Scots pine responses to elevated temperature and carbon dioxide concentration: growth and wood properties

Growth and wood properties of 20-year-old Scots pine (Pinus sylvestris L.) trees were studied for 6 years in 16 closed chambers providing a factorial combination of two temperature regimes (ambient and elevated) and two carbon dioxide concentrations ([CO2]) (ambient and twice ambient). The elevation of temperature corresponded to the predicted effect at the site of a doubling in atmospheric [CO2]. Annual height and radial growth and wood properties were analyzed during 1997-2002. Physical wood properties analyzed included early- and latewood widths and their proportions, intra-ring wood densities, early- and latewood density and mean fiber length. Chemical wood properties analyzed included concentrations of acetone-soluble extractives, lignin, cellulose and hemicellulose. There were no significant treatment effects on height growth during the 6-year study. Elevated [CO2] increased ring width by 66 and 47% at ambient and elevated temperatures, respectively. At ambient [CO2], elevated temperature increased ring width by 19%. Increased ring width in response to elevated [CO2] resulted from increases in both early- and latewood width; however, there was no effect of the treatments on early- and latewood proportions. Mean wood density, earlywood density and fiber length increased in response to elevated temperature. The chemical composition of wood was affected by elevated [CO2], which reduced the cellulose concentration, and by elevated temperature, which reduced the concentration of acetone-soluble extractives. Thus, over the 6-year period, radial growth was significantly increased by elevated [CO2], and some wood properties were significantly affected by elevated temperature or elevated [CO2], or both, indicating that climate change may affect the material properties of wood.     

Diameter growth of Scots pine (Pinus sylvestris) trees grown at elevated temperature and carbon dioxide concentration under boreal conditions

We investigated the impacts of elevated temperature and carbon dioxide concentration ([CO2]) on diameter growth of Scots pine (Pinus sylvestris L.), aged about 20 years, grown with a low nitrogen supply in closed chambers at (i) ambient temperature and [CO2] (AT+AC), (ii) ambient temperature and elevated [CO2] (AT+EC), (iii) elevated temperature and ambient [CO2] (ET+AC), and (iv). elevated temperature and [CO2] (ET+EC). Each treatment was replicated four times. Diameter growth was monitored with a band dendrograph at 15-min intervals throughout the growing seasons of 1997, 1998 and 1999. Over the monitoring period, diameter growth began 2-3 weeks earlier in trees in the ET+EC and ET+AC chambers than in trees in the AT+AC and AT+EC chambers. However, the cessation of growth occurred about a week later in trees in the ET+EC, ET+AC and AT+EC chambers compared with the AT+AC chambers. The duration of the growing season was 115 and 108 days in the ET+EC and ET+AC chambers, respectively, and 95 and 84 days in the AT+EC and AT+AC chambers, respectively. The ET+AC and ET+EC treatments enhanced diameter growth most early in the growing season, whereas in trees in the AT+AC and AT+EC treatments diameter growth rate was highest in the middle of the growing season. Diameter growth rate leveled off more slowly in trees in the ET+EC and AT+EC treatments than in the other treatments. The growth response to elevated T, elevated [CO2] or both decreased with time and it was less than the maximum observed in other studies for small seedlings and under optimal growth conditions. Nevertheless, cumulative diameter growth for the 3-year period was 67% greater in trees in the ET+EC treatment, and 57 and 26% greater in trees in the AT+EC and ET+AC treatments, respectively, compared with trees in the AT+AC treatment. Over the 3 years, [CO2] had a statistically significant (P < 0.10) effect on both absolute and relative diameter growth, but the interaction between [CO2] and temperature was not significant.     

