Photosynthetic responses of forest understory tree species to long-term exposure to elevated carbon dioxide concentration at the Duke Forest FACE experiment

We examined the photosynthetic responses of four species of saplings growing in the understory of the Duke Forest FACE experiment during the seventh year of exposure to elevated CO2 concentration ([CO2]). Saplings of these same species were measured in the first year of the Duke Forest FACE experiment and at that time showed only seasonal fluctuations in acclimation of photosynthesis to elevated [CO2]. Based on observations from the Duke Forest FACE experiment, we hypothesized that after seven years of exposure to elevated [CO2] significant photosynthetic down-regulation would be observed in these tree species. To test our hypothesis, photosynthetic CO2-response and light-response curves, along with chlorophyll fluorescence, chlorophyll concentration and foliar N were measured twice during the summer of 2003. Exposure to elevated [CO2] continued to increase photosynthesis in all species measured after seven years of treatment with the greatest photosynthetic increase observed near saturating irradiances. In all species, elevated [CO2] increased electron transport efficiency but did not significantly alter carboxylation efficiency. Quantum yield estimated by light curves, chlorophyll concentration, and foliar nitrogen concentrations were unaffected by elevated [CO2]. Contrary to our hypothesis, there is little evidence of progressive N limitation of leaf-level processes in these understory tree species after seven years of exposure to elevated [CO2] in the Duke Forest FACE experiment.     

Growth and photosynthetic responses of four Virginia Piedmont tree species to shade

To determine the effects of shade on biomass, carbon allocation patterns and photosynthetic response, seedlings of loblolly pine (Pinus taeda L.), white pine (Pinus strobus L.), red maple (Acer rubrum L.), and yellow-poplar (Liriodendron tulipifera L.) were grown without shade or in shade treatments providing a 79 or 89% reduction of full sunlight for two growing seasons. The shade treatments resulted in less total biomass for all species, with loblolly pine showing the greatest shade-induced growth reduction. Yellow-poplar was the only species to show increased stem height growth in the 89% shade treatment. The shade treatments increased specific leaf area of all species. Quantum efficiency, dark respiration and light compensation point were generally not affected by the shade treatments. Quantum efficiency, dark respiration, maximum photosynthesis and light compensation point did not change consistently between the first and second growing seasons. We conclude that differences in shade tolerance among these species are not the result of changes in the photosynthetic mechanism in response to shade.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

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

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

Ecophysiological responses and carbon distribution ofPinus koraiensis seedlings to elevated carbon dioxide

The net CO₂ assimilation rate, stomatal conductance, RuBPcase (ribulose 1,5-biphosphate carboxylose) activity, dry weight of aboveground and belowgroud part, plant height, the length and diameter of taproot ofPinus koraiensis seedlings were measured and analyzed after six-week exposure to elevated CO₂ in an open-top chamber in Changbai Mountain of China from May to Oct. 1999. Seedlings were planted in four different conditions: on an open site, control chamber, 500 μL·L⁻¹ and 700 μL·L⁻¹ CO₂ chambers. The results showed that the total biomass of the seedlings increased whereas stomatal conductance decreased. The physiological responses and growth to 500 μL·L⁻¹ and 700 μL·L⁻¹ CO₂ varied greatly. The acclimation of photosynthesis was downward to 700 μL·L⁻¹ CO₂ but upward to 500 μL·L⁻¹ CO₂. The RuBPcase activity, chlorophyll and soluble sugar contents of the seedlings grown at 500 μL·L⁻¹ CO₂ were higher than that at 700 μL·L⁻¹ CO₂. The concentration 500 μL·L⁻¹ CO₂ enhanced the growth of aboveground part whereas 700 μL·L⁻¹ CO₂ allocated more carbon to belowground part. Elevated CO₂ changed the carbon distribution pattern. The ecophysiological responses were significantly different between plants grown under 500 μL·L⁻¹ CO₂ and 700 μL·L⁻¹ CO₂. Foundation Item: This paper was supported by Chinese Academy of Sciences. Biography: HAN Shi-jie (1956-), male, Ph. Doctor, Professor in Laboratory of Ecological Process of Trace Substance in Terrestrial Ecosystem, Institute of Applied Ecology, Chinese Academy of Sciences. Responsible editor: Chai Ruihai     

