Interactive effects of elevated [CO2] and drought on cherry (Prunus avium) seedlings I. Growth, whole-plant water use efficiency and water loss

Seeds of cherry ( Prunus avium ) were germinated and grown for two growing seasons in ambient (∼350 μmol mol⁻¹) or elevated (ambient+∼350 μmol mol⁻¹) CO₂ mole fractions in six open-top chambers. The seedlings were fertilized once a week, following Ingestad principles in order to supply mineral nutrients at free-access rates. In the first growing season gradual drought was imposed on rapidly growing cherry seedlings by withholding water for a 6-wk drying cycle. In the second growing season, the rapid onset of drought was imposed at the height of the growing season on the seedlings which had already experienced drought in the first growing season. Elevated [CO₂] significantly increased total dry-mass production in both water regimes, but did not ameliorate the growth response to drought of the cherry seedlings subjected to two sequential drying cycles. Water loss did not differ in either well watered or droughted seedlings between elevated and ambient [CO₂]; consequently whole-plant water- use efficiency (the ratio of total dry mass produced to total water consumption) was significantly increased. Similar patterns of carbon allocation between shoot and root were found in elevated and ambient [CO₂] when the seedlings were the same size. Thus, elevated [CO₂] did not improve drought tolerance, but it accelerated ontogenetic development irrespective of water status.     

Interaction of enriched CO2 and water stress on the physiology of and biomass production in sweet potato grown in open-top chambers

Abstract. The objective of this study was to investigate the effects of water stress in sweet potato ( Ipomoea batatas L. [Lam] 'Georgia Jet') on biomass production and plant-water relationships in an enriched CO₂ atmosphere. Plants were grown in pots containing sandy loam soil (Typic Paleudult) at two concentrations of elevated CO₂ and two water regimes in open-top field chambers. During the first 12 d of water stress, leaf xylem potentials were higher in plants grown in a CO₂ concentration of 438 and 666 μmol mol⁻¹ than in plants grown at 364 μmol mol⁻¹. The 364 μmol mol⁻¹ CO₂ grown plants had to be rewatered 2 d earlier than the high CO₂-grown plants in response to water stress. For plants grown under water stress, the yield of storage roots and root: shoot ratio were greater at high CO₂ than at 364 μmol mol⁻¹; the increase, however, was not linear with increasing CO₂ concentrations. In well-watered plants, biomass production and storage root yield increased at elevated CO₂, and these were greater as compared to water-stressed plants grown at the same CO₂ concentration.     

Growth and physiological responses of canola (Brassica napus) to three components of global climate change: temperature, carbon dioxide and drought

Elevated CO₂ appears to be a significant factor in global warming, which will likely lead to drought conditions in many areas. Few studies have considered the interactive effects of higher CO₂, temperature and drought on plant growth and physiology. We grew canola ( Brassica napus cv. 45H72) plants under lower (22/18°C) and higher (28/24°C) temperature regimes in controlled-environment chambers at ambient (370 μmol mol⁻¹) and elevated (740 μmol mol⁻¹) CO₂ levels. One half of the plants were watered to field capacity and the other half at wilting point. In three separate experiments, we determined growth, various physiological parameters and content of abscisic acid (ABA), indole-3-acetic acid and ethylene. Drought-stressed plants grown under higher temperature at ambient CO₂ had decreased stem height and diameter, leaf number and area, dry matter, leaf area ratio, shoot/root weight ratio, net CO₂ assimilation and chlorophyll fluorescence. However, these plants had increased specific leaf weight, leaf weight ratio and chlorophyll concentration. Elevated CO₂ generally had the opposite effect, and partially reversed the inhibitory effects of higher temperature and drought on leaf dry weight accumulation. This study showed that higher temperature and drought inhibit many processes but elevated CO₂ partially mitigate some adverse effects. As expected, drought stress increased ABA but higher temperature inhibited the ability of plants to produce ABA in response to drought.     

