ALBERT

Notes: The effects of long-term CO2 enhancement and varying nutrient availability on photosynthesis and ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) were studied on loblolly pine (Pinus taeda L.) seedlings grown in two atmospheric CO2 partial pressures (35 and 65 Pa) and three nutrient treatments (low N, low P, and high N and P). Measurements taken in late autumn (November) after 2 years of CO2 enrichment and nutrient addition showed that photosynthetic rates were higher for plants grown at elevated CO2 only when they received supplemental N. Total rubisco activity and rubisco content decreased at elevated CO2, but there was an increase in activation state. At elevated CO2, proportionately less N was found in rubisco and more N was found in the light reaction components. These results demonstrate acclimation of photosynthetic processes to elevated CO2 through reallocation of N. Loblolly pine grown in nutrient conditions similar to native soils (low N availability) had lower needle N and chlorophyll content, lower total rubisco activity and content, and lower photosynthetic rates than plants grown at high N and P. This suggests that the magnitude of the photosynthetic response to a future, high-CO2 environment will be dependent on soil fertility in the system.

Notes: Atmospheric CO2 partial pressure may have been as low as 18 Pa during the Pleistocene and is expected to increase from 35 to 70 Pa before the end of the next century. Low CO2 reduces the growth and reproduction of C3 plants, whereas elevated CO2 often increases growth and reproduction. Plants at high elevation are exposed to reduced CO2 partial pressure and may be better adapted to the low CO2 of the Pleistocene. We examined genotypes of Arabidopsis thaliana from different elevations for variation in growth and reproduction at the CO2 levels of the Pleistocene, the present and the future. Genotypes exhibited limited genetic variation in the response of the production of biomass to changes in CO2, but showed significant variation in reproductive characters. We found evidence that plants from high elevations may be better adapted to low CO2 when considering seed number, which is an important component of fitness. Genotypes showed greater variation in the response of seed number between 35 and 20 Pa CO2 compared to 35 and 70 Pa CO2. We conclude that present-day C3 annuals may have greater potential for evolution in response to the low CO2 of the Pleistocene relative to the elevated CO2 predicted for the future.

Notes: We grew loblolly and ponderosa pine seedlings in a factorial experiment with two CO2 partial pressures (35 and 70 Pa), and two nitrogen treatments (1.0 and 3.5 mol m−3 NH4+), for one growing season to examine the effects of carbon and nitrogen availability on leaf construction cost. Growth in elevated CO2 reduced leaf nitrogen concentrations by 17 to 40%, and increased C:N by 22 to 68%. Elevated N availability increased leaf N concentrations and decreased C:N. Non-structural carbohydrates increased in high-CO2-grown loblolly seedlings, except in fascicles from low N, and in ponderosa primary and fascicle leaves grown in high N. In loblolly, increases in starch were nearly 2-fold greater than the increases in soluble sugars. In ponderosa, only the soluble sugars were affected by CO2. Leaf construction cost (g glucose g−1 dm) varied by 9.3% across all treatments. All of the variation in loblolly leaf construction cost could be explained by changes in non-structural carbohydrates. A model of the response of construction cost to changes in the mass of different biochemical fractions suggests that the remainder of the variation in ponderosa, not explained by non-structural carbohydrates, is probably attributable to changes in lignin, phenolic or protein concentrations.

Notes: Forest trees are major components of the terrestrial biome and their response to rising atmospheric CO2 plays a prominent role in the global carbon cycle. In this study, loblolly pine seedlings were planted in the field in recently disturbed soil of high fertility, and CO2 partial pressures were maintained at ambient CO2 (Amb) and elevated CO2 (Amb + 30 Pa) for 4 years. The objective of the study was to measure seasonal and long-term responses in growth and photosynthesis of loblolly pine exposed to elevated CO2 under ambient field conditions of precipitation, light, temperature and nutrient availability. Loblolly pine trees grown in elevated CO2 produced 90% more biomass after four growing seasons than did trees grown in ambient CO2. This large increase in final biomass was primarily due to a 217% increase in leaf area in the first growing season which resulted in much higher relative growth rates for trees grown in elevated CO2. Although there was not a sustained effect of elevated CO2 on relative growth rate after the first growing season, absolute production of biomass continued to increase each year in trees grown in elevated CO2 as a consequence of the compound interest effect of increased leaf area on the production of more new leaf area and more biomass. Allometric analyses of biomass allocation patterns demonstrated size-dependent shifts in allocation, but no direct effects of elevated CO2 on partitioning of biomass. Leaf photosynthetic rates were always higher in trees grown in elevated CO2, but these differences were greater in the summer (60–130% increase) than in the winter (14–44% increase), reflecting strong seasonal effects of temperature on photosynthesis. Our results suggest that seasonal variation in the relative photosynthetic response to elevated CO2 will occur in natural ecosystems, but total non-structural carbohydrate (TNC) levels in leaves indicate that this variation may not always be related to sink activity. Despite indications of canopy-level adjustments in carbon assimilation, enhanced levels of leaf photosynthesis coupled with increased total leaf area indicate that net carbon assimilation for the whole tree was greater for trees grown under elevated CO2 compared with ambient CO2. If the large growth enhancement observed in loblolly pine were maintained after canopy closure, then these trees could be a large sink for fossil carbon emitted to the atmosphere and produce a negative feedback on atmospheric CO2.

