Altered physiological and growth responses to elevated [CO2] in offspring from holm oak (Quercus ilex L.) mother trees with lifetime exposure to naturally elevated [CO2]

An important question with respect to plant performance in future climatic scenarios is whether the offspring of mature trees that have experienced lifelong exposure to elevated [CO₂] show altered physiological responses to elevated [CO₂] compared with those originating from current ambient CO₂ concentrations. To investigate this question, acorns were collected from two seed sources, denoted as ‘control’ and ‘spring’, from Quercus ilex mother trees grown at ambient (36 Pa) and at about twice ambient CO₂ concentrations, respectively, close to a natural CO₂ spring, Laiatico, central Italy. The seedlings were raised for 8 months under controlled conditions at ambient and elevated [CO₂] in a reciprocal experimental design and were used for the determination of biomass, photosynthesis and foliar carbohydrate concentrations, as well as the accumulation of structural biomass and lignin during leaf maturation. Under ambient [CO₂], biomass and foliar carbon acquisition in control progeny were not significantly different from spring progeny. However, under elevated [CO₂], spring seedlings showed less CO₂ acclimation than control seedlings but no significant differences in non‐structural carbohydrate concentrations and structural biomass per unit leaf dry mass. Developmental lignin accumulation in leaves was delayed under elevated [CO₂] compared with ambient [CO₂], but only in control progeny. Under elevated [CO₂], whole‐plant biomass, leaf area and stem diameter were significantly increased in Quercus ilex seedlings from both seed sources but with a higher stimulation of above‐ground biomass in spring than in control seedlings and a higher stimulation of below‐ground biomass in control seedlings. These results indicate that life history and/or progeny may determine the species‐specific CO₂ response and suggest that positive CO₂ acclimation is possible.     

Atmospheric CO2 enrichment increases growth and photosynthesis of Pinus taeda: a 4 year experiment in the field

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.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Carbon balance of young birch trees grown in ambient and elevated atmospheric CO2 concentrations

We constructed a carbon budget for young birch trees grown in ambient and elevated CO₂ concentrations over their fourth year of growth. The annual total of net leaf photosynthesis was 110% more in elevated CO₂ than in ambient CO₂. However, the trees in elevated CO₂ grew only 59% more biomass than the trees in ambient CO₂ over the year. Modelling studies showed that larger loss of carbon from fine-root production and growth of the root-associated mycorrhiza by the trees in elevated CO₂ probably accounted for all the remaining difference in net photosynthesis between the two treatments. Our modelling also showed that the fraction of net photosynthate consumed by respiration of nonleaf tissue was similar in the two CO₂ treatments, and was 26% and 24% for trees in ambient and elevated CO₂, respectively. Trees in elevated CO₂ had 43% more leaves, and produced 110% more net photosynthate than trees in ambient CO₂, even though the maximum rate of carboxylation per unit leaf nitrogen decreased by 21%. Sensitivity studies showed that down-regulation reduced the annual net photosynthetic production of the trees in elevated CO₂ by only 6%. Direct effects of higher CO₂ on photosynthesis and greater leaf area of the trees in elevated CO₂ increased the net photosynthesis of the trees by 68% and 60%, respectively; and together accounted for most of the difference in net photosynthesis between the two treatments.     

Long-term response of Sitka spruce (Picea sitchensis (Bong.) Carr.) to CO2 enrichment and nitrogen supply. I. Growth,biomass allocation and physiology

ABSTRACT

After a 3-year exposure to elevated CO₂, young trees of Sitka spruce (Picea sitchensis (Bong.) Carr.) were planted in native, nutrient-deficient forest soil and grown for two more years with three CO₂ treatments in open-top chambers, and with two nutrient treatments (with and without supplied N). Elevated CO₂ resulted in larger fresh mass, dry mass, leaf area and leaf thickness in two-year old needles, but had no effect on one-year old and current needles. Tree height, basal diameter and biomass production also increased, regardless of N supply. In trees without added N, elevated CO₂ resulted in higher root-to-shoot and absorbing roots-to-stump ratios. Regardless of N supply, trees grown in elevated CO₂ had lower photosynthetic rates on a leaf area basis. Photosynthesis reduction was accompanied by a decline in Rubisco activity and leaf N concentration. Under elevated CO₂, added N elevated photosynthesis and Rubisco activity, suggesting a dependence on N availability of the photosynthetic response to elevated CO₂. Stomatal conductance of trees grown with added N decreased in response to elevated CO₂. This may account for the larger reduction in intercellular CO₂ concentration, and hence photosynthesis, in the trees supplied with N than in those without N supply.     