Disturbance, rainfall and contrasting species responses mediated aboveground biomass response to 11 years of CO2 enrichment in a Florida scrub-oak ecosystem

This study reports the aboveground biomass response of a fire-regenerated Florida scrub-oak ecosystem exposed to elevated CO₂ (1996–2007), from emergence after fire through canopy closure. Eleven years exposure to elevated CO₂ caused a 67% increase in aboveground shoot biomass. Growth stimulation was sustained throughout the experiment; although there was significant variability between years. The absolute stimulation of aboveground biomass generally declined over time, reflecting increasing environmental limitations to long-term growth response. Extensive defoliation caused by hurricanes in September 2004 was followed by a strong increase in shoot density in 2005 that may have resulted from reopening the canopy and relocating nitrogen from leaves to the nutrient-poor soil. Biomass response to elevated CO₂ was driven primarily by stimulation of growth of the dominant species, Quercus myrtifolia , while Quercus geminata , the other co-dominant oak, displayed no significant CO₂ response. Aboveground growth also displayed interannual variation, which was correlated with total annual rainfall. The rainfall × CO₂ interaction was partially masked at the community level by species-specific responses: elevated CO₂ had an ameliorating effect on Q. myrtifolia growth under water stress. The results of this long-term study not only show that atmospheric CO₂ concentration had a consistent stimulating effect on aboveground biomass production, but also showed that available water is the primary driver of interannual variation in shoot growth and that the long-term response to elevated CO₂ may have been caused by other factors such as nutrient limitation and disturbance.     

Effects of source-sink relations on photosynthetic acclimation to elevated CO2

Abstract. While photosynthesis of C₃ plants is stimulated by an increase in the atmospheric CO₂ concentration, photosynthetic capacity is often reduced after long-term exposure to elevated CO₂. This reduction appears to be brought about by end product inhibition, resulting from an imbalance in the supply and demand of carbohydrates. A review of the literature revealed that the reduction of photosynthetic capacity in elevated CO₂ was most pronounced when the increased supply of carbohydrates was combined with small sink size. The volume of pots in which plants were grown affected the sink size by restricting root growth. While plants grown in small pots had a reduced photosynthetic capacity, plants grown in the field showed no reduction or an increase in this capacity. Pot volume also determined the effect of elevated CO₂ on the root/shoot ratio: the root/shoot ratio increased when root growth was not restricted and decreased in plants grown in small pots. The data presented in this paper suggest that plants growing in the field will maintain a high photosynthetic capacity as the atmospheric CO₂ level continues to rise.     

Growth, dry matter partitioning, and diurnal activities of RuBP carboxylase in citrus seedlings maintained at two levels of CO2

The long-term response of citrus rootstock seedlings to CO₂ enrichment was examined in Carrizo estrange ( Poncirua trifoliata (L.) Raf. x Citrus sinensis (L.) Osbeck] and Swingle citrumelo ( P. trifoliate x C. parodist Macf.]. Plaotlets 14 weeks old were transferred to outdoor controlled-environment chambers and maintained for 5 months from Feb. 14 to July 21. During this period, new growth (cm) of citrange and citrumelo shoots at 660 μl1⁻¹ was 94 and 69% greater, respectively, than at 330 μ1 1⁻¹. Total dry weight of both rootstock shoots had increased by over 100%. Growth of few species is affected this markedly by elevated CO₂ levels.
More carbon was partitioned to above-ground organs in CO₂-enriched citrus seedlings. Stem dry matter per unit length was also 32 and 44% greater in citrange and citrumelo, respectively. Total leaf area was increased by 124% in citrange and 85% in citrumelo due to greater leaf number and size. Variations in overall relative growth rate appeared to be related to the rapid, sequential, flush-type growth in citrus, in which an entire shoot segment with its associated leaves remains an active sink until fully expanded. RuBP carboxylase (EC 4.1.1.39) activity in leaves of recently-expanded flushes was higher in citrumelo plants grown at 660 vs 330 μ1 1⁻¹ CO₂ and changed diurnally for citrange (but not citrumelo) leaves at both CO₂ levels. The results are consistent with the hypothesis that positive long-term effects of CO² enrichment may be greater in species or during growth periods where sink capacity for carbon utilization is high.     

