Carbon isotope discrimination and stomatal responses of mature Pinus sylvestris L. trees exposed in situ for three years to elevated CO2 and temperature

The Climate Change Experiment (CLIMEX) is a unique large scale facility in which an entire undisturbed catchment of boreal vegetation has been exposed to elevated CO₂ (560 ppm) and temperature (+3°C summer, +5°C winter) for the past three years with all the soil-plant-atmosphere linkages intact. Here, carbon isotope composition and stomatal density have been analysed from sequential year classes of needles of mature Scots pine trees (Pinus sylvestris L.) to investigate the response of time-integrated water-use efficiency (UWE) and stomatal density to CO₂ enrichment and climate change. Carbon isotope discrimination decreased and WUE increased in cohorts of needles developing under increased CO₂ and temperature, compared to needles on the same trees developing in pretreatment years. Mid-season instantaneous gas exchange, measured on the same trees for the past four years, indicated that these responses resulted from higher needle photosynthetic rates and reduced stomatal conductance. Needles of P. sylvestris developing under increased CO₂ and temperature had consistently lower stomatal densities than their ambient grown counterparts on the same trees. The stomatal density of P. sylvestris needles was inversely correlated with δ¹³C-derived WUE, implying some effect of this morphological response on leaf gas exchange. Future atmospheric CO₂ and temperature increases are therefore likely to improve the water economy of P. sylvestris, at least at the scale of individual needles, by affecting stomatal density and gas exchange processes.     

Variable photosynthetic acclimation in consecutive cohorts of Scots pine needles during 3 years of growth at elevated CO2 and elevated temperature

In this experiment, the photosynthetic acclimation of successive needle cohorts of Scots pine were studied during 3 years of growth at elevated CO₂ and temperature. Naturally regenerated Scots pine (Pinus sylvestris L.) trees were subjected to elevated CO₂ concentration (+CO₂, 700 p.p.m), elevated temperature (+T, ambient +2 to +6 °C) and to a combination of elevated CO₂ and temperature (+CO₂ + T) in closed‐top chambers, starting in August 1996. Trees growing in chambers with ambient CO₂ and ambient temperature served as controls (AmbC). Elevated CO₂ influenced the dark reactions more than the light reactions of photosynthesis, as in the 1996 and 1997 cohorts the carboxylation capacity of Rubisco was reduced in the first and second year of exposure, but there was no consistent change in chlorophyll fluorescence. Net photosynthesis measured at growth concentration of CO₂ was higher at +CO₂ than at AmbC on only one measuring occasion, was generally lower at +T and was not changed at +CO₂ + T. However, trees grown at +T tended to invest more nitrogen (N) in Rubisco, as Rubisco/chlorophyll and the proportion of the total needle N bound to Rubisco occasionally increased. The interaction of +CO₂ and +T on Rubisco was mostly negative; consequently, in the second and third year of the experiment the carboxylation capacity decreased at +CO₂ + T. In the 1996, 1997 and 1998 cohorts, the structural N concentration of needles was lower at +CO₂ than at AmbC. Elevated CO₂ and elevated temperature generally had a positive interaction on N concentration; consequently, N concentration in needles decreased less at +CO₂ + T than at +CO₂. At +CO₂ + T, the acclimation response of needles varied between years and was more pronounced in the 1‐year‐old needles of the 1997 cohort than in those of the 1998 cohort. Thus, acclimation was not always greater in 1‐year‐old needles than in current‐year needles. In the +CO₂ + T treatment, elevated temperature had a greater effect on acclimation of needles than elevated CO₂.     