A meta-analysis of leaf gas exchange and nitrogen in trees grown under elevated carbon dioxide

The response of trees to rising atmospheric CO₂ concentration ([CO₂]) is of concern to forest ecologists and global carbon modellers and is the focus of an increasing body of research work. I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or specific leaf area (SLA) in woody plants grown at <400 μmol mol^?1 CO₂ or at 600–800 μmol mol^?1 CO₂. The resulting data from 41 species were categorized according to growth conditions (unstressed versus stressed), length of CO₂ exposure, pot size and exposure facility [growth chamber (GC), greenhouse (GH), or open-top chamber (OTC)] and interpreted using meta-analytic methods. Overall, A showed a large and significant increase at elevated [CO₂] but length of CO₂ exposure and the exposure facility were important modifiers of this response. Plants exposed for < 50 d had a significantly greater response, and those from GCs had a significantly lower response than plants from longer exposures or from OTC studies. Negative acclimation of A was significant and general among stressed plants, but in unstressed plants was influenced by length of CO₂ exposure, the exposure facility and/or pot size. Growth at elevated [CO₂] resulted in moderate reductions in gs in unstressed plants, but there was no significant effect of CO₂ on gs in stressed plants. Leaf dark respiration (mass or area basis) was reduced strongly by growth at high [CO₂] > while leaf N was reduced only when expressed on a mass basis. This review is the first meta-analysis of elevated CO₂ studies and provides statistical confirmation of several general responses of trees to elevated [CO₂]. It also highlights important areas of continued uncertainty in our understanding of these responses.     

Effects of atmospheric CO2 enrichment on early growth of Vivia faba, a plant with large cotyledons

Seedlings of Vicia faba L. were grown in open-top growth chambers at present (P=350μmol^?1) and at elevated (E=700μmol mol^?1) atmospheric CO₂ concentration. The effects of CO₂ enrichment on the first phase of growth after germination were examined over 45 d. There were no positive effects of CO₂ enrichment on growth of the seedlings during this early phase. No differences were observed in leaf area or in total dry weight. No differences were found in morphology or anatomy of the leaves. The numbers of stomatal and epidermal cells, thickness of leaf, of epidermis and of mesophyll cell-layers were unaffected by CO₂ enrichment. Also, no differences were observed in leaf concentrations of chlorophyll, reducing carbohydrates or starch. These results contrast markedly with results from similar experiments on poplar hybrids and Phaseolus vulgaris obtained in the same growth facility. It seems that the intitial growth is under internal control such that the atmospheric CO₂ concentration has no effects. The lack of response in this case may be attributed to the presence and longevity of the large cotyledons which provided available substrate for growth.     

No evidence that elevated CO2 gives tropical lianas an advantage over tropical trees

Recent studies indicate that lianas are increasing in size and abundance relative to trees in neotropical forests. As a result, forest dynamics and carbon balance may be altered through liana‐induced suppression of tree growth and increases in tree mortality. Increasing atmospheric CO₂ is hypothesized to be responsible for the increase in neotropical lianas, yet no study has directly compared the relative response of tropical lianas and trees to elevated CO₂. We explicitly tested whether tropical lianas had a larger response to elevated CO₂ than co‐occurring tropical trees and whether seasonal drought alters the response of either growth form. In two experiments conducted in central Panama, one spanning both wet and dry seasons and one restricted to the dry season, we grew liana (n = 12) and tree (n = 10) species in open‐top growth chambers maintained at ambient or twice‐ambient CO₂ levels. Seedlings of eight individuals (four lianas, four trees) were grown in the ground in each chamber for at least 3 months during each season. We found that both liana and tree seedlings had a significant and positive response to elevated CO₂ (in biomass, leaf area, leaf mass per area, and photosynthesis), but that the relative response to elevated CO₂ for all variables was not significantly greater for lianas than trees regardless of the season. The lack of differences in the relative response between growth forms does not support the hypothesis that elevated CO₂ is responsible for increasing liana size and abundance across the neotropics.     