Ecophysiological responses and carbon distribution of Pinus koraiensis seedlings to elevated carbon dioxide

The net CO2 assimilation rate, stomatal conductance, RuBPcase (ribulose 1,5-biphosphate carboxylose) activity, dry weight of aboveground and belowground part, plant height, the length and diameter of taproot of Pinus koraiensis seedlings were measured and analyzed after six-week exposure to elevated CO2 in an open-top chamber in Changbai Mountain of China from May to Oct. 1999. Seedlings were planted in four different conditions: on an open site, control chamber, 500 μ L.L-1 and 700 μL.L-1 CO2 chambers. The results showed that the total biomass of the seedlings increased whereas stomatal conductance decreased. The physiological responses and growth to 500 μL.L-1 and 700 μ L.L-1 CO2 varied greatly. The acclimation of photosynthesis was downward to 700 μL.L-1 CO2 but upward to 500 μ L.L-1 CO2. The RuBPcase activity, chlorophyll and soluble sugar contents of the seedlings grown at 500 μL.L-1 CO2 were higher than that at 700 μ L.L-1 CO2. The concentration 500 μ L.L-1 CO2 enhanced the growth of aboveground part whereas 700 μL.L-1 CO2 allocated more carbon to belowground part. Elevated CO2 changed the carbon distribution pattern. The ecophysiological responses were significantly different between plants grown under 500 μL.L-1 CO2 and 700 μL.L-1 CO2.     

Carbon budget of Pinus sylvestris saplings after four years of exposure to elevated atmospheric carbon dioxide concentration

To study the responses of Scots pine (Pinus sylvestris L.), a commercially important tree species in Europe, to future increases in atmospheric CO2 concentration ([CO2]), we grew saplings for 4 years in the ground in open-top chambers in ambient or ambient + 400 micromol mol(-1) CO2, without supplemental addition of nutrients and water. Carbon (C) budgets were developed for trees in both CO2 treatments based on productivity and biomass data obtained from destructive harvests at the end of the third and fourth years of treatment, and simulations of annual gross photosynthesis (P(tot)) and maintenance respiration by the model MAESTRA. Simulated P(tot) was enhanced by elevated [CO2], despite significant down-regulation of photosynthetic capacity. The subsequent increase in C uptake was allocated primarily to tissues with limited longevity (needles and fine roots), which explains why the measured annual increment in woody biomass did not differ between CO2 treatments. Thus, our results suggest that accelerated stem growth only occurs in the first 2 years in the presence of elevated [CO2] and that forest rotations will not be shortened significantly in response to increasing [CO2]. In elevated [CO2], a higher proportion of available C was allocated below ground, resulting in altered biomass distribution patterns. In trees of equal size, measured ratios of fine root/needle biomass and belowground/aboveground biomass were almost twice as large in the elevated [CO2] treatment. Although there are uncertainties in scaling from saplings to mature canopies, the data indicate that, in nutrient-limited Scots pine forests, elevated [CO2] is unlikely to accelerate tree growth significantly, but is likely to increase C inputs to soil.     

Photosynthetic response ofPinus sylvestriformis to elevated carbon dioxide and its influential factor analysis

The photosynthetic response of 12-year oldPinus sylvestriformis to elevated CO₂ and its influential factors were tested and analyzed in the forest region of Changbai Mountain in 1999. Trees grown at the natural condition were controlled at three levels of CO₂ concentration (350 μL·L⁻¹, 500 μL·L⁻¹ and 700 μL·L⁻¹) by CO₂ rich settlement designed by us. Net photosynthetic rates (NPR), temperature, relative humidity, stomatal conductance, intercellular CO₂ concentration and photosynthetic active radiation (PAR) were measured at 6:00, 8:00, 10:00, 14:00, 16:00 and 18:00 hours a day. Experimental results showed that the NPR ofPinus sylvestriformis increased by 32.6% and 123.0% at 500 μL·L⁻¹ and 700 μL·L⁻¹ CO₂ concentration respectively, compared to ambient atmospheric CO₂ concentration (350 μL·L⁻¹). The relations between NPR and influential factors, including temperature, relative humidity, intercellular CO₂ concentration and photosynthetic active radiation, were analyzed respectively by regression analysis at different CO₂ concentrations. Foundation Item: This project was supported by Chinese Academy of Sciences. Biography: WANG Chen-rui (1970-), male, Assistant Research Fellow in Institute of Applied Ecology, Chinese Academy of Sciences. Responsible editor: Chai Ruihai     

