Effects of increasing UV-B radiation and atmospheric CO2 on photosynthesis and growth: implications for terrestrial ecosystems

Increases in UV-B radiation reaching the earth as a result of stratospheric ozone depletion will most likely accompany increases in atmospheric CO₂ concentrations. Many studies have examined the effects of each factor independently, but few have evaluated the combined effects of both UV-B radiation and elevated CO₂. In general the results of such studies have shown independent effects on growth or seed yield. Although interspecific variation is large, high levels of UV-B radiation tends to reduce plant growth in sensitive species, while CO₂ enrichment tends to promote growth in most C₃ species. However, most previous studies have not looked at temporal effects or at the relationship between photosynthetic acclimation to CO₂ and possible photosynthetic limitations imposed by UV-B radiation. Elevated CO₂ may provide some protection against UV-B for some species. In contrast, UV-B radiation may limit the ability to exploit elevated CO₂ in other species. Interactions between the effects of CO₂ enrichment and UV-B radiation exposure have also been shown for biomass allocation. Effects on both biomass allocation and photosynthetic acclimation may be important to ecosystem structure in terms of seedling establishment, competition and reproductive output. Few studies have evaluated ecosystem processes such as decomposition or nutrient cycling. Interactive effects may be subtle and species specific but should not be ignored in the assessment of the potential impacts of increases in CO₂ and UV-B radiation on plants.     

Effects of UV-B radiation on terrestrial plants and ecosystems: interaction with CO2 enrichment

UV-B radiation is just one of the environmental factors, that affect plant growth. It is now widely accepted that realistic assessment of plant responses to enhanced UV-B should be performed at sufficiently high Photosynthetically Active Radiation (PAR), preferably under field conditions. This will often imply, that responses of plants to enhanced UV-B in the field will be assessed under simultaneous water shortage, nutrient deficiency and variation of temperature. Since atmospheric CO₂ enrichment, global warming and increasing UV-B radiation represent components of global climatic change, interactions of UV-B with CO₂ enrichment and temperature are particularly relevant. Only few relevant UV-B× CO₂ interaction studies have been published. Most of these studies refer to greenhouse experiments. We report a significant CO₂ × UV-B interaction for the total plant dry weight and root dry weight of the C₃-grass Elymus athericus. At elevated CO₂ (720 mol mol^-1, plant growth was much less reduced by enhanced UV-B than at ambient atmospheric CO₂ although there were significant (positive) CO₂ effects and (negative) UV-B effects on plant growth. Most other CO₂ × UV-B studies do not report significant interactions on total plant biomass. This lack of CO₂ × UV-B interactions may result from the fact that primary metabolic targets for CO₂ and UVB are different. UV-B and CO₂ may differentially affect plant morphogenetic parameters: biomass allocation, branching, flowering, leaf thickness, emergence and senescence. Such more subtle interactions between CO₂ and UV-B need careful and long term experimentation to be detected. In the case of no significant CO₂× UV-B interactions, combined CO₂ and UV-B effects will be additive. Plants differ in their response to CO₂ and UV-B, they respond in general positively to elevated CO₂ and negatively to enhanced UV-B. Moreover, plant species differ in their responsiveness to CO₂ and UV-B. Therefore, even in case of additive CO₂ and UV-B effects, plant competitive relationships may change markedly under current climatic change with simultaneous enhanced atmospheric CO₂ and solar UV-B radiation.     

The combined effects of elevated CO2 levels and UV-B radiation on growth characteristics ofElymus athericus (= E. pycnanathus)

Elymus athericus (Link) Kerguélen, a C₃ grass, was grown in a greenhouse experiment to determine the effect of enhanced atmospheric CO₂ and elevated UV-B radiation levels on plant growth. Plants were subjected to the following treatments; a) ambient CO₂-control UV-B, b) ambient CO₂-elevated UV-B, c) double CO₂-control UV-B, d) double CO₂-elevated UV-B. Elevated CO₂ concentrations stimulated plant growth, biomass production was 67% higher than at ambient CO₂. Elevated UV-B radiation had a negative effect on growth, biomass production was depressed by 31%. Enhanced CO₂ combined with elevated UV-B levels caused a biomass depression of 8% when compared with the control plants. UV-B induced growth depression can be modified by a growth stimulus caused by high CO₂ concentrations. Growth analysis has been performed and possible physiological mechanisms behind changing growth parameters are discussed.     

