Carbon exchange rates, chlorophyll content, and carbohydrate status of two forest tree species exposed to carbon dioxide enrichment

Seedlings of yellow-poplar (Liriodendron tulipifera L.) and white oak (Quercus alba L.) were exposed continuously to one of three CO(2) concentrations in open-top chambers under field conditions and evaluated after 24 weeks with respect to carbon exchange rates (CER), chlorophyll (Chl) content, and diurnal carbohydrate status. Increasing the CO(2) concentration from ambient to +150 or +300 microl l(-1) stimulated CER of yellow-poplar and white oak seedlings by 60 and over 35%, respectively, compared to ambient-grown seedlings. The increases in CER were not associated with a significant change in stomatal conductance and occurred despite a reduction in the amounts of Chl and accessory pigments in the leaves of plants grown in CO(2)-enriched air. Total Chl contents of yellow-poplar and white oak seedlings grown at +300 microl l(-1) were reduced by 27 and over 55%, respectively, compared with ambient-grown seedlings. Yellow-poplar and white oak seedlings grown at +300 microl l(-1) contained 72 and 67% more morning starch, respectively, than did ambient-grown plants. In contrast, yellow-poplar and white oak seedlings grown at +300 microl l(-1) contained 17 and 27% less evening sucrose, respectively, than did plants grown at ambient CO(2) concentration. Diurnal starch accumulation and the subsequent depletion of sucrose contributed to a pronounced increase in the starch/sucrose ratio of plants grown in CO(2)-enriched air. All seedlings exhibited a substantial reduction in dark respiration as CO(2) concentration increased, but the significance of this increase to the carbohydrate status and carbon economy of plants grown in CO(2)-enriched air remains unclear.     

Influence of elevated CO2 and mycorrhizae on nitrogen acquisition: contrasting responses in Pinus taeda and Liquidambar styraciflua

An understanding of root system capacity to acquire nitrogen (N) is critical in assessing the long-term growth impact of rising atmospheric CO2 concentration ([CO2]) on trees and forest ecosystems. We examined the effects of mycorrhizal inoculation and elevated [CO2] on root ammonium (NH4+) and nitrate (NO3-) uptake capacity in sweetgum (Liquidambar styraciflua L.) and loblolly pine (Pinus taeda L.). Mycorrhizal treatments included inoculation of seedlings with the arbuscular mycorrhizal (AM) fungus Glomus intraradices Schenck & Smith in sweetgum and the ectomycorrhizal (EM) fungus Laccaria bicolor (Maire) Orton in loblolly pine. These plants were then equally divided between ambient and elevated [CO2] treatments. After 6 months of treatment, root systems of both species exhibited a greater uptake capacity for NH4+ than for NO3-. In both species, mycorrhizal inoculation significantly increased uptake capacity for NO3-, but not for NH4+. In sweetgum, the mycorrhizal effect on NO3- and NH4+ uptake capacity depended on growth [C02]. Similarly, in loblolly pine, the mycorrhizal effect on NO3- uptake capacity depended on growth [CO2], but the effect on NH4+ uptake capacity did not. Mycorrhizal inoculation significantly enhanced root nitrate reductase activity (NRA) in both species, but elevated [CO2] increased root NRA only in sweetgum. Leaf NRA in sweetgum did not change significantly with mycorrhizal inoculation, but increased in response to [CO2]. Leaf NRA in loblolly pine was unaffected by either treatment. The results indicate that the mycorrhizal effect on specific root N uptake in these species depends on both the form of inorganic N and the mycorrhizal type. However, our data show that in addressing N status of plants under high [CO2], reliable prediction is possible only when information about other root system adjustments (e.g., biomass allocation to fine roots) is simultaneously considered.     