Elevated temperature and CO(2) concentration effects on xylem anatomy of Scots pine

We studied the effects of elevated temperature and carbon dioxide concentration ([CO(2)]) alone and together on wood anatomy of 20-year-old Scots pine (Pinus sylvestris L.) trees. The study was conducted in 16 closed chambers, providing a factorial combination of two temperature regimes and two CO(2) concentrations (ambient and elevated), with four trees in each treatment. The climate scenario included a doubling of [CO(2)] and a corresponding increase of 2-6 degrees C in temperature at the site depending on the season. Anatomical characteristics analyzed were annual earlywood, latewood and ring widths, intra-ring wood densities (earlywood, latewood and mean wood density), tracheid width, length, wall thickness, lumen diameter, wall thickness:lumen diameter ratio and mass per unit length (coarseness), and numbers of rays, resin canals and tracheids per xylem cross-sectional area. Elevated [CO(2)] increased ring width in four of six treatment years; earlywood width increased in the first two years and latewood width in the third year. Tracheid walls in both the earlywood and latewood tended to become thicker over the 6-year treatment period when temperature or [CO(2)] was elevated alone, whereas in the combined treatment they tended to become thinner relative to the tracheids of trees grown under ambient conditions. Latewood tracheid lumen diameters were larger in all the treatments relative to ambient conditions over the 6-year period, whereas lumen diameters in earlywood increased only in response to elevated [CO(2)] and were 3-6% smaller in the treatments with elevated temperature than in ambient conditions. Tracheid width, length and coarseness were greater in trees grown in elevated than in ambient temperature. The number of resin canals per mm(2) decreased in the elevated [CO(2)] treatment and increased in the elevated temperature treatments relative to ambient conditions. The treatments decreased the number of rays and tracheids per mm(2) of cross-sectional area, the greatest decrease occurring in the elevated [CO(2)] treatment. It seemed that xylem anatomy was affected more by elevated temperature than by elevated [CO(2)] and that the effects of temperature were confined to the earlywood.     

Impact of needle age on the response of respiration in Scots pine to long-term elevation of carbon dioxide concentration and temperature

Sixteen 20-year-old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed in environment-controlled chambers that for 4 years maintained: (1) ambient conditions (CON); (2) elevated atmospheric carbon dioxide concentration [CO2] (ambient + 350 micromol mol-1; EC); (3) elevated temperature (ambient + 2-3 degrees C; ET); or (4) elevated [CO2] and temperature (EC+ET). Dark respiration rate, specific leaf area (SLA) and the concentrations of starch and soluble sugars in needles were measured in the fourth year. Respiration rates, on both an area and a mass basis, and SLA decreased in EC relative to CON, but increased in ET and EC+ET, regardless of needle age class. Starch and soluble sugar concentrations for a given needle age class increased in EC, but decreased slightly in ET and EC+ET. Respiration rates and SLA were highest in current-year needles in all treatments, whereas starch and soluble sugar concentrations were highest in 1-year-old needles. Relative to that of older needles, respiration of current-year needles was inhibited less by EC, but increased in response to ET and EC+ET. All treatments enhanced the difference in respiration between current-year and older needles relative to that in CON. Age had a greater effect on needle respiration than any of the treatments. There were no differences in carbohydrate concentration or SLA between needle age classes in response to any treatment. Relative to CON, the temperature coefficient (Q10) of respiration increased slightly in EC, regardless of age, but declined significantly in ET and EC+ET, indicating acclimation of respiration to temperature.     

Respiratory responses of Scots pine stems to 5 years of exposure to elevated CO2 concentration and temperature