Photosynthetic response of Pinus sylvestriformis to elevated carbon dioxide and its influential factor analysis

IntroductionOur world is changing in the way and at the speedthat are describable, but we are unable to predictthese changes with any degree of accuracy. Radioactive and chemical properties of the atmosphere,global climate, and global ecology are dynamic andmeasurable, but they also linked to each other incomplex and poorly understood ways (Rayal andRamanathan 1989). While many of the physical andbiological sub-processes are understood and modeled in detail, predictive capabilities are poor i…     

Patterns of photosynthesis and starch allocation in seedlings of four bottomland hardwood tree species subjected to flooding

Effects of short-term (32 days) flooding on photosynthesis, stomatal conductance, relative growth rate and tissue starch concentrations of flood-intolerant Quercus alba L. (white oak), bottomland Quercus nigra L. (water oak), bottomland Fraxinus pennsylvanica Marshall. (green ash) and flood-tolerant Nyssa aquatica L. (water tupelo) seedlings were studied under controlled conditions. Net photosynthetic rates of flooded N. aquatica seedlings were reduced by 25% throughout the 32-day flooding period. Net photosynthetic rates of flooded Q. alba seedlings fell rapidly to 25% of those of the control seedlings by Day 4 of the flooding treatment and to 5% by Day 16. In F. pennsylvanica and Q. nigra, net photosynthetic rates were reduced to 50% of control values by Day 8 but remained at approximately 30 and 23%, respectively, of control values by Day 32. Leaves of flooded Q. alba seedlings accumulated approximately twice as much starch as leaves of non-flooded control plants, whereas root starch concentrations decreased to 67% of those of control plants by the end of the 32-day flooding treatment. In contrast, flooding caused only a small increase in leaf starch concentrations of N. aquatica plants, but it increased root starch concentrations to 119% of those of the control plants by the end of the experiment. The co-occurring bottomland species, Fraxinus pennsylvanica and Q. nigra, differed from each other in their patterns of stomatal conductance and root starch concentrations. We conclude that the maintenance of low leaf starch concentrations, and high pre-flood root tissue starch concentrations are important characteristics allowing flood-tolerant species to survive in flooded soils.     

Short- and long-term responses of total soil organic carbon to harvesting in a northern hardwood forest

The long-term response of total soil organic carbon pools (‘total SOC’, i.e. soil and dead wood) to different harvesting scenarios in even-aged northern hardwood forest stands was evaluated using two soil carbon models, CENTURY and YASSO, that were calibrated with forest plot empirical data in the Green Mountains of Vermont. Overall, 13 different harvesting scenarios that included four levels of aboveground biomass removal (20%, 40%, 60% and 90%) and four different rotation lengths (60 year, 90 year, 120 year, and No Rotation (NR)) were simulated for a 360 year period. Simulations indicate that following an initial post-harvest increase, total SOC decreases for several decades until carbon inputs into the soil pool from the re-growth are greater than losses due to decomposition. At this point total SOC begins to gradually increase until the next harvest. One consequence of this recovery pattern is that between harvests, the size of the SOC pool in a stand may change from −7 to 18% of the pre-harvest pool, depending on the soil pool considered. Over 360 years, the average annual decrease in total SOC depends on the amount of biomass removed, the rotation length, and the soil pool considered. After 360 years a stand undergoing the 90yr-40% scenario will have 15% less total SOC than a non-harvested stand. Long-term declines in total SOC greater than 10% were observed in the 60yr-60%, 60yr-90%, and 90yr-90% scenarios. Long-term declines less than 5% were observed in scenarios with 120 year rotations that remove 60% or less of the aboveground biomass. The long-term decreases simulated here for common management scenarios in this region would require intensive sampling procedures to be detectable.     