Notes: Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO2 partial pressures (pCO2) and with a CO2-enriched atmosphere (+ 30 Pa pCO2 compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO2. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO2-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO2 enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.

Notes: Abstract Steady-state labelling with 11CO2 was used to observe the blocking of phloem transport, induced by chilling short regions of stems or petioles of velvetleaf (Abutilon theophrasti Medic.) or cotton (Gossipium hirsutum L.). The abruptness of these blockages was evidenced by sharp decreases in 11C activity below, and increases above a 2 to 3 cm region cooled from 28°C to 18 or 13°C for periods as short as 2 min. Abrupt unblocking of transport in velvetleaf occurred a few minutes after rewarming, as evidenced by a sharp rise and and overshoot in 11C activity. Recovery of transport in cotton was more prolonged and was marked by occasional spontaneous blocking and unblocking of transport at various points along the petiole or stem, not necessarily in the cooled region. Similar spontaneous events were often observed in undisturbed cotton plants, but only rarely in velvetleaf.

Notes: Elevated atmospheric carbon dioxide partial pressures have been shown to have variable direct and indirect effects on plant respiration rates. In this study, growth, leaf respiration, and leaf nitrogen and carbohydrate partitioning were measured in Gossypium hirsutum L. grown in 35 and 65 Pa CO2 for 30d. Growth and maintenance coefficients of leaf respiration were estimated using gas exchange techniques both at night and during the day. Elevated CO2 stimulated biomass production (107%) and net photo-synthetic rates (35–50%). Total day-time respiration (Rd) was not significantly affected by growth CO2 partial pressure. However, night respiration (Rn) of leaves grown in 65 Pa CO2 was significantly greater than that of plants grown in 35 Pa CO2. Correlation of Rd and Rn with leaf expansion rates indicated that plants in both CO2 treatments had equivalent growth respiration coefficients but maintenance respiration was significantly greater in elevated CO2. Increased maintenance coefficients in elevated CO2 appeared to be related to increased starch accumulation rather than to changes in leaf nitrogen.

Notes: Cotton plants were grown in CO2-controlled growth chambers in atmospheres of either 35 or 65 Pa CO2. A widely accepted model of C3 leaf photosynthesis was parameterized for leaves from both CO2 treatments using non-linear least squares regression techniques, but in order to achieve reasonable fits, it was necessary to include a phosphate limitation resulting from inadequate triose phosphate utilization. Despite the accumulation of large amounts of starch (〉50 g m−2) in the high CO2 plants, the photosynthetic characteristics of leaves in both treatments were similar, although the maximum rate of Rubisco activity (Vcmax), estimated from A versus Ci response curves measured at 29°C, was ∼10% lower in leaves from plants grown in high CO2. The relationship between key model parameters and total leaf N was linear, the only difference between CO2 treatments being a slight reduction in the slope of the line relating Vcmax to leaf N in plants grown at high CO2. Stomatal conductance of leaves of plants grown and measured at 65 Pa CO2 was approximately 32% lower than that of plants grown and measured at 35 Pa. Because photosynthetic capacity of leaves grown in high CO2 was only slightly less than that of leaves grown in 35 Pa CO2, net photosynthesis measured at the growth CO2, light and temperature conditions was approximately 25% greater in leaves of plants grown in high CO2, despite the reduction in leaf conductance. Greater assimilation rate was one factor allowing plants grown in high CO2 to incorporate 30% more biomass during the first 36 d of growth.

Notes: The nitrogen requirement of plants is predominantly supplied by NH4+ and/or NO3− from the soil solution, but the energetic cost of uptake and assimilation is generally higher for NO3− than for NH4+. We found that CO2 enrichment of the atmosphere enhanced the root uptake capacity for NO3−, but not for NH4+, in field-grown loblolly pine saplings. Increased preference for NO3− at the elevated CO2 concentration was accompanied by increased carbohydrate levels in roots. The results have important implications for the potential consequences of global climate change on plant-and ecosystem-level processes in many temperate forest ecosystems.

Notes: Abstract Increasing atmospheric CO2 may result in alleviation of salinity stress in salt-sensitive plants. In order to assess the effect of enriched CO2 on salinity stress in Andropogon glomeratus, a C4 non-halophyte found in the higher regions of salt marshes, plants were grown at 350, 500, and 650 cm3 m−3 CO2 with 0 or 100 mol m−3 NaCl watering treatments. Increases in leaf area and biomass with increasing CO2 were measured in salt-stressed plants, while decreases in these same parameters were measured in non-salt-stressed plants. Tillering increased substantially with increasing CO2 in salt-stressed plants, resulting in the increased biomass. Six weeks following initiation of treatments, there was no difference in photosynthesis on a leaf area basis with increasing CO2 in salt-stressed plants, although short-term increases probably occurred. Stomatal conductance decreased with increasing CO2 in salt-stressed plants, resulting in higher water-use efficiency, and may have improved the diurnal water status of the plants. Concentrations of Na+ and Cl− were higher in salt stressed-plants while the converse was found for K +. There were no differences in leaf ion content between CO2 treatments in the salt-stressed plants. Decreases in photosynthesis in salt-stressed plants occurred primarily as a result of decreased internal (non-stomatal) conductance.