Physiological adjustment of two full-sib families of ponderosa pine to elevated CO(2)

Seeds from two full-sib families of ponderosa pine (Pinus ponderosa) with known differences in growth rates were germinated and grown in an ambient (350 micro l l(-1)) or elevated (700 micro l l(-1)) CO(2) concentration. Gas exchange at both ambient and elevated CO(2) concentrations was measured 1, 6, 39, and 112 days after the seed coat was shed. Initial stimulation of CO(2) exchange rate (CER) by elevated CO(2) was large (> 100%). On Day 1, CER of seedlings grown in elevated CO(2) and measured at ambient CO(2) was significantly lower than the CER of seedlings grown and measured at ambient CO(2), indicating physiological adjustment of the seedlings exposed to elevated CO(2). Physiological acclimation to elevated CO(2) was complete by Day 39 when there was no significant difference in CER between seedlings grown and measured at ambient CO(2) and seedlings grown and measured at elevated CO(2). After 4 months, the light response of seedlings in the two treatments was determined at both ambient and elevated CO(2). Light compensation point, CER at light saturation, and apparent quantum efficiency of seedlings grown and measured at ambient CO(2) were not significantly different from those of seedlings grown and measured at elevated CO(2). With a short-term increase in CO(2), CER at light saturation (5.16 +/- 0.52 versus 3.13 +/- 0.30 micro mol CO(2) m(-2) s(-1)) and apparent quantum efficiency (0.082 +/- 0.011 versus 0.045 +/- 0.003 micro mol CO(2) micro mol(-1) quanta) were significantly increased. Leaf C/N ratio was significantly increased in the elevated CO(2) treatment. There were few significant differences between families for any response to elevated CO(2). Under the experimental conditions, high growth rate was not correlated with a greater response to elevated CO(2).     

Production efficiency of loblolly pine and sweetgum in response to four years of intensive management

We tested the hypothesis that productivity of intensively managed loblolly pine (Pinus taeda L.) and sweetgum (Liquidambar styraciflua L.) stands is dependent not only on leaf area, but also on foliar photosynthetic rate. Effects of irrigation (irrigation treatment), irrigation with a fertilizer solution (fertigation treatment), and fertigation + pest control (loblolly pine only; fertigation + pest control treatment) on leaf physiology and growth were compared with control plots during the third and fourth growing seasons. Complete weed control was maintained on all plots. Aboveground net primary productivity of loblolly pine and sweetgum increased from 16.3 to 40.5 Mg ha(-1) and from 4.2 to 23.9 Mg ha(-1), respectively, in response to the most intensive treatment. Relative to the control treatment, neither fertigation of sweetgum nor fertigation + pest control of loblolly pine had a significant or consistent influence on foliar N concentration, quantum yield, carboxylation efficiency, net photosynthesis, stomatal conductance, or production efficiency (increment in woody biomass per unit leaf area). Irrigation increased predawn leaf water potential and photosynthesis of loblolly pine, but it had no effect on production efficiency. Leaf area was the predominant determinant of maximum productivity in these rapidly growing plantations.     

Responses of loblolly pine seedlings to elevated CO(2) and fluctuating water supply

Osmotic adjustment of loblolly pine (Pinus taeda L.) seedlings to fluctuating water supply in elevated CO(2) was investigated. Seedlings were grown in controlled-environment chambers in either 350 or 700 micro l l(-1) CO(2) with weekly watering for four months, after which they were either watered weekly (well-watered treatment) or every two weeks (water-stress treatment) for 59 days. Osmotic adjustment was assessed by pressure-volume analysis of shoots and by analysis of soluble carbohydrates and free amino acids in roots during the last drying cycle. In well-watered seedlings, elevated CO(2) increased the concentration of soluble sugars in roots by 68%. Water stress reduced the soluble sugar concentration in roots of seedling growing in ambient CO(2) to 26% of that in roots of well-watered seedlings. Elevated CO(2) mitigated the water stress-induced decrease in the concentration of soluble sugars in roots. However, this was probably due, in part, to carbohydrate loading during the first four months when all seedlings were grown in the presence of a high water supply, rather than to osmotic adjustment to water stress. Water stress caused a doubling in the concentration of free primary amino acids in roots, whereas elevated CO(2) reduced primary amino acid and nitrogen concentrations to 32 and 74%, respectively, of those in roots of seedlings grown in ambient CO(2). There was no indication of large-scale osmotic adjustment to water stress or that elevated CO(2) enhanced osmotic adjustment in loblolly pine.     