Long-term effects of elevated carbon dioxide concentration and provenance on four clones of Sitka spruce (Picea sitchensis). II. Photosynthetic capacity and nitrogen use efficiency

Four clones of Sitka spruce (Picea sitchensis (Bong.) Carr.) from two provenances, at 53.2 degrees N (Skidegate a and Skidegate b) and at 41.3 degrees N (North Bend a and North Bend b), were grown for three growing seasons in ambient (~350 micromol per mol) and elevated (~700 micromol per mol) CO2 concentrations. The clones were grown in stress-free conditions (adequate nutrition and water) to assess the effect of elevated [CO2] on tree physiology. Growth in elevated [CO2] significantly increased instantaneous photosynthetic rates of the clonal Sitka spruce saplings by about 62%. Downward acclimation of photosynthesis (A) was found in all four clones grown in elevated [CO2]. Rubisco activity and total chlorophyll concentration were also significantly reduced in elevated [CO2]. Provenance did not influence photosynthetic capacity. Best-fit estimates of Jmax (maximum rate of electron transport), Vcmax (RuBP-saturated rate of Rubisco) and Amax (maximum rate of assimilation) were derived from responses of A to intercellular [CO2] by using the model of Farquhar et al. (1980). At any leaf N concentration, the photosynthetic parameters were reduced by growth in elevated [CO2]. However, the ratio between Jmax and Vcmax was unaffected by CO2 growth concentration, indicating a tight coordination in the allocation of N between thylakoid and soluble proteins. In elevated [CO2], the more southerly clones had a higher initial N use efficiency (more carbon assimilated per unit of leaf N) than the more northerly clones, so that they had more N available for those processes or organs that were most limiting to growth at a particular time. This may explain the initial higher growth stimulation by elevated [CO2] in the North Bend clones than in the Skidegate clones.     

Photosynthesis and carbon allocation of six boreal tree species grown in understory and open conditions

One-year-old seedlings of Abies balsamea (L.) Mill, Picea glauca (Moench) Voss, Pinus contorta Loudon, Betula papyrifera Marsh., Populus tremuloides Michx. and Populus balsamifera L. were transplanted in the spring, in pots, to the understory of a mixed P. tremuloides-P. balsamifera stand or to an adjacent open site. Growth and leaf characteristics were measured and photosynthetic light response curves determined in mid-August. Overall, the coniferous seedlings showed less photosynthetic plasticity in response to growth conditions than the deciduous species. Abies balsamea, P. glauca and B. papyrifera responded to the understory environment with higher leaf area ratios, and lower photosynthetic light saturation points and area-based leaf respiration relative to values for open-grown seedlings, while they matched or exceeded the height growth of open-grown seedlings. In contrast, seedlings of Pinus contorta, P. tremuloides and P. balsamifera displayed characteristics that were not conducive to survival in the understory. These characteristics included a high light saturation point and leaf dark respiration rate in P. contorta, and lower leaf area variables combined with higher carbon allocation to roots in P. tremuloides and P. balsamifera. By the second growing season, all seedlings of P. tremuloides and P. balsamifera growing in the understory had died.     

Short-term light and leaf photosynthetic dynamics affect estimates of daily understory photosynthesis in four tree species