Effects of elevated O3 and CO2 concentrations on photosynthesis and stomatal conductance in Scots pine

Naturally regenerated Scots pines (Pinus sylvestris L.), aged 28–30 years old, were grown in open-top chambers and subjected in situ to three ozone (O₃) regimes, two concentrations of CO₂, and a combination of O₃ and CO₂ treatments From 15 April to 15 September for two growing seasons (1994 and 1995). The gas exchanges of current-year and 1-year-old shoots were measured, along with the nitrogen content of needles. In order to investigate the factors underlying modifications in photosynthesis, five parameters linked to photosynthetic performance and three to stomatal conductance were determined. Elevated O₃ concentrations led to a significant decline in the CO₂ compensation point (Г^*), maximum RuP₂-saturated rate of carboxylation (V_cmax), maximum rate of electron transport (J_max), maximum stomatal conductance (g_smax), and sensitivity of stomatal conductance to changes in leaf-to-air vapour pressure difference (?g_s/?D_v) in both shoot-age classes. However, the effect of elevated O₃ concentrations on the respiration rate in light (R_d) was dependent on shoot age. Elevated CO₂(700 μmol mol^?1) significantly decreased J_max and g_smax but increased R_d in 1-year-old shoots and the ?g_s/?D_v in both shoot-age classes. The interactive effects of O₃ and CO₂ on some key parameters (e.g. V_cmax and J_max) were significant. This may be closely related to regulation of the maximum stomatal conductance and stomatal sensitivity induced by elevated CO₂. As a consequence, the injury induced by O₃ was reduced through decreased ozone uptake in 1-year-old shoots, but not in the current-year shoots. Compared to ambient O₃ concentration, reduced O₃ concentrations (charcoal-filtered air) did not lead to significant changes in any of the measured parameters. Compared to the control treatment, calculations showed that elevated O₃ concentrations decreased the apparent quantum yield by 15% and by 18%, and the maximum rate of photosynthesis by 21% and by 29% in the current-year and 1-year-old shoots, respectively. Changes in the nitrogen content of needles resulting from the various treatments were associated with modifications in photosynthetic components.     

The effects on Arbutus unedo L. of long-term exposure to elevated CO2

Arbutus unedo is a sclerophyllous evergreen, characteristic of Mediterranean coastal scrub vegetation. In Italy, trees of A. unedo have been found close to natural CO₂ vents where the mean atmospheric carbon dioxide concentration is about 2200 μmol mol^?1. Comparisons were made between trees growing in elevated and ambient CO₂ concentrations to test for evidence of adaptation to long-term exposure to elevated CO₂. Leaves formed at elevated CO₂ have a lower stomatal density and stomatal index and higher specific leaf area than those formed at ambient CO₂, but there was no change in carbon to nitrogen ratios of the leaf tissue. Stomatal conductance was lower at elevated CO₂ during rapid growth in the spring. In mid-summer, under drought stress, stomatal closure of all leaves occurred and in the autumn, when stress was relieved, the conductance of leaves at both elevated and ambient CO₂ increased. In the spring, the stomatal conductance of the new flush of leaves at ambient CO₂ was higher than the leaves at elevated CO₂, increasing instantaneous water use efficiency at elevated CO₂. Chlorophyll fluorescence measurements suggested that elevated CO₂ provided some protection against photoinhibition in mid-summer. Analysis of A/C_i curves showed that there was no evidence of either upward or downward regulation of photosynthesis at elevated CO₂. It is therefore anticipated that A. unedo will have higher growth rates as the ambient CO₂ concentrations increase.     

The influence of elevated O3 and CO2 concentrations on secondary metabolites of Scots pine (Pinus sylvestris L. ) seedlings