Radiation-use efficiency of a forest exposed to elevated concentrations of atmospheric carbon dioxide

We compared radiation-use efficiency of growth (epsilon;), defined as rate of biomass accumulation per unit of absorbed photosynthetically active radiation, of forest plots exposed to ambient (approximately 360 micro l l-1) or elevated (approximately 560 micro l l-1) atmospheric CO2 concentration ([CO2]). Large plots (30-m diameter) in a loblolly pine (Pinus taeda L.) plantation, which contained several hardwood species in the understory, were fumigated with a free-air CO2 enrichment system. Biomass accumulation of the dominant loblolly pines was calculated from monthly measurements of tree growth and site-specific allometric equations. Depending on the species, leaf area index (L*) was estimated by three methods: optical, allometric and litterfall. Based on the relationship between tree height and diameter during the first 3 years of exposure, we conclude that elevated [CO2] did not alter the pattern of aboveground biomass allocation in loblolly pine. There was considerable variation in L* estimates by the different methods; total L* was 18-42% lower when estimated by the optical method compared with estimates from allometric calculations, and this discrepancy was reduced when optical measurements were corrected for the non-random distribution of loblolly pine foliage. The allometric + litterfall approach revealed a seasonal maximum total L* of 6.2-7.1 with about 1/3 of the total from hardwood foliage. Elevated [CO2] had only a slight effect on L* in the first 3 years of this study. Mean epsilon; (+/- SD), calculated for loblolly pine only, was 0.49 +/- 0.05 and 0.62 +/- 0.04 g MJ-1 for trees in the ambient and elevated [CO2] plots, respectively. The 27% increase in epsilon; in response to CO2 enrichment was caused primarily by the stimulation of biomass increment, as there was only a small effect of elevated [CO2] on L* during the initial years of fumigation. Long-term increases in atmospheric [CO2] can increase epsilon; in closed-canopy forests but the absolute magnitude and duration of this increase remain uncertain.     

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.     

Chlorophyll and carotenoid concentrations in two varieties of Pinus ponderosa seedlings subjected to long-term elevated carbon dioxide

Two varieties of ponderosa pine (Pinus ponderosa Dougl. var. scopulorum (Rocky Mountain variety) and P. ponderosa var. ponderosa (Sierran variety)) seedlings were subjected to elevated atmospheric CO(2) for two and a half years. The CO(2) concentrations were ambient, ambient + 75 microl l(-1), ambient + 150 microl l(-1) and ambient + 300 microl l(-1), or approximately 350, 425, 500 and 650 microl l(-1) CO(2). After one and a half years of exposure to elevated CO(2) and until the end of the study, seedlings of both varieties showed symptoms of stress including mottling, mid-needle abscission and early senescence. In both varieties, exposure to CO(2) concentrations greater than ambient + 75 microl l(-1) resulted in lower chlorophyll a, chlorophyll b and carotenoid concentrations. At elevated CO(2) concentrations, the concentrations of pigments in needles of the Sierran variety were lower than those in the Rocky Mountain variety. Also, at elevated CO(2) concentrations, the pigment concentrations in the 1-year-old needles of both P. ponderosa varieties were lower than those in current-season needles.     

Effect of elevated CO2 on monoterpene emission of young Quercus ilex trees and its relation to structural and ecophysiological parameters

We investigated growth, leaf monoterpene emission, gas exchange, leaf structure and leaf chemical composition of 1-year-old Quercus ilex L. seedlings grown in ambient (350 microl l(-1)) and elevated (700 microl l(-1)) CO2 concentrations ([CO2]). Monoterpene emission and gas exchange were determined at constant temperature and irradiance (25 degrees C and 1000 micromol m(-2) s(-1) of photosynthetically active radiation) at an assay [CO2] of 350 or 700 microl l(-1). Measurements were made on intact shoots after the end of the growing season between mid-October and mid-February. On average, plants grown in elevated [CO2] had significantly increased foliage biomass (about 50%). Leaves in the elevated [CO2] treatment were significantly thicker and had significantly higher concentrations of cellulose and lignin and significantly lower concentrations of nitrogen and minerals than leaves in the ambient [CO2] treatment. Leaf dry matter density and leaf concentrations of starch, soluble sugars, lipids and hemi-cellulose were not significantly affected by growth in elevated [CO2]. Monoterpene emissions of seedlings were significantly increased by elevated [CO2] but were insensitive to short-term changes in assay [CO2]. On average, plants grown in elevated [CO2] had 1.8-fold higher monoterpene emissions irrespective of the assay [CO2]. Conversely, assay [CO2] rapidly affected photosynthetic rate, but there was no apparent long-term acclimation of photosynthesis to growth in elevated [CO2]. Regardless of growth [CO2], photosynthetic rates of all plants almost doubled when the assay [CO2] was switched from 350 to 700 microl l(-1). At the same assay [CO2], mean photosynthetic rates of seedlings in the two growth CO2 treatments were similar. The percentage of assimilated carbon lost as monoterpenes was not significantly altered by CO2 enrichment. Leaf emission rates were correlated with leaf thickness, leaf concentrations of cellulose, lignin and nitrogen, and total plant leaf area. In all plants, monoterpene emissions strongly declined during the winter independently of CO2 treatment. The results are discussed in the context of the acquisition and allocation of resources by Q. ilex seedlings and evaluated in terms of emission predictions.     