The effects of increased atmospheric carbon dioxide and temperature on carbon partitioning, source-sink relations and respiration

Abstract. Herbaceous C₃ plants grown in elevated CO₂ show increases in carbon assimilation and carbohydrate accumulation (particularly starch) within source leaves. Although changes in the partitioning of biomass between root and shoot occur, the proportion of this extra assimilate made available for sink growth is not known. Root:shoot ratios tend to increase for CO₂-enriched herbaceous plants and decrease for CO₂-enriched trees. Root:shoot ratios for cereals tend to remain constant. In contrast, elevated temperatures decrease carbohydrate accumulation within source and sink regions of a plant and decrease root:shoot ratios. Allometric analysis of at least two species showing changes in root: shoot ratios due to elevated CO₂ show no alteration in the whole-plant partitioning of biomass. Little information is available for interactions between temperature and CO₂. Cold-adapted plants show little response to elevated levels of CO₂, with some species showing a decline in biomass accumulation. In general though, increasing temperature will increase sucrose synthesis, transport and utilization for CO₂-enriched plants and decrease carbohydrate accumulation within the leaf. Literature reports are discussed in relation to the hypothesis that sucrose is a major factor in the control of plant carbon partitioning. A model is presented in support.     

CO2 enrichment and development of freezing tolerance in Norway spruce

Plant growth and adaptation to cold and freezing temperatures in a CO₂-enriched atmosphere have received little attention despite their predicted effects on plant distribution and productivity. In this study we looked at the interaction between elevated CO₂ and development of freezing tolerance in Norway spruce ( Picea abies (L.) Karst.). First-year seedlings were grown under controlled conditions in an atmosphere enriched in CO₂ (70 Pa) for one simulated growth season. We measured shoot growth, registered the timing of growth cessation and bud set, measured needle net photosynthetic rate, and determined needle carbohydrate concentration (fructose+pinitol, glucose, sucrose, inositol, raffinose and starch). Freezing tolerance (LT₅₀) was determined after exposing whole seedlings to temperatures ranging from −6.5 to −36.0°C and scoring for visual needle browning. Elevated CO₂ did not affect height growth or the timing of growth cessation and bud set. The only statistically significant effects of CO₂ treatment were on seedling dry weight, percent dry matter and starch content. During the three weeks after growth cessation and bud set, freezing tolerance increased from −10 to −35°C, and there was a marked increase in all soluble sugars except inositol. However, neither freezing tolerance nor the concentration of soluble sugars was significantly influenced by elevated CO₂.     

Effect of elevated CO2, temperature and drought on photosynthesis of nodulated alfalfa during a cutting regrowth cycle

Rising atmospheric CO₂ may increase potential net leaf photosynthesis under short-term exposure, but this response decreases under long-term exposure because plants acclimate to elevated CO₂ concentrations through a process known as downregulation. One of the main factors that may influence this phenomenon is the balance between sources and sinks in the plant. The usual method of managing a forage legume like alfalfa requires the cutting of shoots and subsequent regrowth, which alters the source/sink ratio and thus photosynthetic behaviour. The aim of this study was to determine the effect of CO₂ (ambient, around 350 vs. 700 µmol mol⁻¹), temperature (ambient vs. ambient + 4° C) and water availability (well-irrigated vs. partially irrigated) on photosynthetic behaviour in nodulated alfalfa before defoliation and after 1 month of regrowth. At the end of vegetative normal growth, plants grown under conditions of elevated CO₂ showed photosynthetic acclimation with lower photosynthetic rates, V_cmax and ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) activity. This decay was probably a consequence of a specific rubisco protein reduction and/or inactivation. In contrast, high CO₂ during regrowth did not change net photosynthetic rates or yield differences in V_cmax or rubisco total activity. This absence of photosynthetic acclimation was directly associated with the new source-sink status of the plants during regrowth. After cutting, the higher root/shoot ratio in plants and remaining respiration can function as a strong sink for photosynthates, avoiding leaf sugar accumulation, the negative feed-back control of photosynthesis, and as a consequence, photosynthetic downregulation.     

The response of nodulated alfalfa to water supply, temperature and elevated CO2: photosynthetic downregulation

Plants grown in an environment of elevated CO₂ and temperature often show reduced CO₂ assimilation capacity, providing evidence of photosynthetic downregulation. The aim of this study was to analyse the downregulation of photosynthesis in elevated CO₂ (700 µmol mol⁻¹) in nodulated alfalfa plants grown at different temperatures (ambient and ambient + 4°C) and water availability regimes in temperature gradient tunnels. When the measurements were taken in growth conditions, a combination of elevated CO₂ and temperature enhanced the photosynthetic rate; however, when they were carried out at the same CO₂ concentration (350 and 700 µmol mol⁻¹), elevated CO₂ induced photosynthetic downregulation, regardless of temperature and drought. Intercellular CO₂ concentration measurements revealed that photosynthetic acclimation could not be accounted for by stomatal limitations. Downregulation of plants grown in elevated CO₂ was a consequence of decreased carboxylation efficiency as a result of reduced rubisco activity and protein content; in plants grown at ambient temperature, downregulation was also induced by decreased quantum efficiency. The decrease in rubisco activity was associated with carbohydrate accumulation and depleted nitrogen availability. The root nodules were not sufficiently effective to balance the source–sink relation in elevated CO₂ treatments and to provide the required nitrogen to counteract photosynthetic acclimation.     