Leaf gas exchange and nitrogen dynamics of N2-fixing, field-grown Alnus glutinosa under elevated atmospheric CO2

Few studies have investigated the effects of elevated CO₂ on the physiology of symbiotic N₂-fixing trees. Tree species grown in low N soils at elevated CO₂ generally show a decline in photosynthetic capacity over time relative to ambient CO₂ controls. This negative adjustment may be due to a reallocation of leaf N away from the photosynthetic apparatus, allowing for more efficient use of limiting N. We investigated the effect of twice ambient CO₂ on net CO₂ assimilation (A), photosynthetic capacity, leaf dark respiration, and leaf N content of N2-fixing Alnus glutinosa (black alder) grown in field open top chambers in a low N soil for 160 d. At growth CO₂, A was always greater in elevated compared to ambient CO₂ plants. Late season A vs. internal leaf p(CO₂) response curves indicated no negative adjustment of photosynthesis in elevated CO₂ plants. Rather, elevated CO₂ plants had 16% greater maximum rate of CO₂ fixation by Rubisco. Leaf dark respiration was greater at elevated CO₂ on an area basis, but unaffected by CO₂ on a mass or N basis. In elevated CO₂ plants, leaf N content (μg N cm^?2) increased 50% between Julian Date 208 and 264. Leaf N content showed little seasonal change in ambient CO₂ plants. A single point acetylene reduction assay of detached, nodulated root segments indicated a 46% increase in specific nitrogenase activity in elevated compared to ambient CO₂ plants. Our results suggest that N₂-fixing trees will be able to maintain high A with minimal negative adjustment of photosynthetic capacity following prolonged exposure to elevated CO₂ on N-poor soils.     

Responses of communities of tropical tree species to elevated CO2 in a forest clearing

Communities of ten species of tropical forest tree seedlings from three successional classes were grown at ambient and elevated CO₂ in large open-top chambers on the edge of a forest in Panamá. Communities grew from 20?cm to approximately 2?m in height in 6 months. No enhancements in plant biomass accumulation occurred under elevated CO₂ either in the whole communities or in growth of individual species. Reductions in leaf area index under elevated CO₂ were observed, as were decreases in leaf nitrogen concentrations and increases in the C:N ratio of leaf tissue. Species tended to respond individualistically to elevated CO₂, but some generalizations of how successional groupings responded could be made. Early and mid-successional species generally showed greater responses to elevated CO₂ than late-successional species, particularly with respect to increases in photosynthetic rates and leaf starch concentrations, and reductions in leaf area ratio. Late-successional species showed greater increases in C:N ratios in response to elevated CO₂ than did other species. Our results indicate that there may not be an increase in the growth of regenerating tropical forest under elevated CO₂, but that there could be changes in soil nutrient availability because of reductions in leaf tissue quality, particularly in late-successional species.     

Interactive effects of elevated CO2 and drought stress on leaf water potential and growth in Caragana intermedia

We studied the responses of leaf water potential (Ψ_w), morphology, biomass accumulation and allocation, and canopy productivity index (CPI) to the combined effects of elevated CO₂ and drought stress in Caragana intermedia seedlings. Seedlings were grown at two CO₂ concentrations (350 and 700 μmol mol⁻¹) interacted with three water regimes (60–70%, 45–55%, and 30–40% of field capacity of soil). Elevated CO₂ significantly increased Ψ_w, decreased specific leaf area (SLA) and leaf area ratio (LAR) of drought-stressed seedlings, and increased tree height, basal diameter, shoot biomass, root biomass as well as total biomass under the all the three water regimes. Growth responses to elevated CO₂ were greater in well-watered seedlings than in drought-stressed seedlings. CPI was significantly increased by elevated CO₂, and the increase in CPI became stronger as the level of drought stress increased. There were significant interactions between elevated CO₂ and drought stress on leaf water potential, basal diameter, leaf area, and biomass accumulation. Our results suggest that elevated CO₂ may enhance drought avoidance and improved water relations, thus weakening the effect of drought stress on growth of C. intermedia seedings.     