Aboveground biomass and nitrogen in four short-rotation woody crop species growing with different water and nutrient availabilities

We quantified the effect of water and nutrient availability on aboveground biomass and nitrogen accumulation and partitioning in four species from the southeastern United States, loblolly pine (Pinus taeda), slash pine (Pinus elliottii), sweetgum (Liquidambar styraciflua), and sycamore (Platanus occidentalis). The 6-year-old stands received five levels of resource input (control, irrigation with 3.05 cm water week⁻¹, irrigation + 57 kg N ha⁻¹ year⁻¹, irrigation + 85 kg N ha⁻¹ year⁻¹, and irrigation + 114 kg N ha⁻¹ year⁻¹). Irrigation significantly increased foliage, stem, and branch biomass for sweetgum and sycamore, culminating in 103% and 238% increases in total aboveground biomass. Fertilization significantly increased aboveground components for all species resulting in 49, 58, 281, and 132% increases in total aboveground biomass for loblolly pine, slash pine, sweetgum, and sycamore, respectively. Standing total aboveground biomass of the fertilized treatments reached 79, 59, 48, and 54 Mg ha⁻¹ for loblolly pine, slash pine, sweetgum, and sycamore, respectively. Fertilization increased foliar nitrogen concentration for loblolly pine, sweetgum, and sycamore foliage. Irrigation increased total stand nitrogen content by 6, 14, 93, and 161% for loblolly pine, slash pine, sweetgum, and sycamore, respectively. Fertilization increased total nitrogen content by 62, 53, 172, and 69% with maximum nitrogen contents of 267, 212, 237, and 203 kg ha⁻¹ for loblolly pine, slash pine, sweetgum, and sycamore, respectively. Growth efficiency (stem growth per unit of leaf biomass) and nitrogen use efficiency (stem growth per unit of foliar nitrogen content) increased for the sycamore and sweetgum, but not the loblolly or slash pine.     

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.     

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.     

Carbon dioxide concentration and nitrogen input affect the C and N storage pools in Amanita muscaria-Picea abies mycorrhizae

We studied the influence of elevated atmospheric CO2 concentration ([CO2]) on the vacuolar storage pool of nitrogen-containing compounds and on the glycogen pool in the hyphal sheath of Amanita muscaria (L. ex Fr.) Hooker-Picea abies L. Karst. mycorrhizae grown with two concentrations of ammonium in the substrate. Mycorrhizal seedlings were grown in petri dishes on agar containing 5.3 or 53 mg N l(-1) and exposed to 350 or 700 microl CO2 l(-1) for 5 or 7 weeks, respectively. Numbers and area of nitrogen-containing bodies in the vacuoles of the mycorrhizal fungus were determined by light microscopy linked to an image analysis system. The relative concentration of nitrogen in the vacuolar bodies was measured by electron energy loss spectroscopy (EELS). Glycogen stored in the cytosol was determined at the ultrastructural level by image analysis after staining the sections (PATAg test). Shoot dry weight, net photosynthesis and relative amounts of N in vacuolar bodies were greater at the higher N and CO2 concentrations. The numbers and areas of vacuolar N-containing bodies were significantly greater at the higher N concentration only at ambient [CO2]. In the same treatment the percentage of hyphae containing glycogen declined to nearly zero. We conclude that, in the high N/low [CO2] treatment, the mycorrhizal fungus had an insufficient carbohydrate supply, partly because of increased amino acid synthesis by the non-mycorrhizal rootlets. When [CO2] was increased, the equilibrium between storage of glycogen and N-containing compounds was reestablished.     