Stem respiration in 20-year-old Scots pine (Pinus sylvestris L.) trees was examined following 5 years of exposure to ambient conditions (CON), elevated atmospheric carbon dioxide concentration ([CO2]) (ambient + 350 micromol mol(-1), (EC)), elevated temperature (ambient + 2-6 degrees C, (ET)) or a combination of elevated [CO2] and elevated temperature (ECT). Stem respiration varied seasonally regardless of the treatment and displayed a similar trend to temperature, with maximum rates occurring around Day 190 in summer and minimum rates in winter. Respiration normalized to 15 degrees C (R15) was higher in the growing season than in the non-growing season, whereas the temperature coefficient (Q10) was lower in the growing season. Annually averaged R15 was 0.36, 0.43, 0.40 and 0.44 micromol m(-2) s(-1) under CON, EC, ET and ECT conditions, respectively, whereas the corresponding values for total stem respiration were 6.55, 7.69, 7.50 and 7.90 mol m(-2) year(-1). The EC, ET and ECT treatments increased R15 by 18, 11 and 22%, respectively, relative to CON, and increased the modeled annual total stem respiration by 18, 15 and 21%. The increase in modeled annual stem respiration under EC and ECT conditions was caused mainly by higher maintenance respiration (22 and 25%, respectively, whereas the increase in growth respiration was 9 and 12%). Growth respiration was unaltered by ET. The treatments did not significantly affect the respiratory response to stem temperature; the mean Q10 value was 2.04, 2.10, 1.99 and 2.12 in the CON, EC, ET and ECT treatments, respectively. It is suggested that the increase in stem respiration was partly a result of the increased growth rate. We conclude that elevated [CO2] increased the maintenance component of respiration more than the growth component.     

Contrasting effects of elevated carbon dioxide concentration and temperature on Rubisco activity,chlorophyll fluorescence,needle ultrastructure and secondary metabolites in conifer seedlings

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were grown for 50 days in growth chambers in an ambient or twice ambient carbon dioxide concentration ([CO2]) at a day/night temperature of 19/12 degrees C or 23/16 degrees C. Although elevated [CO2] (EC) had only slight effects on the growth parameters measured, elevated temperature (ET) increased above ground dry mass of both species. Among treatments, biomass accumulation of both species was greatest in the combined EC + ET treatment. The EC treatment induced thylakoid swelling and increased numbers of plastoglobuli observed in Scots pine needles. Although EC had little effect on Rubisco protein or N concentration of needles, ET had a large effect on N-containing compounds and enhanced N allocation from 1-year-old needles. Terpenoids were more responsive to EC and ET than total phenolics. Generally, terpene concentrations were reduced by EC and increased by ET. Increased terpenoid concentrations in response to ET might be associated with thermotolerance of photosynthesis. In Norway spruce, EC decreased total phenolic concentrations in needles, probably as a result of increased growth. We conclude that, in seedlings of these boreal species, the effects of elevated [CO2] on the studied parameters were small compared with the effects of elevated temperature.     

Leaf photosynthetic characteristics of silver birch during three years of exposure to elevated concentrations of CO2 and O3 in the field

Effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on photosynthesis and related biochemistry of two European silver birch (Betula pendula Roth) clones were studied under field conditions during 1999-2001. Seven-year-old trees of Clones 4 and 80 were exposed for 3 years to the following treatments in an open-top chamber experiment: outside control (OC), chamber control (CC), 2x ambient [CO2] (EC), 2x ambient [O3] (EO) and 2x ambient [CO2] + 2x ambient [O3] (EC+EO). During the experiment, gas exchange, chlorophyll fluorescence, amount and activity of Rubisco, concentrations of chlorophyll, soluble protein, soluble sugars, starch, nitrogen (N) and carbon:nitrogen (C:N) ratio were determined in short- and long-shoot leaves. Elevated [CO2] increased photosynthetic rate by around 30% when measurements were made at the growth [CO2]. When measured at ambient [CO2], photosynthesis was around 15% lower in EC trees than in CC trees. This was related to a approximately 10% decrease in total leaf N, to 26 and 20% decreases in the amount and activity of Rubisco, respectively, and to a 49% increase in starch concentration in elevated [CO2]. Elevated [O3] had no significant effect on gas exchange parameters and its effect on biochemistry was small in both clones. However, elevated [O3] decreased the proportion of Rubisco in total soluble proteins and the apparent quantum yield of photosystem II (PSII) photochemistry in light and increased non-photochemical quenching in 2000. The interactive effect of CO2 and O3 was variable. Elevated [O3] decreased chlorophyll concentration only in EO trees, and the EC+EO treatment decreased the total activity of Rubisco and increased the C:N ratio more than the EO treatment alone. The small effect of elevated [O3] on photosynthesis indicates that these young silver birches were fairly tolerant to annual [O3] exposures that were 2-3 times higher than the AOT40 value of 10 ppm.h. set as a critical dose for forest trees.     

