to foliar N effects on carbon fixation rates or on stomatal control of water loss. Photosynthesis and transpiration were measured at different light intensities and ambient CO(2) molar fractions and comparisons were made between current-year shoots with average foliar N concentrations of 1.58% (High-N) and 1.25% (Low-N). Photosynthetic rates and foliar N concentrations were positively correlated. In response to light, photosynthesis and stomatal conductance were closely coupled and a similar coupling was observed in response to different ambient CO(2) concentrations. Partitioning the photosynthetic responses into mesophyll and stomatal components indicated that foliar N altered mesophyll conductance but not stomatal control of water loss. High-N shoots had significantly greater rates of photosynthesis and transpiration than Low-N shoots and, as a result, instantaneous WUE did not differ significantly between High-N and Low-N shoots.     

Variations in dark respiration and mitochondrial numbers within needles of Pinus radiata grown in ambient or elevated CO2 partial pressure

Within-leaf variations in cell size, mitochondrial numbers and dark respiration rates were compared in the most recently expanded tip, the mid-section and base of needles of Pinus radiata D. Don trees grown for 4 years in open-top chambers at ambient (36 Pa) or elevated (65 Pa) carbon dioxide partial pressure (p(CO2)a). Mitochondrial numbers and respiratory activity varied along the length of the needle, with the highest number of mitochondria per unit cytoplasm and the highest rate of respiration per unit leaf area at the base of the needle. Regardless of the location of the cells (tip, middle or basal sections), needles collected from trees grown in elevated p(CO2)a had nearly twice the number of mitochondria per unit cytoplasm as those grown in ambient p(CO2)a. This stimulation of mitochondrial density by growth at elevated p(CO2)a was greater at the tip of the needle (2.7 times more mitochondria than in needles grown in ambient CO2) than at the base of the needle (1.7 times). The mean size of individual mitochondria was unaffected either by growth at elevated p(CO2)a or by position along the needle. Tree growth at elevated p(CO2)a had a variable effect on respiration per unit leaf area, significantly increasing respiration in the tip of the needles (+25%) and decreasing respiration at the mid-section and base of the needles (-14% and -25%, respectively). Although a simple relationship between respiration per unit leaf area and mitochondrial number per unit cytoplasm was found within each CO2 treatment, the variable effect of growth at elevated p(CO2)a on respiration along the length of the needles indicates that a more complex relationship must determine the association between structure and function in these needles.     

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.     

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].     

Effects of elevated carbon dioxide concentration on growth and nitrogen fixation in Alnus glutinosa in a long-term field experiment

Nitrogen-fixing plant species may respond more positively to elevated atmospheric carbon dioxide concentrations ([CO2]) than other species because of their ability to maintain a high internal nutrient supply. A key factor in the growth response of trees to elevated [CO2] is the availability of nitrogen, although how elevated [CO2] influences the rate of N2-fixation of nodulated trees growing under field conditions is unclear. To elucidate this relationship, we measured total biomass, relative growth rate, net assimilation rate (NAR), leaf area and net photosynthetic rate of N2-fixing Alnus glutinosa (L.) Gaertn. (common alder) trees grown for 3 years in open-top chambers in the presence of either ambient or elevated atmospheric [CO2] and two soil N regimes: full nutrient solution or no fertilizer. Nitrogen fixation by Frankia spp. in the root nodules of unfertilized trees was assessed by the acetylene reduction method. We hypothesized that unfertilized trees would show similar positive growth and physiological responses to elevated [CO2] as the fertilized trees. Growth in elevated [CO2] stimulated (relative) net photosynthesis and (absolute) total biomass accumulation. Relative total biomass increased, and leaf nitrogen remained stable, only during the first year of the experiment. Toward the end of the experiment, signs of photosynthetic acclimation occurred, i.e., down-regulation of the photosynthetic apparatus. Relative growth rate was not significantly affected by elevated [CO2] because although NAR was increased, the effect on relative growth rate was negated by a reduction in leaf area ratio. Neither leaf area nor leaf P concentration was affected by growth in elevated [CO2]. Nodule mass increased on roots of unfertilized trees exposed to elevated [CO2] compared with fertilized trees exposed to ambient [CO2]. There was also a biologically significant, although not statistically significant, stimulation of nitrogenase activity in nodules exposed to elevated [CO2]. Root nodules of trees exposed to elevated [CO2] were smaller and more evenly spaced than root nodules of trees exposed to ambient [CO2]. The lack of an interaction between nutrient and [CO2] effects on growth, biomass and photosynthesis indicates that the unfertilized trees maintained similar CO2-induced growth and photosynthetic enhancements as the fertilized trees. This implies that alder trees growing in natural conditions, which are often limited by soil N availability, should nevertheless benefit from increasing atmospheric [CO2].