Instantaneous measurements of photosynthesis are often implicitly or explicitly scaled to longer time frames to provide an understanding of plant performance in a given environment. For plants growing in a forest understory, results from photosynthetic light response curves in conjunction with diurnal light data are frequently extrapolated to daily photosynthesis (A(day)), ignoring dynamic photosynthetic responses to light. In this study, we evaluated the importance of two factors on A(day) estimates: dynamic physiological responses to photosynthetic photon flux density (PPFD); and time-resolution of the PPFD data used for modeling. We used a dynamic photosynthesis model to investigate how these factors interact with species-specific photosynthetic traits, forest type, and sky conditions to affect the accuracy of A(day) predictions. Increasing time-averaging of PPFD significantly increased the relative overestimation of A(day) similarly for all study species because of the nonlinear response of photosynthesis to PPFD (15% with 5-min PPFD means). Depending on the light environment characteristics and species-specific dynamic responses to PPFD, understory tree A(day) can be overestimated by 6-42% for the study species by ignoring these dynamics. Although these overestimates decrease under cloudy conditions where direct sunlight and consequently understory sunfleck radiation is reduced, they are still significant. Within a species, overestimation of A(day) as a result of ignoring dynamic responses was highly dependent on daily sunfleck PPFD and the frequency and irradiance of sunflecks. Overall, large overestimates of A(day) in understory trees may cause misleading inferences concerning species growth and competition in forest understories with < 2% full sunlight. We conclude that comparisons of A(day) among co-occurring understory species in deep shade will be enhanced by consideration of sunflecks by using high-resolution PPFD data and understanding the physiological responses to sunfleck variation.     

Photosynthetic and transpirational responses of red spruce understory trees to light and temperature

Understory red spruce (Picea rubens Sarg.) trees, between 20 and 50 cm in height and 12 years or more in age, were collected from mid- and high-elevation stands in north-central Vermont and placed in a closed-cuvette system to measure photosynthetic and transpirational responses to photosynthetic photon flux density (PPFD) and temperature. Photosynthesis, dark respiration, transpiration and water-use efficiency of trees from both stands responded to changes in PPFD and temperature in similar ways. Trees from both stands exhibited maximum rates of net photosynthesis at temperatures between 15 and 20 degrees C, and exposure to higher temperatures resulted in reduced rates of photosynthesis and increased rates of respiration. Net photosynthetic rates generally increased with increasing light intensity but began to level off at 250 micro mol m(-2) s(-1). Water-use efficiency was maximal when temperature and PPFD were at 15 degrees C and above 400 micro mol m(-2) s(-1), respectively.     

Forecasting development of understory American beech after partial cutting in uneven-aged northern hardwood stands

American beech (Fagus grandifolia Ehrh.) in the understory of northern hardwood stands can interfere with the development of more desirable species such as sugar maple (Acer saccharum Marsh.). Yet, managers have had only limited methods for identifying stands where beech might dominate a new cohort. We examined five uneven-aged stands in New York State to determine the degree of influence that understory beech has on species development after a cutting treatment. No significant differences between stands were found, allowing us to develop a single model for evaluation. Understory beech was assessed using a species index value (SIV), which measures the proportion of stems, weighted by height, for a species on a milacre plot. We used a 5×5 transition matrix to compare pre- and post-cut beech importance levels. Stand-level trends showed an overall increase in the proportion of sample plots with higher levels of understory beech after cutting, with the likelihood of change on individual plots dependent on their pre-cut status. Plots with either no beech or very high levels of beech changed little after cutting, while beech was more likely to reach critical levels of dominance when pre-cut SIV was medium or higher. The transition matrix gives forest managers a means to forecast the probable level of future beech importance following cutting, based upon precursor conditions.     

Species selection in hardwoods research: variations in baseline physiological responses of select temperate hardwood tree species

Drought periods are becoming more extreme worldwide and the ability of plants to contribute towards atmospheric flux is being compromised. Properly functioning stomata provide an exit for water that has been absorbed by the roots, funneled into various cell parts, and eventually released into the atmosphere via transpiration. By observing the effects that weather conditions such as climate change may have on stomatal density, distribution, and functioning, it may be possible to elucidate a portion of the mechanisms trees use to survive longer periods of water stress. This study analyzed stomatal density (SD), stomatal conductance (gs ), CO2 assimilation (A), instantaneous water-use efficiency (WUEi ), and transpiration (E) rates in six native tree species in the Midwestern USA and showed that trees within the same ecotype followed similar trends, but that trees within the same family did not when exposed to identical greenhouse conditions. Naturally drought tolerant tree species demonstrated lower g s and higher WUEi , while intolerant species had higher SD. This study showed negative or no correlation between SD and g s , A, E, and WUEi and positive correlations between E and A and gs and E.     

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

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