Terpene, resin acid and total phenolic concentrations in five‐year‐old Scots pine (Pinus sylvestris L.) seedlings were analysed after exposure to ambient and realistically elevated (2 × ambient) O₃ and CO₂ concentrations and their combination in open‐top chambers during two growing seasons. Under O₃ exposure, limonene concentration in needles and isopimaric concentration in stems decreased significantly. As a response to elevated CO₂, α‐pinene and total phenolic concentrations in needles increased significantly, while bornyl acetate concentration in needles and palustric + levopimaric and neoabietic acid concentrations in stems decreased significantly. Some terpenes and resin acids were found at lower concentrations in the combined O₃ and CO₂ treatment than in O₃ exposure or elevated CO₂. A negative chamber effect was found: seedlings growing inside the chambers with ambient air had significantly lower concentrations of some terpenes and resin acids than seedlings growing outside the chambers. There was a lot of between‐tree variation in terpene and resin acid concentrations, which is typical of open‐pollinated populations. The results of this study suggest that, at least in short‐term experiments, Scots pine secondary metabolites are relatively insensitive to climate change factors. Total phenolics in the needles were the most responsive group showing about 25% increase in elevated CO₂, and O₃ exposure did not mitigate this CO₂ effect. Terpenes and resin acids were less responsive, although some individual compounds showed notable responses, e.g. α‐pinene in needles, which increased about 50% in response to elevated CO₂. As a consequence, although there were only slight effects on total pools of needle secondary metabolites, considerable O₃ and CO₂ effects on certain individual compounds might have ecological significance via trophic amplification, e.g. in decomposing processes of needle litter.     

Elevated CO2 and temperature alter nitrogen allocation in Douglas-fir

The effects of elevated CO₂ and temperature on principal carbon constituents (PCC) and C and N allocation between needle, woody (stem and branches) and root tissue of Pseudotsuga menziesii Mirb. Franco seedlings were determined. The seedlings were grown in sun‐lit controlled‐environment chambers that contained a native soil. Chambers were controlled to reproduce ambient or ambient +180 ppm CO₂ and either ambient temperature or ambient +3.5 °C for 4 years. There were no significant CO₂ × temperature interactions; consequently the data are presented for the CO₂ and temperature effects. At the final harvest, elevated CO₂ decreased the nonpolar fraction of the PCC and increased the polar fraction and amount of sugars in the needles. In contrast, elevated temperature increased the nonpolar fraction of the PCC and decreased sugars in needles. There were no CO₂ or temperature effects on the PCC fractions in the woody tissue or root tissue. Elevated CO₂ and temperature had no significant effects on the C content of any of the plant tissues or fractions. In contrast, the foliar N content declined under elevated CO₂ and increased under elevated temperature; there were no significant effects in other tissues. The changes in the foliar N concentrations were in the cellulose and lignin fractions, the fractions, which contain protein, and are the consequences of changes in N allocation under the treatments. These results indicate reallocation of N among plant organs to optimize C assimilation, which is mediated via changes in the selectivity of Rubisco and carbohydrate modulation of gene expression.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Effects of carbon dioxide enrichment on response of mycorrhizal pitch pine (Pinus rigida) to aluminum: growth and mineral nutrition

 Carbon dioxide enrichment may increase the Al tolerance of trees by increasing root growth, root exudation and/or mycorrhizal colonization. The effect of elevated CO₂ on the response of mycorrhizal pitch pine (Pinus rigida Mill.) seedlings to Al was determined in two experiments with different levels of nutrients, 0.1- or 0.2-strength Clark solution. During each experiment, seedlings inoculated with the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker & Couch were grown 13 weeks in sand irrigated with nutrient solution (pH 3.8) containing 0, 6.25, 12.5, or 25 mg/l Al (0, 232, 463, or 927 μM Al) in growth chambers fumigated with 350 (ambient) or 700 (elevated) μl/l CO₂. At ambient CO₂, in the absence of Al, mean total dry weights (DW) of seedlings at the high nutrient level were 164% higher than those at the low level. Total DW at elevated CO₂, in the absence of Al, was significantly greater than that in ambient CO₂ at the low (+34%) and high (+16%) nutrient levels. Root and shoot DW at both nutrient levels decreased with increasing Al concentrations with Al reducing root growth more than shoot growth. Although visible symptoms of Al toxicity in roots and needles were reduced by CO₂ enrichment, there were no significant CO₂ × Al interactions for shoot or root DW. The percentage of seedling roots that became mycorrhizal was negatively related to nutrient level and was greater at elevated than at ambient CO₂ levels. Generally, elevated CO₂ had little effect on concentration of mineral nutrients in roots and needles. Aluminum reduced concentrations of most nutrients by inhibiting uptake. Received: 18 June 1997 / Accepted: 8 December 1997     