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.     

System-level adjustments to elevated CO2 in model spruce ecosystems

Atmospheric carbon dioxide enrichment and increasing nitrogen deposition are often predicted to increase forest productivity based on currently available data for isolated forest tree seedlings or their leaves. However, it is highly uncertain whether such seedling responses will scale to the stand level. Therefore, we studied the effects of increasing CO₂ (280, 420 and 560 μL L^-1) and increasing rates of wet N deposition (0, 30 and 90 kg ha^-1 y^-1) on whole stands of 4-year-old spruce trees (Picea abies). One tree from each of six clones, together with two herbaceous understory species, were established in each of nine 0.7 m² model ecosystems in nutrient poor forest soil and grown in a simulated montane climate for two years. Shoot level light-saturated net photosynthesis measured at growth CO₂ concentrations increased with increasing CO₂, as well as with increasing N deposition. However, predawn shoot respiration was unaffected by treatments. When measured at a common CO₂ concentration of 420 μL L^-1 37% down-regulation of photosynthesis was observed in plants grown at 560 μL CO₂ L^-1. Length growth of shoots and stem diameter were not affected by CO₂ or N deposition. Bud burst was delayed, leaf area index (LAI) was lower, needle litter fall increased and soil CO₂ efflux increased with increasing CO₂. N deposition had no effect on these traits. At the ecosystem level the rate of net CO₂ exchange was not significantly different between CO₂ and N treatments. Most of the responses to CO₂ studied here were nonlinear with the most significant differences between 280 and 420 μL CO₂ L^-1 and relatively small changes between 420 and 560 μL CO₂ L^-1. Our results suggest that the lack of above-ground growth responses to elevated CO₂ is due to the combined effects of physiological down-regulation of photosynthesis at the leaf level, allometric adjustment at the canopy level (reduced LAI), and increasing strength of below-ground carbon sinks. The non-linearity of treatment effects further suggests that major responses of coniferous forests to atmospheric CO₂ enrichment might already be under way and that future responses may be comparatively smaller.     

Effects of elevated carbon dioxide and/or ozone on endogenous plant hormones in the leaves of <Emphasis Type="Italic">Ginkgo biloba</Emphasis>

Four-year-old Gingko (Ginkgo biloba L.) trees were exposed to ambient and elevated concentrations of CO₂ and O₃, and their combination for 1 year, using open-top chambers in Shenyang, China in 2006. Growth parameters and endogenous plant hormones were measured simultaneously over the experiment period. Elevated CO₂ increased leaf area and leaf dry weight but had no effect on shoot length, increased indole-3-acetic acid (IAA), gibberellins A₃ (GA₃), zeatin riboside (ZR), dihydrozeatin (DHZR) and isopentenyl-adenosine (iPA) content but decreased abscisic acid (ABA) content. Elevated O₃ significantly decreased leaf area, leaf dry weight, shoot length, and decreased IAA, GA₃, ZR content but increased ABA content and had a little effect on iPA, DHZR content. Elevated CO₂ + O₃ decreased IAA, iPA and DHZR content while increased ABA and GA₃ content in the early stage of the exposure and then decreased in the late stage. The evidence from this study indicates that elevated CO₂ ameliorated the effects of elevated ozone on tree growth, and elevated CO₂ may have a largely positive impact on forest tree growth while elevated O₃ will likely have a negative impact.     

Where temperate meets tropical: multi-factorial effects of elevated CO2, nitrogen enrichment, and competition on a mangrove-salt marsh community

