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.     

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).     

Carbon dioxide enrichment improves growth, water relations and survival of droughted honey mesquite (Prosopis glandulosa) seedlings

Low water availability reduces the establishment of the invasive shrub Prosopis on some grasslands. Water deficit survival and traits that may contribute to the postponement or tolerance of plant dehydration were measured on seedlings of P. glandulosa Torr. var. glandulosa (honey mesquite) grown at CO(2) concentrations of 370 (ambient), 710, and 1050 micro mol mol(-1). Because elevated CO(2) decreases stomatal conductance, the number of seedlings per container in the elevated CO(2) treatments was increased to ensure that soil water content was depleted at similar rates in all treatments. Seedlings grown at elevated CO(2) had a greater root biomass and a higher ratio of lateral root to total root biomass than those grown at ambient CO(2) concentration; however, these seedlings also shed more leaves and retained smaller leaves. These changes, together with a reduced transpiration/leaf area ratio at elevated CO(2), may have contributed to a slight increase in xylem pressure potentials of seedlings in the 1050 micro mol mol(-1) CO(2) treatment during the first 37 days of growth (0.26 to 0.40 MPa). Osmotic potential was not affected by CO(2) treatment. Increasing the CO(2) concentration to 710 and 1050 micro mol mol(-1) more than doubled the percentage survival of seedlings from which water was withheld for 65 days. Carbon dioxide enrichment significantly increased survival from 0% to about 40% among seedlings that experienced the lowest soil water content. By increasing seedling survival of drought, rising atmospheric CO(2) concentration may increase abundance of P. glandulosa on grasslands where low water availability limits its establishment.     

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.     

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 μmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.     

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.     

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.     

Carbohydrate reserve accumulation and depletion in Engelmann spruce (Picea engelmannii Parry): effects of cold storage and pre-storage CO(2) enrichment

The effects of pre-storage CO(2) enrichment on growth, non-structural carbohydrates and post-storage root growth potential of Engelmann spruce (Picea engelmannii Parry) seedlings were studied. Seedlings were grown from seed for 202 days in growth chambers with ambient (340 micro l l(-1)) or CO(2) enriched (1000 micro l l(-1)) air. Some seedlings were transferred between CO(2) treatments at 60 and 120 days. Photoperiod was reduced at 100 days to induce bud set and temperature was reduced at 180 days to promote frost hardiness development for storage at -5 degrees C for 2 or 4 months. Stored seedlings were planted in a growth chamber after thawing for one week at +5 degrees C. At 80, 120, 140 and 202 days, and at each planting time after storage, seedlings were harvested for growth measurements and analysis of starch and soluble sugar concentrations. Planted seedlings were assessed for bud break every two days and new roots > 5 mm long were counted after four weeks. Carbon dioxide enrichment increased root collar diameter and almost doubled seedling biomass, with the most obvious effects occurring after bud set. Stem height was affected only slightly and shoot/root ratios were not affected at all. Carbon dioxide enrichment increased the rate of reserve carbohydrate accumulation, but did not influence the final concentration attained before storage (accounting for 32% of seedling dry weight). Needles were the major storage organ for soluble sugars, whereas roots were the major storage organ for starch. Soluble sugars were not strongly affected by two or four months of storage, but starch was reduced by more than 50% in all plant parts. None of the CO(2) treatments had an impact on bud break or root growth potential.     

Movement of respiratory CO(2) in stems of loblolly pine (Pinus taeda L.) seedlings

Temperature-independent fluctuations in stem CO(2) efflux were measured in Pinus taeda L. seedlings. Stem CO(2) efflux was measured during high and low transpiration rates, high and low net photosynthesis rates, and normal and interrupted substrate supply conditions. Stem CO(2) efflux rates were an average of 6.7% lower during periods of high transpiration compared to periods of low transpiration. This difference in stem CO(2) efflux rates was not due to water stress. The most likely cause was movement of respiratory CO(2) in the transpiration stream. Interruption of substrate supply to the stem by phloem girdling reduced stem CO(2) efflux rates. Increasing net photosynthesis rates from low to high had no effect on stem CO(2) efflux, but decreasing net photosynthesis from high to low caused relatively small reductions in stem CO(2) efflux. These results indicate that diurnal changes in net photosynthesis rate may play a small role in temperature-independent afternoon depressions of stem CO(2) efflux. The transport of respiratory CO(2) by the transpiration stream compromises measurements of woody tissue respiration obtained by commonly accepted gas exchange techniques. This phenomenon could also affect measurement of leaf net photosynthesis and branch woody tissue respiration.     