Photosynthetic acclimation to long-term exposure to elevated CO2 concentration in Pinus radiata D. Don. is related to age of needles

The effects of CO₂ enrichment on photosynthesis and ribulose-1,5-bisphosphate carboxylase/ oxygenase (Rubisco) in current year and 1-year-old needles on the same branch were studied on Pinus radiata D. Don. trees growing for 4 years in large, open-top chambers at ambient (36 Pa) and elevated (65 Pa) CO₂ partial pressures. At this age trees were 3·5–4 m tall. Measurements made late in the growing cycle (March) showed that photosynthetic rates at the growth CO₂ concentration [(pCO₂)_a] were lower in 1-year-old needles of trees grown at elevated CO₂ concentrations than in those of trees grown at ambient (pCO₂)_a. At elevated CO₂ concentrations V_cmax (maximum carboxylation rate) was reduced by 13% and J_max (RuBP regeneration capacity mediated by maximum electron transport rate) by 17%. This corresponded with photosynthetic rates at the growth (pCO₂)_a of 4·68 ± 0·41 μmol m^–2 s^–1 and 6·15 ± 0·46 μmol m^–2 s^–1 at 36 and 65 Pa, respectively (an enhancement of 31%). In current year needles photosynthetic rates at the growth (pCO₂)_a were 6·2 ± 0·72 μmol m^–2 s^–1 at 36 Pa and 10·15 ± 0·64 μmol m^–2 s^–1 at 65 Pa (an enhancement of 63%). The smaller enhancement of photosynthesis in 1-year-old needles at 65 Pa was accompanied by a reduction in Rubisco activity (39%) and content (40%) compared with that at 36 Pa. Starch and sugar concentrations in 1-year-old needles were not significantly different in the CO₂ treatments. There was no evidence in biochemical parameters for down-regulation at elevated (pCO₂)_a in fully fexpanded needles of the current year cohort. These data show that enhancement of photosynthesis continues to occur in needles after 4 years’ exposure to elevated CO₂ concentrations. Photosynthetic acclimation reduces the degree of this enhancement, but only in needles after 1 year of growth. Thus, responses to elevated CO₂ concentration change during the lifetime of needles, and acclimation may not be apparent in current year needles. This transitory effect is most probably attributable to the effects of developmental stage and proximity to actively growing shoots on sink strength for carbohydrates. The implications of such age-dependent responses are that older trees, in which the contribution of older needles to the photosynthetic biomass is greater than in younger trees, may become progressively more acclimated to elevated CO₂ concentration.     

Decomposition of secondary compounds from needle litter of Scots pine grown under elevated CO2 and O3

Elevated atmospheric carbon dioxide (CO₂) and ozone (O₃) concentrations have both been shown to affect plant tissue quality, which in turn could affect litter decomposition and carbon (C) and nutrient cycling. In order to evaluate effects of climate change on litter chemistry, needle litter was collected from Scots pine (Pinus sylvestris L.) saplings exposed to elevated CO₂ or O₃ concentration and their combination over three growing seasons in open‐top chambers. The decomposition of needle litter was followed for 19 months in a pine forest. During decomposition, needle samples for secondary compound analysis were collected and the mass loss of needles was followed. Main nutrients and total phenolics were analysed from litter in the beginning and at the end of the experiment. After 19‐month decomposition, the accumulated mass loss was about 34%; however, no significant differences were found in decomposition rates of needle litter between various treatments. Concentrations of total monoterpenes were about 4%, total resin acids 21% and total phenolics 14% of the initial concentrations in litter after 19‐month decomposition. In the beginning of litter decomposition, concentrations of individual monoterpenes –α‐pinene and β‐pinene – were significantly higher in needle litter grown under elevated CO₂. However, concentrations of total monoterpenes during the whole decomposition period were not significantly affected by CO₂ or O₃ treatments. Concentrations of some individual and total resin acids were higher in needle litter grown under elevated CO₂ or O₃ than under ambient air. There were no significant differences in concentrations of total phenolics as well as nitrogen (N) and the main nutrient concentrations between treatments during decomposition. High concentrations of monoterpenes and resin acids in needles might slightly delay C recycling in forest soils. It is concluded that elevated CO₂ and O₃ concentrations do not have remarkable impacts on litter decomposition processes in Scots pine forests.     