Effect of elevated CO2 on carbon partitioning and exudate release from Plantago lanceolata seedlings

Plantago lanceolata L. seedlings were grown in sand microcosm units over a 43‐day experimental period under two CO₂ regimes (800 or 400 µmol mol⁻¹) to investigate the effect of elevated atmospheric CO₂ concentration on carbon partitioning and exudate release. Total organic carbon (TOC) content of the collected exudate material was measured throughout the experimental period. After 42 days growth the seedlings were labelled with [¹⁴C]‐CO₂ and the fate of the label within the plant and its release by the roots monitored. Elevated CO₂ significantly (P ≤ 0.001) enhanced shoot, root and total dry matter production although the R:S ratio was unaltered, suggesting no alteration in gross carbon partitioning. The cumulative release of TOC (in mg C) over 0‐42 days was unaltered by CO₂ treatment however, when expressed as a percentage of net assimilated C, ambient‐grown plants released a significantly (P≤ 0.001) higher percentage from their roots compared to elevated CO₂‐grown plants (i.e. 8 vs 3%). The distribution of ¹⁴C‐label was markedly altered by CO₂ treatment with significantly (P≤ 0.001) greater per cent label partitioned to the roots under elevated CO₂. This indicates increased partitioning of recent assimilate below‐ground under elevated CO₂ treatment although there was no significant difference in the percentage of ¹⁴C‐label released by the roots. Comparison of plant C budgets based on ¹⁴C‐pulse‐chase methodology and TOC measurements is discussed.     

Biomass Production and Carbohydrate Content of Arabidopsis thaliana at Atmospheric CO2 Concentrations from 390 to 1680 μl l-1

Abstract: The concentration dependency of the impact of elevated atmospheric CO₂ concentrations on Arabidopsis thaliana L. was studied. Plants were exposed to nearly ambient (390), 560, 810, 1240 and 1680 μl I^-1 CO₂ during the vegetative growth phase for 8 days. Shoot biomass production and dry matter content were increased upon exposure to elevated CO₂. Maximal increase in shoot fresh and dry weight was obtained at 560 μl I^-1 CU₂, which was due to a transient stimulation of the relative growth rate for up to 3 days. The shoot starch content increased with increasing CO₂ concentrations up to two-fold at 1680 μl I^-1 CO₂, whereas the contents of soluble sugars and phenolic compounds were hardly affected by elevated CO₂. The chlorophyll and carotenoid contents were not substantially affected at elevated CO₂ and the chlorophyll a/b ratio remained unaltered. There was no acclimation of photosynthesis at elevated CO₂; the photosynthetic capacity of leaves, which had completely developed at elevated CO₂ was similar to that of leaves developed in ambient air. The possible consequences of an elevated atmospheric CO₂ concentration to Arabidopsis thaliana in its natural habitat is discussed.     

Elevated CO2 and ozone reduce nitrogen acquisition by Pinus halepensis from its mycorrhizal symbiont

The effects of 700 μmol mol⁻¹ CO₂ and 200 nmol mol⁻¹ ozone on photosynthesis in Pinus halepensis seedlings and on N translocation from its mycorrhizal symbiont, Paxillus involutus, were studied under nutrient-poor conditions. After 79 days of exposure, ozone reduced and elevated CO₂ increased net assimilation rate. However, the effect was dependent on daily accumulated exposure. No statistically significant differences in total plant mass accumulation were observed, although ozone-treated plants tended to be smaller. Changes in atmospheric gas concentrations induced changes in allocation of resources: under elevated ozone, shoots showed high priority over roots and had significantly elevated N concentrations. As a result of different shoot N concentration and net carbon assimilation rates, photosynthetic N use efficiency was significantly increased under elevated CO₂ and decreased under ozone. The differences in photosynthesis were mirrored in the growth of the fungus in symbiosis with the pine seedlings. However, exposure to CO₂ and ozone both reduced the symbiosis-mediated N uptake. The results suggest an increased carbon cost of symbiosis-mediated N uptake under elevated CO₂, while under ozone, plant N acquisition is preferentially shifted towards increased root uptake.     