The combined effects of CO2 concentration and solar UV-B radiation on faba bean grown in open-top chambers

The response of faba bean seedlings to the combined effects of increased atmospheric CO2 concentrations ([CO2]) and solar UV-B irradiance was studied using open-top chambers transparent to UV-B radiation. The purpose of the study was to determine whether effects of increased [CO2] on growth and physiology are modified by the present solar UV-B fluence rate in the Netherlands. Seedlings were exposed to 350 or 700 micromoles mol-1 CO2. At both [CO2], solar UV-B irradiance was either present or reduced using polyester foil opaque to UV-B radiation. To obtain information on the time dependence of increased [CO2] and UV-B radiation effects, three harvests were performed during the experiment. CO2 enrichment resulted in increased biomass production at all harvests. At the final harvest, UV-B radiation did not affect biomass production but a significant decrease was observed after 14 d of treatment. A reduction of the UV-B fluence increased shoot length at both [CO2] throughout the experiment. UV-B radiation slightly altered biomass allocation. Plants grown at reduced levels of UV-B radiation invested less biomass in flowers and more in stem material compared to plants grown at ambient UV-B levels. CO2 enrichment resulted in a stimulation of net photosynthesis after 26 and 38 d of treatment. UV-B reduction did not alter this response. After 26 d of treatment, photosynthetic acclimation to CO2 enrichment was observed, which was probably the result of accumulation of carbohydrates in the leaves. After 38 d, photosynthetic acclimation was no longer present. The UV absorbance of methanolic leaf extracts was increased by CO2 enrichment only. Both CO2 enrichment and solar UV-B reduced the transmittance of radiation through intact attached leaves. Interaction between [CO2] and UV-B radiation was limited to UV-A transmittance of leaves. Under prevalent experimental conditions, UV-B radiation did not affect the measured physiological parameters. Most open-top chambers used for climate change research are constructed of materials which do not transmit UV-B radiation. Our results indicate that part of the 'chamber effects' on plant height often described in the literature might be explained by the absence of solar UV-B radiation in these chambers.     

The effects of UV-B radiation on loblolly pine. 3. Interaction with CO2 enhancement

Projected depletions in the stratospheric ozone layer will result in increases in solar ultraviolet-B radiation (290–320 nm) reaching the earth's surface, These increases will likely occur in concert with other environmental changes such as increases in atmospheric carbon dioxide concentrations. Currently very little information is available on the effectiveness of UV-B radiation within a CO₂-enriched atmosphere, and this is especially true for trees. Loblolly pine (Pinus taeda L.) seedlings were grown in a factorial experiment at the Duke University Phytotron with either 0, 8.8 or 13.8 kJ m⁻² of biologically effective UV-B radiation (UV-B_BE). The CO₂ concentrations used were 350 and 650 μmol mol⁻¹. Measurements of chlorophyll fluorescence were made at 5-week intervals and photosynthetic oxygen evolution and leaf pigments were measured after 22 weeks, prior to harvest. The results of this study demonstrated a clear growth response to CO₂ enrichment but neither photosynthetic capacity nor quantum efficiency were altered by CO₂. The higher UV-B irradiance reduced total biomass by about 12% at both CO₂ levels but biomass partitioning was altered by the interaction of CO₂ and UV-B radiation. Dry matter was preferentially allocated to shoot components by UV-B radiation at 350 μmol mol⁻¹ CO₂ and towards root components at 650 μmol mol⁻¹ CO₂. These subtle effects on biomass allocation could be important in the future to seedling establishment and competitive interactions in natural as well as agricultural communities.     