Effects of climate change on labile and structural carbon in Douglas-fir needles as estimated by δ13C and Carea measurements

Models of photosynthesis, respiration, and export predict that foliar labile carbon (C) should increase with elevated CO₂ but decrease with elevated temperature. Sugars, starch, and protein can be compared between treatments, but these compounds make up only a fraction of the total labile pool. Moreover, it is difficult to assess the turnover of labile carbon between years for evergreen foliage. Here, we combined changes in foliar C_area (C concentration on an areal basis) as needles aged with changes in foliar isotopic composition (δ¹³C) caused by inputs of ¹³C‐depleted CO₂ to estimate labile and structural C in needles of different ages in a four‐year, closed‐chamber mesocosm experiment in which Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were exposed to elevated temperature (ambient + 3.5 °C) and CO₂ (ambient + 179 ppm). Declines in δ ¹³C of needle cohorts as they aged indicated incorporation of newly fixed labile or structural carbon. The δ ¹³C calculations showed that new C was 41 ± 2% and 28 ± 3% of total needle carbon in second‐ and third‐year needles, respectively, with higher proportions of new C in elevated than ambient CO₂ chambers (e.g. 42 ± 2% vs. 37 ± 6%, respectively, for second‐year needles). Relative to ambient CO₂, elevated CO₂ increased labile C in both first‐ and second‐year needles. Relative to ambient temperature, elevated temperature diminished labile C in second‐year needles but not in first‐year needles, perhaps because of differences in sink strength between the two needle age classes. We hypothesize that plant‐soil feedbacks on nitrogen supply contributed to higher photosynthetic rates under elevated temperatures that partly compensated for higher turnover rates of labile C. Strong positive correlations between labile C and sugar concentrations suggested that labile C was primarily determined by carbohydrates. Labile C was negatively correlated with concentrations of cellulose and protein. Elevated temperature increased foliar %C, possibly due to a shift of labile constituents from low %C carbohydrates to relatively high %C protein. Decreased sugar concentrations and increased nitrogen concentrations with elevated temperature were consistent with this explanation. Because foliar constituents that vary in isotopic signature also vary in concentrations with leaf age or environmental conditions, inferences of c_i/c_a values from δ ¹³C of bulk leaf tissue should be done cautiously. Tracing of ¹³C through foliar carbon pools may provide new insight into foliar C constituents and turnover.     

Effects of elevated CO2 on leaf gas exchange in beech and oak at two levels of nutrient supply: consequences for sensitivity to drought in beech

Beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) were grown from seed for two whole seasons at two CO₂ concentrations (ambient and ambient + 250 μmol mol^?1) with two levels of soil nutrient supply. Measurements of net leaf photosynthetic rate (A) and stomatal conductance (g_s) of well-watered plants were taken over both seasons; a drought treatment was applied in the middle of the second growing season to a separate sample of beech drawn from the same population. The net leaf photosynthetic rate of well-watered plants was stimulated in elevated CO₂ by an average of 75% in beech and 33% in oak; the effect continued through both growing seasons at both nutrient levels. There were no interactive effects of CO₂ concentration and nutrient level on A or g_s in beech or oak. Stomatal conductance was reduced in elevated CO₂ by an average of 34% in oak, but in beech there were no significant reductions in g_s except under cloudy conditions (–22% in elevated CO₂). During drought, there was no effect of CO₂ concentration on g_s in beech grown with high nutrients, but for beech grown with low nutrients, g_s was significantly higher in elevated CO₂, causing more rapid soil drying. With high nutrient supply, soil drying was more rapid at elevated CO₂ due to increased leaf area. It appears that beech may substantially increase whole-plant water consumption in elevated CO₂, especially under conditions of high temperature and irradiance when damage due to high evaporative demand is most likely to occur, thereby putting itself at risk during periods of drought.     