Elevated atmospheric CO2 alters wood production, wood quality and wood strength of Scots pine (Pinus sylvestris L) after three years of enrichment

Scots pine ( Pinus sylvestris L.) trees were grown in open top chambers for three years under ambient and elevated CO₂ concentrations. The trees were aged 3 y at the beginning of the CO₂ exposure, and the effects of the treatment on total stem volume, stem wood biomass, wood quality and wood anatomy were examined at the end of the exposure. The elevated CO₂ treatment lead to a 49% and 38% increase in stem biomass and stem wood volume, respectively. However, no significant effects of the elevated CO₂ treatment on wood density were observed, neither when green wood density was estimated from stem biomass and stem volume, nor when oven-dry wood density was measured on small wood samples. Under elevated CO₂ significantly wider growth rings were observed. The effect of elevated CO₂ on growth ring width was primarily the result of an increase in earlywood width. Wood compression strength decreased under elevated CO₂ conditions, which could be explained by significantly larger tracheids and the increased earlywood band, that has thinner walls and larger cavities. A significant decrease of the number of resin canals in the third growth ring was observed under the elevated treatment; this might indicate that trees produced and contained less resin, which has implications for disease and pest resistance. So, although wood volume yield in Scots pine increased significantly with elevated CO₂ after three years of treatment, wood density remained unchanged, while wood strength decreased. Whilst wood volume and stem biomass production may increase in this major boreal forest tree species, wood quality and resin production might decrease under future elevated CO₂ conditions.     

Elevated CO2 does not ameliorate effects of ozone on carbon allocation in Pinus halepensis and Betula pendula in symbiosis with Paxillus involutus

The effect of 700 μmol CO₂ mol⁻¹, 200 nmol ozone mol⁻¹ and a combination of the two on carbon allocation was examined in Pinus halepensis co-cultured with Betula pendula in symbiosis with the ectomycorrhizal fungus Paxillus involutus . The results show that under low nutrient and ozone levels, elevated CO₂ has no effect on the growth of B. pendula or P. halepensis seedlings nor on net carbon partitioning between plant parts. Elevated CO₂ did not enhance the growth of the fungus in symbiosis with the birch. On the other hand, ozone had a strong negative effect on the growth of the birch, which corresponded with the significantly reduced growth rates of the fungus. Exposure to elevated CO₂ did not ameliorate the negative effects of ozone on birch; in contrast, it acted as an additional stress factor. Neither ozone nor CO₂ had significant effects on biomass accumulation in the pine seedlings. Ozone stimulated the spread of mycorrhizal infection from the birch seedlings to neighbouring pines and had no statistically significant effects on phosphoenolpyruvate carboxylase (PEPC) or ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity in the pine needles or on PEPC activity in pine roots.     

Water vapour fluxes and their impact under elevated CO2 in a C4-tallgrass prairie

We measured leaf-level stomatal conductance, xylem pressure potential, and stomate number and size as well as whole plant sap flow and canopy-level water vapour fluxes in a C4-tallgrass prairie in Kansas exposed to ambient and elevated CO₂. Stomatal conductance was reduced by as much as 50% under elevated CO₂ compared to ambient. In addition, there was a reduction in stomate number of the C4 grass, Andropogon gerardii Vitman, and the C3 dicot herb, Salvia pitcheri Torr., under elevated CO₂ compared to ambient. The result was an improved water status for plants exposed to elevated CO₂ which was reflected by a less negative xylem pressure potential compared to plants exposed to ambient CO₂. Sap flow rates were 20 to 30% lower for plants exposed to elevated CO₂ than for those exposed to ambient CO₂. At the canopy level, evapotranspiration was reduced by 22% under elevated CO₂. The reduced water use by the plant canopy under elevated CO₂ extended the photosynthetically-active period when water became limiting in the ecosystem. The result was an increased above- and belowground biomass production in years when water stress was frequent.     