Plant responses to atmospheric carbon dioxide enrichment: interactions with some soil and atmospheric conditions

In general, C₃ plant species are more responsive to atmospheric carbon dioxide (CO₂) enrichment than C₄-plants. Increased relative growth rate at elevated CO₂ primarily relates to increased Net Assimilation Rate (NAR), and enhancement of net photosynthesis and reduced photorespiration. Transpiration and stomatal conductance decrease with elevated CO₂, water use efficiency and shoot water potential increase, particularly in plants grown at high soil salinity. Leaf area per plant and leaf area per leaf may increase in an early growth stage with increased CO₂, after a period of time Leaf Area Ratio (LAR) and Specific Leaf Area (SLA) generally decrease. Starch may accumulate with time in leaves grown at elevated CO₂. Plants grown under salt stress with increased (dark) respiration as a sink for photosynthates, may not show such acclimation to increased atmospheric CO₂ levels. Plant growth may be stimulated by atmospheric carbon dioxide enrichment and reduced by enhanced UV-B radiation but the limited data available on the effect of combined elevated CO₂ and ultraviolet B (280–320 nm) (UV-B) radiation allow no general conclusion. CO₂-induced increase of growth rate can be markedly modified at elevated UV-B radiation. Plant responses to elevated atmospheric CO₂ and other environmental factors such as soil salinity and UV-B tend to be species-specific, because plant species differ in sensitivity to salinity and UV-B radiation, as well as to other environmental stress factors (drought, nutrient deficiency). Therefore, the effects of joint elevated atmospheric CO₂ and increased soil salinity or elevated CO₂ and enhanced UV-B to plants are physiologically complex.     

The combined effects of CO2 concentration and enhanced UV-B radiation on faba bean. 3. Leaf optical properties,pigments, stomatal index and epidermal cell density

Seedlings of Vicia faba L. (cv. Minica) were grown in a factorial experiment in a greenhouse. The purpose of the study was to determine whether CO₂ enrichment and supplemental UV-B radiation affect leaf optical properties and whether the combined effects differ from single factor effects. Seedlings were grown at either 380 mol mol^-1 or 750 mol mol^-1 CO₂ and at four levels of UV-B radiation. After 20 and 40 days of treatment, absorptance, transmittance and reflectance of photosynthetically active radiation (PAR) were measured on the youngest fully developed leaf. On the same leaf, the specific leaf area on a fresh weight basis (SLA_fw), chlorophyll content, UV-B absorbance, transmittance of UV light and stomatal index were measured. UV-B radiation significantly increased PAR absorptance and decreased PAR transmittance. The increased PAR absorptance can be explained by an increased chlorophyll content in response to UV-B radiation. Leaf transmittance of UV radiation decreased with increasing UV-B levels mainly caused by increased absorbance of UV absorbing compounds. UV-B radiation decreased both the stomatal density and epidermal cell density of the abaxial leaf surface, leaving the stomatal index unchanged. Effects of CO₂ enrichment were less pronounced than those of UV-B radiation. The most important CO₂ effect was an increase in stomatal density and epidermal cell density of the adaxial leaf surface. The stomatal index was not affected. No interaction between CO₂ and UV-B radiation was found. The results are discussed in relation to the internal light environment of the leaf.     

The effects of UV-B radiation on European heathland species

The effects of enhanced UV-B radiation on three examples of European shrub-dominated vegetation were studied in situ. The experiments were in High Arctic Greenland, northern Sweden and Greece, and at all sites investigated the interaction of enhanced UV-B radiation (simulating a 15% reduction in the ozone layer) with artificially increased precipitation. The Swedish experiment also involved a study of the interaction between enhanced UV-B radiation and elevated CO₂ (600 ppm). These field studies were supported by an outdoor controlled environment study in the United Kingdom involving modulated enhancement of UV-B radiation in combination with elevated CO₂ (700 ppm). Effects of the treatments on plant growth, morphology, phenology and physiology were measured. The effects observed were species specific, and included both positive and negative responses to the treatments. In general the negative responses to UV-B treatments of up to three growing seasons were small, but included reductions in shoot growth and premature leaf senescence. Positive responses included a marked increase in flowering in some species and a stimulation of some photosynthetic processes. UV-B treatment enhanced the drought tolerance of Pinus pinea and Pinus halepensis by increasing leaf cuticle thickness. In general, there were few interactions between the elevated CO₂ and enhanced UV-B treatments. There was evidence to suggest that although the negative responses to the treatments were small, damage may be increasing with time in some long-lived woody perennials. There was also evidence in the third year of treatments for effects of UV-B on insect herbivory in Vaccinium species. The experiments point to the necessity for long-term field investigations to predict the likely ecological consequences of increasing UV-B radiation.     