Effects of carbon dioxide concentration on the interactive effects of temperature and water vapour on stomatal conductance in soybean

Soybeans were grown at three CO₂ concentrations in outdoor growth chambers and at two concentrations in controlled-environment growth chambers to investigate the interactive effects of CO₂, temperature and leaf-to-air vapour pressure difference (LAVPD) on stomatal conductance. The decline in stomatal conductance with CO₂ was a function of both leaf temperature and LAVPD. In the field measurements, stomatal conductance was more sensitive to LAVPD at low CO₂ at 30 °C but not at 35 °C. There was also a direct increase in conductance with temperature, which was greater at the two elevated carbon dioxide concentrations. Environmental growth chamber results showed that the relative stomatal sensitivity to LAVPD decreased with both leaf temperature and CO₂. Measurements in the environmental growth chamber were also performed at the opposing CO₂, and these experiments indicate that the stomatal sensitivity to LAVPD was determined more by growth CO₂ than by measurement CO₂. Two models that describe stomatal responses to LAVPD were compared with the outdoor data to evaluate whether these models described adequately the interactive effects of CO₂, LAVPD and temperature.     

Water Relations of Cotton Plants under Nitrogen Deficiency: III. STOMATAL CONDUCTANCE, PHOTOSYNTHESIS, AND ABSCISIC ACID ACCUMULATION DURING DROUGHT

Nitrogen nutrition exerted a strong effect on stomatal sensitivity to water stress in cotton. In well-watered plants grown with 0.31 millimolar N in the nutrient solution, stomata closed at a water potential of -9 bars even though the wilting point was below -15 bars. For each doubling of nutrient N level, the water potential for stomatal closure decreased by about 2 bars. Elevated intercellular CO₂ concentrations caused only slight stomatal closure regardless of N nutrition. Exogenous abscisic acid (ABA) greatly increased stomatal sensitivity to elevated CO₂ concentrations.     

Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass

Native tallgrass prairie in NE Kansas was exposed to elevated (twice ambient) or ambient atmospheric CO₂ levels in open-top chambers. Within chambers or in adjacent unchambered plots, the dominant C₄ grass, Andropogon gerardii, was subjected to fluctuations in sunlight similar to that produced by clouds or within canopy shading (full sun > 1500 μmol m⁻² s⁻¹ versus 350 μmol m⁻² s⁻¹ shade) and responses in gas exchange were measured. These field experiments demonstrated that stomatal conductance in A. gerardii achieved new steady state levels more rapidly after abrupt changes in sunlight at elevated CO₂ when compared to plants at ambient CO₂. This was due primarily to the 50% reduction in stomatal conductance at elevated CO₂, but was also a result of more rapid stomatal responses. Time constants describing stomatal responses were significantly reduced (29–33%) at elevated CO₂. As a result, water loss was decreased by as much as 57% (6.5% due to more rapid stomatal responses). Concurrent increases in leaf xylem pressure potential during periods of sunlight variability provided additional evidence that more rapid stomatal responses at elevated CO₂ enhanced plant water status. CO₂-induced alterations in the kinetics of stomatal responses to variable sunlight will likely enhance direct effects of elevated CO₂ on plant water relations in all ecosystems.     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

Variations of stomatal density and carbon isotope values of<Emphasis Type="Italic"> Picea crassifolia</Emphasis> at different altitudes in the Qilian Mountains