Warming and elevated CO2 affect the relationship between seed mass, germinability and seedling growth in Austrodanthonia caespitosa, a dominant Australian grass

While the influence of elevated CO₂ on the production, mass and quality of plant seeds has been well studied, the effect of warming on these characters is largely unknown; and there is practically no information on possible interactions between warming and elevated CO₂, despite the importance of these characters in population maintenance and recovery. Here, we present the impacts of elevated CO₂ and warming, both in isolation and combination, on seed production, mass, quality, germination success and subsequent seedling growth of Austrodanthonia caespitosa , a dominant temperate C₃ grass from Australia, using seeds collected from the TasFACE experiment. Mean seed production and mass were not significantly affected by either elevated CO₂ or warming, but elevated CO₂ more than doubled the proportion of very light, inviable seeds ( P < 0.05) and halved mean seed N concentration ( P < 0.04) and N content ( P < 0.03). The dependence of seed germination success on seed mass was affected by an elevated CO₂× warming interaction ( P < 0.004), such that maternal exposure to elevated CO₂ or warming reduced germination if applied in isolation, but not when applied in combination. Maternal effects were retained when seedlings were grown in a common environment for 6 weeks, with seedlings descended from warmed plants 20% smaller ( P < 0.008) with a higher root : shoot ratio ( P < 0.001) than those from unwarmed plants. Given that both elevated CO₂ and warming reduced seed mass, quality, germinability or seedling growth, it is likely that global change will reduce population growth or distribution of this dominant species.     

Adaptation to high CO2 concentration in an optimal environment: radiation capture, canopy quantum yield and carbon use efficiency

The effect of elevated [CO₂] on wheat (Triticum aestivum L. Veery 10) productivity was examined by analysing radiation capture, canopy quantum yield, canopy carbon use efficiency, harvest index and daily C gain. Canopies were grown at either 330 or 1200 μ mol mol^–1[CO₂] in controlled environments, where root and shoot C fluxes were monitored continuously from emergence to harvest. A rapidly circulating hydroponic solution supplied nutrients, water and root zone oxygen. At harvest, dry mass predicted from gas exchange data was 102·8 ± 4·7% of the observed dry mass in six trials. Neither radiation capture efficiency nor carbon use efficiency were affected by elevated [CO₂], but yield increased by 13% due to a sustained increase in canopy quantum yield. CO₂ enrichment increased root mass, tiller number and seed mass. Harvest index and chlorophyll concentration were unchanged, but CO₂ enrichment increased average life cycle net photosynthesis (13%, P < 0·05) and root respiration (24%, P < 0·05). These data indicate that plant communities adapt to CO₂ enrichment through changes in C allocation. Elevated [CO₂] increases sink strength in optimal environments, resulting in sustained increases in photosynthetic capacity, canopy quantum yield and daily C gain throughout the life cycle.     

Growth and physiological responses of canola (Brassica napus) to UV-B and CO2 under controlled environment conditions

Few studies have investigated the interaction of ultraviolet (UV)-B radiation and CO₂ concentration on plants. We studied the combined effects of UV-B radiation and CO₂ concentration on canola ( Brassica napus cv. 46A65) under four growth conditions – ambient CO₂ with UV-B (control), elevated CO₂ with UV-B, ambient CO₂ without UV-B, and elevated CO₂ without UV-B – to determine whether the adverse effects of UV-B are mitigated by elevated CO₂. Elevated CO₂ significantly increased plant height and seed yield, whereas UV-B decreased them. Elevated CO₂ ameliorated the adverse effects of UV-B in plant height. UV-B did not affect the physical characteristics of leaf but CO₂ did. Certain flower and fruit characteristics were affected negatively by UV-B and positively by CO₂. UV-B did not affect net photosynthesis, transpiration and stomatal conductance but decreased water use efficiency (WUE). Elevated CO₂ significantly increased net photosynthesis and WUE. Neither UV-B nor CO₂ affected chlorophyll (Chl) fluorescence. UV-B significantly decreased Chl b and increased the ratio of Chl a / b . Elevated CO₂ decreased only the ratio of Chl a / b . UV-B significantly increased UV-absorbing compounds while CO₂ had no effect on them. Both UV-B and CO₂ significantly increased epicuticular wax content. Many significant relationships were found between morphological, physiological, and chemical parameters. This study showed that elevated CO₂ can partially ameliorate some of the adverse effects of UV-B radiation in B . napus .