Effects of ozone on growth and gas exchange of Eucalyptus globulus seedlings

To study the effects of a low concentration of ozone on growth and gas exchange in Eucalyptus globulus Labill. seedlings, ozone was applied for 37 days at a concentration of 50 ppb for 7 h daily under conditions of low light (250 micro mol m(-2) s(-1) (PAR)) and controlled temperature (20 degrees C). The seedlings exhibited extreme sensitivity to ozone. The ozone treatment reduced total plant biomass but had no effect on the partitioning of assimilate. Photosynthesis, stomatal conductance and internal CO(2) concentration were all reduced by ozone. The decline in photosynthesis was partly the result of direct effects of ozone on the stomata.     

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)].     

Interaction of nutrient limitation and elevated CO2 concentration on carbon assimilation of a tropical tree seedling (Cedrela odorata)

Carbon assimilation by Cedrela odorata L. (Meliaceae) seedlings was investigated in ambient and elevated CO2 concentrations ([CO2]) for 119 days, using small fumigation chambers. A solution containing macro- and micronutrients was supplied at two rates. The 5% rate (high rate) was designed to avoid nutrient limitation and allow a maximum rate of growth. The 1% rate (low rate) allowed examination of the effect of the nutrient limitation-elevated CO2 interaction on carbon assimilation. Root growth was stimulated by 23% in elevated [CO2] at a high rate of nutrient supply, but this did not lead to a change in the root:shoot ratio. Total biomass did not change at either rate of nutrient supply, despite an increase in relative growth rate at the low nutrient supply rate. Net assimilation rates and relative growth rates were stimulated by the high rate of nutrient addition, irrespective of [CO2]. We used a biochemical model of photosynthesis to investigate assimilation at the leaf level. Maximum rate of electron transport (Jmax) and maximum velocity of carboxylation (Vcmax) did not differ significantly with CO2 treatment, but showed a substantial reduction at the low rate of nutrient supply. Across both CO2 treatments, mean Jmax for seedlings grown at a high rate of nutrient supply was 75 micromol m(-2) s(-1) and mean Vcmax was 27 micromol m(-2) s(-1). The corresponding mean values for seedlings grown at a low rate of nutrient supply were 36 micromol m(-2) s(-1) and 15 micromol m(-2) s(-1), respectively. Concentrations of leaf nitrogen, on a mass basis, were significantly decreased by the low nutrient supply rate, in proportion to the observed decrease in photosynthetic parameters. Chlorophyll and carbohydrate concentrations of leaves were unaffected by growth [CO2]. Because there was no net increase in growth in response to elevated [CO2], despite increased assimilation of carbon at the leaf level, we hypothesize that the rate of respiration of non-photosynthetic organs was increased.     

Elevated atmospheric CO2 concentration alters the effect of phosphate supply on growth of Japanese red pine (Pinus densiflora) seedlings

We demonstrated that the inorganic phosphate (P(i)) requirement for growth of Japanese red pine (Pinus densiflora Sieb. & Zucc.) seedlings is increased by elevated CO(2) concentration ([CO(2)]) and that responses of the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker & Couch to P(i) supply are also altered. To investigate the growth response of non-mycorrhizal seedlings to P(i) supply in elevated [CO(2)], non-mycorrhizal seedlings were grown for 73 days in ambient or elevated [CO(2)] (350 or 700 micromol mol(-1)) with nutrient solutions containing one of seven phosphate concentrations (0, 0.02, 0.04, 0.06, 0.08, 0.10 and 0.20 mM). In ambient [CO(2)], the growth response to P(i) was saturated at about 0.1 mM P(i), whereas in elevated [CO(2)], the growth response to P(i) supply did not saturate, even at the highest P(i) supply (0.2 mM), indicating that the P(i) requirement is higher in elevated [CO(2)] than in ambient [CO(2)]. The increased requirement was due mainly to an altered shoot growth response to P(i) supply. The enhanced P(i) requirement in elevated [CO(2)] was not associated with a change in photosynthetic response to P(i) or a change in leaf phosphorus (P) status. We investigated the effect of P(i) supply (0.04, 0.08 and 0.20 mM) on the ectomycorrhizal fungus P. tinctorius in mycorrhizal seedlings grown in ambient or elevated [CO(2)]. Root ergosterol concentration (an indicator of fungal biomass) decreased with increasing P(i) supply in ambient [CO(2)], but the decrease was far less in elevated [CO(2)]. In ambient [CO(2)] the ratio of extramatrical mycelium to root biomass decreased with increasing P(i) supply but did not change in elevated [CO(2)]. We conclude that, because elevated [CO(2)] increased the P(i) requirement for shoot growth, the significance of the ectomycorrhizal association was also increased in elevated [CO(2)].     