Silver birch and climate change: variable growth and carbon allocation responses to elevated concentrations of carbon dioxide and ozone

We studied the effects of elevated concentrations of carbon dioxide ([CO2]) and ozone ([O3]) on growth, biomass allocation and leaf area of field-grown O3-tolerant (Clone 4) and O3-sensitive clones (Clone 80) of European silver birch (Betula pendula Roth) trees during 1999-2001. Seven-year-old trees of Clones 4 and 80 growing outside in open-top chambers were exposed for 3 years to the following treatments: outside control (OC); chamber control (CC); 2 x ambient [CO2] (EC); 2 x ambient [O3] (EO); and 2 x ambient [CO2] + 2 x ambient [O3] (EC+EO). When the results for the two clones were analyzed together, elevated [CO2] increased tree growth and biomass, but had no effect on biomass allocation. Total leaf area increased and leaf abscission was delayed in response to elevated [CO2]. Elevated [O3] decreased dry mass of roots and branches and mean leaf size and induced earlier leaf abscission in the autumn; otherwise, the effects of elevated [O3] were small across the clones. However, there were significant interactions between elevated [CO2] and elevated [O3]. When results for the clones were analyzed separately, stem diameter, volume growth and total biomass of Clone 80 were increased by elevated [CO2] and the stimulatory effects of elevated [CO2] on stem volume growth and total leaf area increased during the 3-year study. Clone 80 was unaffected by elevated [O3]. In Clone 4, elevated [O3] decreased root and branch biomass by 38 and 29%, respectively, whereas this clone showed few responses to elevated [CO2]. Elevated [CO2] significantly increased total leaf area in Clone 80 only, which may partly explain the smaller growth responses to elevated [CO2] of Clone 4 compared with Clone 80. Although we observed responses to elevated [O3], the responses to the EC+EO and EC treatments were similar, indicating that the trees only responded to elevated [O3] under ambient [CO2] conditions, perhaps reflecting a greater quantity of carbohydrates available for detoxification and repair in elevated [CO2].     

Temperature responses of growth and wood anatomy in European beech saplings grown in different carbon dioxide concentrations

Effects of temperature on growth and wood anatomy were studied in young European beech (Fagus sylvatica L.) grown in 7-l pots for 2.5 years in field-phytotron chambers supplied with an ambient (approximately 400 micromol mol-1) or elevated (approximately 700 micromol mol-1) carbon dioxide concentration ([CO2]). Temperatures in the chambers ranged in increments of 2 degrees C from -4 degrees C to +4 degrees C relative to the long-term mean monthly (day and night) air temperature in Berlin-Dahlem. Soil was not fertilized and soil water and air humidity were kept constant. Data were evaluated by regression analysis. At final harvest, stem diameter was significantly greater at increased temperature (Delta1 degrees C: 2.4%), stems were taller (Delta1 degrees C: 8.5%) and stem mass tree-1 (Delta1 degrees C: 10.9%) and leaf area tree-1(Delta1 degrees C: 6.5%) were greater. Allocation pattern was slightly influenced by temperature: leaf mass ratio and leaf area ratio decreased with increasing temperature (Delta1 degrees C: 2.3% and 2.2% respectively), whereas stem mass/total mass increased (Delta1 degrees C: 2.1%). Elevated [CO2] enhanced height growth by 8.8% and decreased coarse root mass/total mass by 10.3% and root/shoot ratio by 11.7%. Additional carbon was mainly invested in aboveground growth. At final harvest a synergistic interaction between elevated [CO2] and temperature yielded trees that were 3.2% taller at -4 degrees C and 12.7% taller at +4 degrees C than trees in ambient [CO2]. After 2.5 seasons, cross-sectional area of the oldest stem part was approximately 32% greater in the +4 degrees C treatment than in the -4 degrees C treatment, and in the last year approximately 67% more leaf area/unit tree ring area was produced in the highest temperature regime compared with the lowest. Elevated [CO2] decreased mean vessel area of the 120 largest vessels per mm2 by 5.8%, causing a decrease in water conducting capacity. There was a positive interaction between temperature and elevated [CO2] for relative vessel area, which was approximately 38% higher at +4 degrees C than at -4 degrees C in elevated [CO2] compared with ambient [CO2]. Overall, temperature had a greater effect on growth than [CO2], but elevated [CO2] caused quantitative changes in wood anatomy.     