Prescribed burning and hexazinone herbicide as release treatments in a sapling hardwood-loblolly pine stand

To compare release treatments, a randomized complete block study was established in a 7-year-old hardwood-loblolly pine (Pinus taeda L.) stand in central Louisiana established using chopping and burning. There were 5 blocks of 3 treatments each: (1) check, (2) hexazinone applied once, and (3) prescribed backfiring applied twice. The first burn in December 1985 (7 years after site preparation) had a fire intensity of 90 kJ/s/m. The hexazinone herbicide was applied in April 1986 (the 8th year after site preparation) with a metered spotgun applicator at a rate of 3.0 kg active ingredient/ha. The second burn in March 1989 (the 11th year after site preparation) had a fire intensity of 106 kJ/s/m.The two prescribed burns increased the number of stems less than 1.5 m tall from 1,380 to 2,960/ha red maple (Acer rubrum L.), blackgum (Nyssa sylvatica var. bifora), and sweetgum (Liquidambar styraciflua L.). Burning reduced the number of loblolly pines less than 2.0 m tall, which resulted in a significant increase in average loblolly pine height and diameter. Loblolly pine comprised 62 and 59% of the basal area on the check and burn treatments, respectively, 11 years after site preparation. Hexazinone reduced the number of blackgum, sweetgum, and oak (Quercus spp.) from 6,100 to 4,560 stems/ha and resulted in significantly less hardwood tree basal area than found on the check or burn treatments. Therefore, the herbicide treatment resulted in principally a loblolly pine stand (over 80% of the total tree basal area was pine) four years after hexazinone application.     

Effects of carbon dioxide, fertilization, and irrigation on photosynthetic capacity of loblolly pine trees

Branches of nine-year-old loblolly pine trees grown in a 2 x 2 factorial combination of fertilization and irrigation were exposed for 11 months to ambient, ambient + 175, or ambient + 350 micro mol mol(-1) CO(2). Rates of light-saturated net photosynthesis (A(max)), maximum stomatal conductance to water vapor (g(max)), and foliar nitrogen concentration (% dry mass) were assessed monthly from April 1993 until September 1993 on 1992 foliage (one-year-old) and from July 1993 to March 1994 on 1993 foliage (current-year). Rates of A(max) of foliage in the ambient + 175 CO(2) treatment and ambient + 350 were 32-47 and 83-91% greater, respectively, than that of foliage in the ambient CO(2) treatment. There was a statistically significant interaction between CO(2) treatment and fertilization or irrigation treatment on A(max) on only one measurement date for each age class of foliage. Light-saturated stomatal conductance to water vapor (g(max)) was significantly affected by CO(2) treatment on only four measurement dates. Light-saturated g(max) in winter was only 42% of summer g(max) even though soil water during winter was near field capacity and evaporative demand was low. Fertilization increased foliar N concentration by 30% over the study period when averaged across CO(2) treatments. During the study period, the ambient + 350 CO(2) treatment decreased average foliar N concentration of one-year-old foliage in the control, irrigated, fertilized and irrigated + fertilized plots by 5, 6.4, 9.6 and 11%, respectively, compared with one-year-old foliage in the corresponding ambient CO(2) treatments. The percent increase in A(max) due to CO(2) enrichment was similar in all irrigation and fertilization treatments and the effect persisted throughout the 11-month study period for both one-year-old and current-year foliage.     

Aluminum sensitivity of loblolly pine and slash pine seedlings grown in solution culture

To probe variation in Al sensitivity of two co-occurring pine species, seedlings from six full-sib families of loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Engelm.) were grown in solution culture containing 4.4 mM (high-Al) or 0.01 mM (low-Al) AlCl(3) at pH 4 for 58 days. On average, both pine species had 41% less total dry weight in the high-Al treatment than in the low-Al treatment. Stem volume growth of slash pine was more sensitive to the high-Al treatment than that of loblolly pine. In both species, the high-Al treatment inhibited root dry weight more than shoot dry weight. Within-species variation in Al sensitivity among families was greater in loblolly pine (24 to 52% inhibition of seedling dry weight) than in slash pine (35 to 47% inhibition of seedling dry weight). Foliar Al concentration was positively correlated with Al sensitivity in slash pine but not in loblolly pine; however, in both species, the concentration of Al in roots was 20-fold greater than in foliage.     