The stomatal characteristics, length and dry weight as well as stable carbon isotope composition (i¹³C) of needles and tree rings of Qinghai spruce (Picea crassifolia) at different altitudes in the Qilian mountains were investigated. The results showed that stomatal density, distribution pattern on epidermis, and length and dry weight of needles all increased with altitude below 3,000 m. In contrast, these parameters all decreased with increasing altitude above 3,000 m. Furthermore an assay of tree rings showed that near 3,000 m in altitude was the optimum zone for growth and development of Qinghai spruce. Although atmospheric CO₂ concentration influences stomatal density, the effects of many environmental factors, such as temperature, light and rainfall, could obscure the effects of changes in atmospheric CO₂ concentration on stomatal density at altitudes higher than the optimum for growth. The correlation of stomatal density with atmospheric CO₂ concentration and i¹³C of Qinghai spruce needles was significant below 3,000 m, but was insignificant above 3,000 m altitude. The i¹³C value of needles, however, was negatively correlated with atmospheric CO₂ concentrations. Therefore, the influence of altitude should be considered in research on plant physiological ecology and the relationship of stomatal density with i¹³C value or atmospheric CO₂ concentration.     

The effects of elevated CO2 concentrations on the root growth of Lolium perenne and Trifolium repens grown in a FACE* system

Lolium perenne and Trifolium repens were grown in a Free Air CO₂ Enrichment (FACE) system at elevated (600 μimol mol-¹) and ambient (340 μmol mol-¹) carbon dioxide concentrations during a whole growing season. Using a root ingrowth bag technique the extent to which CO₂ enrichment influenced the growth of L, perenne and T. repens roots under two contrasting nutrient regimes was examined. Root ingrowth bags were inserted for a fixed time into the soil in order to trap roots. It was also possible to follow the mortality of roots in bags inserted for different time intervals. Root ingrowth of both L. perenne and T. repens increased under elevated CO₂ conditions. In L. perenne, root ingrowth decreased with increasing nutrient fertilizer level, but for T. repens the root ingrowth was not affected by the nutrient application rate. Besides biomass measurements, root length estimates were made for T, repens. These showed an increase under elevated CO₂ concentrations. Root decomposition appeared to decrease under elevated CO₂ concentrations. A possible explanation for this effect is the observed changes in tissue composition, such as the increase in the carbon: nitrogen ratio in roots of L. perenne at elevated CO₂ concentrations.     

Recent Bryological Literature

Abstract

Growing Leptobryum pyriforme (Hedw.) Wilson under ten times ambient atmospheric CO₂ concentrations showed that, while stomatal density and index decreased under elevated atmospheric CO₂, the overall number of stomata per spore capsule remained unaltered. Capsule length increased, while the size of stomata and epidermal cells decreased. This study suggests that stomatal frequency measures used to investigate the influence of varying levels of atmospheric CO₂ on angiosperms and gymnosperms may result in misleading conclusions when applied to bryophytes if considered in isolation from growth responses.     

Photosynthetic response of Tempranillo grapevine to climate change scenarios

Photosynthesis in C₃ plants is CO₂ limited and therefore any increase in Rubisco carboxylation substrate may increase net CO₂ fixation, unless plants experience acclimation or other limitations. These aspects are largely unexplored in grapevine. Photosynthesis analysis was used to assess the stomatal, mesophyll, photochemical and biochemical contributions to the decreasing photosynthesis observed in Tempranillo grapevines (Vitis vinifera) from veraison to ripeness, modulated by CO₂, temperature and water availability. Photosynthesis and photosystem II photochemistry decreased from veraison to ripeness. The elevated CO₂ and temperature increased photosynthesis, but transiently, in both well irrigated (WI) and water‐stressed plants. Photosynthetic rates were maxima 1 week after the start of elevated CO₂ and temperature treatments, but differences with treatments of ambient conditions disappeared with time. There were not marked changes in leaf water status, leaf chlorophyll or leaf protein that could limit photosynthesis at ripeness. Leaf total soluble sugars remained at ripeness as high as 2 weeks after the start of treatments. On the other hand, and as expected, CO₂ diffusional limitations impaired photosynthesis in grapevine plants grown under water scarcity, stomatal and mesophyll conductances to CO₂ decreased and in turn low chloroplastic CO₂ concentrations limited photosynthetic CO₂ fixation. In summary, photochemistry and photosynthesis from veraison to ripeness in Tempranillo grapevine were dominated by a developmental‐related decreasing trend that was only transiently influenced by elevated CO₂ concentrations.