Effects of elevated CO(2) on chloroplast components, gas exchange and growth of oak and cherry

Specific chloroplast proteins, gas exchange and dry matter production in oak (Quercus robur L.) seedlings and clonal cherry (Prunus avium L. x pseudocerasus Lind.) plants were measured during 19 months of growth in climate-controlled greenhouses at ambient (350 vpm) or elevated (700 vpm) CO(2). In both species, the elevated CO(2) treatment increased the PPFD saturated-rate of photosynthesis and dry matter production. After two months at elevated CO(2), Prunus plants showed significant increases in leaf (55%) and stem (61%) dry mass but not in root dry mass. However, this initial stimulation was not sustained: treatment differences in net assimilation rate (A) and plant dry mass were less after 10 months of growth than after 2 months of growth, suggesting acclimation of A to elevated CO(2) in Prunus. In contrast, after 10 months of growth at elevated CO(2), leaf dry mass of Quercus increased (130%) along with shoot (356%) and root (219%) dry mass, and A was also twice that of plants grown and measured at ambient CO(2). The amounts of Rubisco and the thylakoid-bound protein cytochrome f were higher in Quercus plants grown for 19 months in elevated CO(2) than in control plants, whereas in Prunus there was less Rubisco in plants grown for 19 months in elevated CO(2) than in control plants. Exposure to elevated CO(2) for 10 months resulted in increased mean leaf area in both species and increased abaxial stomatal density in Quercus. There was no change in leaf epidermal cell size in either species in response to the elevated CO(2) treatment. The lack of acclimation of photosynthesis in oak grown at elevated CO(2) is discussed in relation to the production and allocation of dry matter. We propose that differences in carbohydrate utilization underlie the differing long-term CO(2) responses of the two species.     

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].     

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.     

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

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

Growth 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.     

Influence of water stress on the physiology and growth of red spruce seedlings

Two-year-old, container-grown red spruce (Picea rubens Sarg.) seedlings from a New Hampshire seed source were exposed to 10 or 11 drying cycles in which the seedlings were not watered until their midday (1400 h) xylem water potentials averaged -1.57 MPa. Control seedlings were kept well watered to maintain midday water potentials of about -0.73 MPa. After the final drying cycle, the water-stressed seedlings were rehydrated and osmotic potentials were determined by pressure-volume analysis. Gas exchange at ambient CO(2) concentration (338 ppm) and at an elevated CO(2) concentration (838 ppm) was measured on both groups of plants as they slowly dried down. No osmotic adjustment or photosynthetic acclimation occurred as a result of the water-stress treatment and both groups of seedlings maintained photosynthesis to water potentials as low as -3.0 MPa. Twenty-four hours after rehydration, the water-stressed seedlings had photosynthetic rates as high as the control seedlings. Estimated stomatal limitation to photosynthesis was approximately 30% down to water potentials of -1.4 MPa, but increased steadily as water potentials decreased further. At ambient CO(2) concentrations (338 ppm) and water potentials averaging -2.45 MPa, photosynthetic rates of water-stressed seedlings were 15% those of well-watered seedlings, whereas when the same water-stressed seedlings were measured in the presence of an elevated concentration of CO(2) (838 ppm) their photosynthetic rates were 73% those of well-watered seedlings measured at an ambient CO(2) concentration (338 ppm).     

Effects of dessication on post-planting stress in bare-root Corsican pine seedlings

We examined the post-planting consequences of pre-planting exposure stress on two-year-old, bare-root Corsican pine (Pinus nigra Arnold. ssp. laricio var. Corsicana) seedlings. Seedlings were lifted from a nursery and exposed to ambient conditions for periods of up to 192 h before being planted in minirhizotrons. Exposure decreased seedling water potential, CO(2) assimilation rate, leaf conductance and new root elongation, and increased mortality after planting. During exposure, needle total nonstructural carbohydrates (TNC) concentration (expressed on a dry mass basis) decreased by 0.31 mg g(dm) (-1) h(-1); however, needle and root TNC concentrations remained high (> 100 mg g(dm) (-1)) at planting, even in those treatments leading to severe seedling mortality. More than 90% of the seedlings with predawn water potentials lower than -1.3 MPa at planting did not elongate new roots and did not survive, whereas a similar percentage of seedlings with a predawn water potential above this value at planting elongated new roots and survived, suggesting that this value corresponds to a turgor threshold below which new root formation is inhibited. At planting, embolization of xylem conduits in roots and shoots was low for seedlings in all of the exposure treatments.