Biochemical acclimation patterns of Betula pendula and Pinus sylvestris seedlings to elevated carbon dioxide concentrations

Acclimation of photosynthesis to increasing atmospheric carbon dioxide concentration ([CO2]; 350 to 2,000 micromol mol-1) was followed in silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings for two years. Chlorophyll fluorescence and concentrations of Rubisco, chlorophyll, total soluble protein and nitrogen were monitored together with steady-state gas exchange at three CO2 concentrations (ambient [CO2] (345 +/- 20 micromol mol-1), the growth [CO2] and 1950 +/- 55 micromol mol-1). Rubisco and chlorophyll concentrations decreased in birch and Scots pine with increasing growth [CO2]. A nonlinear response was recorded for Rubisco and chlorophyll concentrations in birch, which was correlated with a significant decrease in specific leaf area. Nitrogen concentration decreased in birch leaves, but was unchanged in Scots pine needles. The species differed substantially in their steady-state CO2 exchange response to increasing growth [CO2]. The principal effect in birch was a significant nonlinear decrease in the steady-state gas exchange rate at the ambient [CO2], whereas in Scots pine the main effect was a significant increase in the steady-state gas exchange rate at the growth [CO2].     

Timing and magnitude of C partitioning through a young loblolly pine (Pinus taeda L.) stand using 13C labeling and shade treatments

The dynamics of rapid changes in carbon (C) partitioning within forest ecosystems are not well understood, which limits improvement of mechanistic models of C cycling. Our objective was to inform model processes by describing relationships between C partitioning and accessible environmental or physiological measurements, with a special emphasis on short-term C flux through a forest ecosystem. We exposed eight 7-year-old loblolly pine (Pinus taeda L.) trees to air enriched with (13)CO(2) and then implemented adjacent light shade (LS) and heavy shade (HS) treatments in order to manipulate C uptake and flux. The impacts of shading on photosynthesis, plant water potential, sap flow, basal area growth, root growth and soil CO(2) efflux rate (CER) were assessed for each tree over a 3-week period. The progression of the (13)C label was concurrently tracked from the atmosphere through foliage, phloem, roots and surface soil CO(2) efflux. The HS treatment significantly reduced C uptake, sap flow, stem growth and fine root standing crop, and resulted in greater residual soil water content to 1 m depth. Soil CER was strongly correlated with sap flow on the previous day, but not the current day, with no apparent treatment effect on the relationship. Although there were apparent reductions in new C flux belowground, the HS treatment did not noticeably reduce the magnitude of belowground autotrophic and heterotrophic respiration based on surface soil CER, which was overwhelmingly driven by soil temperature and moisture. The (13)C label was immediately detected in foliage on label day (half-life = 0.5 day), progressed through phloem by Day 2 (half-life = 4.7 days), roots by Days 2-4, and subsequently was evident as respiratory release from soil which peaked between Days 3 and 6. The δ(13)C of soil CO(2) efflux was strongly correlated with phloem δ(13)C on the previous day, or 2 days earlier. While the (13)C label was readily tracked through the ecosystem, the fate of root C through respiratory, mycorrhizal or exudative release pathways was not assessed. These data detail the timing and relative magnitude of C flux through various components of a young pine stand in relation to environmental conditions.     

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.     

Carbon dioxide enrichment accelerates the decline in nutrient status and relative growth rate of Populus tremuloides Michx. seedlings

Changes in growth dynamics and mineral nutrient concentrations were measured in Populus tremuloides Michx., trembling aspen, grown for 100 days following germination in atmospheres containing 350 or 750 microl l(-1) CO(2). Seedlings were fertilized with nitrogen (N) at concentrations of 15.5 mM (high-N), 1.55 mM (medium-N), or 0.155 mM (low-N). Initially, relative growth rates were enhanced by CO(2) enrichment in each N regime, but the effects did not persist. In plants grown in high-N or medium-N, foliar concentrations of Ca and Mg decreased in response to CO(2) enrichment. During the 100-day study, whole-plant concentrations of N and P decreased in all treatments. The decreases in mineral nutrient concentrations over time were accelerated in CO(2)-enriched plants and accompanied the disappearance of the CO(2)-induced growth enhancement. It is concluded that the depression of relative growth rates often associated with long-term CO(2) enrichment of plants may result from decreases in plant nutrient status.