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.     

Radial variation of wood density components and ring width in cork oak trees

The radial variation of ring width and wood density was studied in cork oaks (Quercus suber) using microdensitometry. The observations were made in young never debarked cork oaks (30–40 years of age) and in mature trees under cork production (37–60 years of age). The cork oak wood is very dense (mean ring density 0.86 g.cm^?3, between 0.79 g.cm^?3 and 0.97 g.cm^?3) with a small intra-ring variability (mean earlywood density 0.80 g.cm^?3 and latewood density 0.90 g.cm^?3). The density components decreased from pith to bark more rapidly until the 15th ring, and then only slightly. There were no significant differences in the mean density components between never debarked trees and trees under cork production but their outwards decrease was accentuated in the never debarked trees. The annual growth was high, with a ring width mean of 3.9 mm (4.2 mm in the first 30 years) and the latewood represented 57% of the annual growth.     

Genetic control of intra-ring wood density variation in hybrid larch (Larix gmelinii var. japonica × L. kaempferi) F1

Genetic parameters for various wood density traits were estimated in 29-year-old trees of 18 full-sib families of hybrid larch (Larix gmelinii var. japonica × Larix kaempferi) F₁. Intra-ring density variation (IDV) was also evaluated using a model that expresses the pattern curve from earlywood to latewood as a power function. A high IDV indicates an abrupt change in wood density from earlywood to latewood. The ring width and wood density traits of individual rings were determined by X-ray densitometry. Overall wood density (RD) was shown to increase with increasing ring number, ranging from 0.42–0.59 g/cm³, whereas IDV of individual rings decreased gradually from pith outwards. Estimates of individual tree narrow-sense heritability of RD and IDV were 0.66 and 0.67, respectively. IDV showed negative genetic and phenotypic correlations with RD (r _g = −0.99, r _p = −0.72). The predicted genetic gains in latewood proportion and IDV were higher than that of RD. These results suggest that the intra-ring density variation is under moderate genetic control equivalent to wood density. The trend of increasing wood density from earlywood to latewood was associated with changes in both tracheid diameter and cell wall thickness.     

Lignification in beech (Fagus sylvatica) grown at elevated CO2 concentrations: interaction with nutrient availability and leaf maturation

Beech (Fagus sylvatica L.) seedlings were grown in an ambient or elevated CO2 concentration ([CO2]) either in small stands in microcosms for three to four seasons or individually in pots fertilized at different nutrient supply rates. Leaves at different stages of development, as well as stems and roots at the end of the growing season, were used for analysis of structural biomass and lignin. In elevated [CO2], lignification of leaves was slightly retarded compared with structural biomass production and showed a strong correlation with the activities of ionically, cell-wall-bound peroxidases but not with total soluble peroxidases or covalently wall-bound peroxidases. The effect of elevated [CO2] on lignin concentration of mature tissues was dependent on nutrient supply rate. In leaves and roots, elevated [CO2] increased the lignin concentration in dry mass in N-limited plants. In seedlings grown with high nutrient supply, the lignin concentration in dry mass was unaffected or diminished by elevated [CO2]. Because elevated [CO2] enhanced seedling growth in the high nutrient supply treatments, the total amount of lignin produced per seedling was higher in these treatments. We predict that long-term sequestration of carbon will increase as long as biomass production is stimulated by elevated [CO2] and that tissue quality will change depending on developmental stage and nutrient availability.     

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii)

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.     

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].     