Our understanding of how elevated CO₂ and interactions with other factors will affect coastal plant communities is limited. Such information is particularly needed for transitional communities where major vegetation types converge. Tropical mangroves (Avicennia germinans) intergrade with temperate salt marshes (Spartina alterniflora) in the northern Gulf of Mexico, and this transitional community represents an important experimental system to test hypotheses about global change impacts on critical ecosystems. We examined the responses of A. germinans (C₃) and S. alterniflora (C₄), grown in monoculture and mixture in mesocosms for 18 months, to interactive effects of atmospheric CO₂ and pore water nitrogen (N) concentrations typical of these marshes. A. germinans, grown without competition from S. alterniflora, increased final biomass (35%) under elevated CO₂ treatment and higher N availability. Growth of A. germinans was severely curtailed, however, when grown in mixture with S. alterniflora, and enrichment with CO₂ and N could not reverse this growth suppression. A field experiment using mangrove seedlings produced by CO₂‐ and N‐enriched trees confirmed that competition from S. alterniflora suppressed growth under natural conditions and further showed that herbivory greatly reduced survival of all seedlings. Thus, mangroves will not supplant marsh vegetation due to elevated CO₂ alone, but instead will require changes in climate, environmental stress, or disturbance to alter the competitive balance between these species. However, where competition and herbivory are low, elevated CO₂ may accelerate mangrove transition from the seedling to sapling stage and also increase above‐ and belowground production of existing mangrove stands, particularly in combination with higher soil N.     

Growth and maintenance respiration in stems of Quercus alba after four years of CO2 enrichment

Atmospheric CO₂ enrichment is increasingly being reported to inhibit leaf and whole-plant respiration. It is not known, however, whether this response is unique to foliage or whether woody-tissue respiration might be affected as well. This was examined for mid-canopy stem segments of white oak (Quercus alba L.) trees that had been grown in open-top field chambers and exposed to either ambient or ambient + 300 µmol mol^?1 CO₂ over a 4-year period. Stem respiration measurements were made throughout 1992 by using an infrared gas analyzer and a specially designed in situ cuvette. Rates of woody-tissue respiration were similar between CO₂ treatments prior to leaf initiation and after leaf senescence, but were several fold greater for saplings grown at elevated concentrations of CO₂ during much of the growing season. These effects were most evident on 7 July when stem respiration rates for trees exposed to elevated CO₂ concentrations were 7.25 compared to 3.44 µmol CO₂ m^?2 s^?1 for ambient-grown saplings. While other explanations must be explored, greater rates of stem respiration for saplings grown at elevated CO₂ concentrations were consistent with greater rates of stem growth and more stem-wood volume present at the time of measurement. When rates of stem growth were at their maximum (7 July to 3 August), growth respiration accounted for about 80 to 85% of the total respiratory costs of stems at both CO₂ treatments, while 15 to 20% supported the costs of stem-wood maintenance. Integrating growth and maintenance respiration throughout the season, taking into account treatment differences in stem growth and volume, indicated that there were no significant effects of elevated CO₂ concentration on either respiratory process. Quantitative estimates that could be used in modeling the costs of woody-tissue growth and maintenance respiration are provided.     

CO2 enrichment reduces the relative contribution of latex and latex-related hydrocarbons to biomass in Euphorbia lathyris

The hypothesis that plants grown under elevated CO₂ allocate more carbon to the production of latex and C‐rich secondary compounds whereas nutrient addition counteracts this effect was tested. Two similar experiments were conducted in two different experimental facilities. In both facilities seedlings of Euphorbia lathyris were exposed to factorial combinations of two CO₂ concentrations and two levels of nutrient availability for 2 months. The CO₂ treatments and growth conditions differed substantially between these two experiments but treatment responses to elevated CO₂ and fertilizer addition were remarkably similar, underlining the robustness of our findings. Elevated CO₂ increased biomass to a greater extent in fertilized than in unfertilized plants and reduced the leaf biomass fraction by accelerating leaf senescence. Concentrations of non‐structural carbohydrates (NSC) increased in elevated CO₂. However, this apparent carbon surplus did not feed into the whole plant latex pool. The latex harvest per leaf (?25%) and the concentration of latex‐related hydrocarbons (?20%) even decreased under elevated CO₂ (both experiments P < 0.05). Fertilization reduced NSC concentrations (?25%) but neither affected latex yield per leaf nor the concentration of latex‐related hydrocarbons. It is concluded that latex and related hydrocarbons in CO₂‐enriched plants are a negligible sink for excess carbon irrespective of nutrient status and thus, vigour of growth.     