Ring characteristics and screw withdrawal resistance of naturally regenerated <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> var. <Emphasis Type="Italic">formosana</Emphasis> trees

The ring characteristics and screw withdrawal resistance (SWR) of naturally regenerated Taiwan yellow cypress (Chamaecyparis obtusa var. formosana) trees were explored. Significant differences in average ring width (RW), earlywood width, latewood width, ring density (RD), earlywood density (ED), latewood density (LD), highest density (D_max), lowest density (D_min), latewood percentage (LWP), and SWR were observed between trees, rings (SWR excluded), and tree height positions. The RW components in the radial direction increased from the pith outward to about the 3rd to 5th ring and then decreased to about the 25th ring; it was almost constantly sustained toward the bark side. The RD in the radial direction slowly decreased from the pith outward to the bark side. Average ring width and ring density were significantly affected by the various tree growth rates, radial ring numbers, and tree height positions. ED, LD, D_max, D_min, and LWP were the most important factors determining the overall RD. RW did not correlate with tree RD. SWR is correlated with ED, RD, D_min, LWP, and intra-ring density variation (IDV). Thus, the SWR can be used to predict wood density and in nondestructive evaluation of a living tree.     

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].     

Dormancy release of Norway spruce under climatic warming: testing ecophysiological models of bud burst with a whole-tree chamber experiment

Ecophysiological models predicting timing of bud burst were tested with data gathered from 40-year-old Norway spruce (Picea abies (L.) Karst.) trees growing in northern Sweden in whole-tree chambers under climatic conditions predicted to prevail in 2100. Norway spruce trees, with heights between 5 and 7 m, were enclosed in individual chambers that provided a factorial combination of ambient (365 micromol mol-1) or elevated (700 micromol mol-1) atmospheric CO2 concentration, [CO2], and ambient or elevated air temperature. Temperature elevation above ambient ranged from +2.8 degrees C in summer to +5.6 degrees C in winter. Compared with control trees, elevated air temperature hastened bud burst by 2 to 3 weeks, whereas elevated [CO2] had no effect on the timing of bud burst. A simple model based on the assumption that bud rest completion takes place on a fixed calendar day predicted timing of bud burst more accurately than two more complicated models in which bud rest completion is caused by accumulated chilling. Together with some recent studies, the results suggest that, in adult trees, some additional environmental cues besides chilling are required for bud rest completion. Although it appears that these additional factors will protect trees under predicted climatic warming conditions, increased risk of frost damage associated with earlier bud burst cannot be ruled out. Inconsistent and partially anomalous results obtained in the model fitting show that, in addition to phenological data gathered under field conditions, more specific data from growth chamber and greenhouse experiments are needed for further development and testing of the models.     

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.     

Wood properties of trembling aspen and paper birch after 5 years of exposure to elevated concentrations of CO(2) and O(3)

We investigated the interactive effects of elevated concentrations of carbon dioxide ([CO(2)]) and ozone ([O(3)]) on radial growth, wood chemistry and structure of five 5-year-old trembling aspen (Populus tremuloides Michx.) clones and the wood chemistry of paper birch (Betula papyrifera Marsh.). Material for the study was collected from the Aspen FACE (free-air CO(2) enrichment) experiment in Rhinelander, WI, where the saplings had been exposed to four treatments: control, elevated [CO(2)] (560 ppm), elevated [O(3)] (1.5 x ambient) and their combination for five growing seasons. Wood properties of both species were altered in response to exposure to the treatments. In aspen, elevated [CO(2)] decreased uronic acids (constituents of, e.g., hemicellulose) and tended to increase stem diameter. In response to elevated [O(3)] exposure, acid-soluble lignin concentration decreased and vessel lumen diameter tended to decrease. Elevated [O(3)] increased the concentration of acetone-soluble extractives in paper birch, but tended to decrease the concentration of these compounds in aspen. In paper birch, elevated [CO(2)] decreased and elevated [O(3)] increased starch concentration. The responses of wood properties to 5 years of fumigation differed from those previously reported after 3 years of fumigation.