Xeromorphy increases in shoots of Pseudotsuga menziesii (Mirb.) Franco seedlings with exposure to elevated temperature but not elevated CO2

Seedling structure influences tree structure and function, ultimately determining the potential productivity of trees and their competitiveness for resources. We investigated changes in shoot structure for seedlings of Pseudotsuga menziesii (Douglas-fir) grown under climate change scenarios of ambient or elevated CO₂ (+180 mol mol^–1) plus ambient or elevated temperature (+3.5°C), for 4 years in outdoor, sunlit chambers. Mass allocation and allometry were measured for buds, leaves, branches, and stems, and anatomy was evaluated for leaves and stems. Seedlings became more xeromorphic with elevated temperature: allocation of total mass to branches over stems and leaves increased, sapwood area to height ratio increased, number of growing points relative to seedling size increased, and stem and branch length and mass decreased for sections initiated during the three full CO₂ and temperature seasons. Neither stem nor leaf anatomy was affected by elevated temperature. Elevated CO₂ increased specific mass of leaves, but had few other effects on mass allocation, allometry, or anatomy for any shoot organ. There were no CO₂ × temperature interactions for any important parameter. Thus, under realistic simulated field environmental conditions representative for in at least some P. menziesii forests (i.e., OR, USA, forests with limited soil nitrogen and summer soil moisture), elevated temperature, but not elevated CO₂, may affect seedling shoot structure and, hence, function.     

CO2 enrichment predisposes foliage of a eucalypt to freezing injury and reduces spring growth

Seedlings of Eucalyptus pauciflora, were grown in open-top chambers fumigated with ambient and elevated [CO₂], and were divided into two populations using 10% light transmittance screens. The aim was to separate the effects of timing of light interception, temperature and [CO₂] on plant growth. The orientation of the screens exposed plants to a similar total irradiance, but incident during either cold mornings (east-facing) or warm afternoons (west-facing). Following the first autumn freezing event elevated CO₂-grown plants had 10 times more necrotic leaf area than ambient CO₂ plants. West-facing plants had significantly greater (25% more) leaf damage and lower photochemical efficiency (F_v/F_m) in comparison with east-facing plants. Following a late spring freezing event east-facing elevated CO₂ plants suffered a greater sustained loss in F_v/F_m than west-facing elevated CO₂- and ambient CO₂-grown plants. Stomatal conductance was lower under elevated CO₂ than ambient CO₂ except during late spring, with the highest leaf temperatures occurring in west-facing plants under elevated CO₂. These higher leaf temperatures apparently interfered with cold acclimation thereby enhancing frost damage and reducing the ability to take advantage of optimal growing conditions under elevated CO₂.     

Thirty years of in situ tree growth under elevated CO2: a model for future forest responses?

Rising concentrations of atmospheric carbon dioxide have been predicted to stimulate the growth of forest trees. However, long-term effects on trees growing to maturity and to canopy closure while exposed to elevated CO₂ have never been examined. We compared tree ring chronologies of Mediterranean Quercus ilex which have been continuously exposed to elevated CO₂ (around 650 μmol mol^–1) since they were seedlings, near two separate natural CO₂ springs with those from trees at nearby ambient-CO₂‘control’ sites. Trees grown under high CO₂ for 30 years (1964–93) showed a 12% greater final radial stem width than those growing at the ambient-CO₂ control sites. However, this stimulation was largely due to responses when trees were young. By the time trees were 25–30 y old the annual difference in tree ring width between low and high CO₂ grown trees had disappeared. At any given tree age, elevated CO₂ had a relatively greater positive effect on tree ring width in years with a dry spring compared to years with more rainfall between April and May. This indicates a beneficial effect of elevated CO₂ on tree water relations under drought stress. Our data suggest that the early regeneration phase of forest stands can be accelerated in CO₂-enriched atmospheres and that maximum biomass per land area may be reached sooner than under lower CO₂ concentrations. In our study, high CO₂ grown Q. ilex trees reached the same stem basal area at the age of 26 y as control trees at 29 y, i.e. three years earlier (faster turnover of carbon?). Reliable predictions of the future development of forests need to account for the variable responses of trees over their entire lifetime. Such responses to elevated CO₂ can presently only be assessed at such unique field sites.