Effects of climate change on productivity of cereals and legumes; model evaluation of observed year-to-year variability of the CO2 response

The effect of elevated [CO₂] on the productivity of spring wheat, winter wheat and faba bean was studied in experiments in climatized crop enclosures in the Wageningen Rhizolab in 1991–93. Simulation models for crop growth were used to explore possible causes for the observed differences in the CO₂ response. Measurements of the canopy gas exchange (CO₂ and water vapour) were made continuously from emergence until harvest. At an external [CO₂] of 700 μmol mol^?1 Maximum Canopy CO₂ Exchange Rate (CCERmax) at canopy closure was stimulated by 51% for spring wheat and by 71% for faba bean. At the end of the growing season, above ground biomass increase at 700 μmol mol^?1 was 58% (faba bean), 35% (spring wheat) and 19% (winter wheat) and the harvest index did not change. For model exploration, weather data sets for the period 1975-88 and 1991–93 were used, assuming adequate water supply and [CO₂] at 350 and 700 μmol mol^?1. For spring wheat the simulated responses (35–50%) were at the upper end of the experimental results. In agreement with experiments, simulations showed smaller responses for winter wheat and larger responses for faba bean. Further model explorations showed that this differential effect in the CO₂ response may not be primarily due to fundamental physiological differences between the crops, but may be at least partly due to differences in the daily air temperatures during comparable stages of growth of these crops. Simulations also showed that variations between years in CO₂ response can be largely explained by differences in weather conditions (especially temperature) between growing seasons.     

Enhanced UV-B radiation alleviates the adverse effects of summer drought in two Mediterranean pines under field conditions

The effects of enhanced UV-B (290-320 nm) radiation on two native Mediterranean pines (Pinus pinea L., Pinus halepensis Mill.) were recorded during a one-year field study. Plants received ambient or ambient plus supplemental UV-B radiation (simulating a 15% stratospheric ozone depletion over Patras. Greece, 38.3°N. 29.1°E) and only natural precipitation, i.e. they were simultaneously exposed to other natural stresses. particularly water stress during summer. Supplemental UV-B irradiation started in early February, 1993 and up to late June, no effects were observed on growth and photochemical efficiency of photosystem II, as measured by chlorophy II fluorescence induction. Water stress during the summer was manifested in the control plants as a decline in the ratio of variable to maximum fluorescence (F_v/F_m), the apparent photon yield for oxygen evolution (φ_l) and the photosynthetic capacity at 5% CO₂ (P_m). In addition, a partial needle loss was evident. Under supplemental UV-B radiation, however, the decreases in F_v/F_m, φ_i, and P_m. as well as needle losses were significantly less. Soon after the first heavy autumn rains. photosynthetic parameters in both control and UV-B treated plants recovered to similar values. but the transient summer superiority of UV-B irradiated plants resulted in a significant increase in their dry weight measured at plant harvest. during late January. 1994. Plant height. UV-B absorbing compounds, photosynthetic pigments and relative water content measured at late spring. late summer and at plant harvest, were not significantly affected by supplemental UV-B radiation. The results indicate that enhanced UV-B radiation may be beneficial for Mediterranean pines through a partial alleviation of the adverse effects of summer drought.     

Intraspecific variation in sensitivity to ultraviolet-B radiation in endogenous hormones and photosynthetic characteristics of 10 wheat cultivars grown under field conditions

Field studies were conducted to determine the potential of altering endogenous hormones and photosynthetic characteristics and intraspecific variation in sensitivity of 10 wheat (Triticum aestivum) cultivars (four tolerant, two middle sensitive and four sensitive) to enhanced ultraviolet-B (UV-B, 280–315 nm) radiation under field conditions. The supplemental UV-B radiation was 5.00 kJ m⁻², simulating a depletion of 20% stratospheric ozone. Responses were cultivar-specific. Out of the 10 tested wheat cultivars, six showed significant decrease in IAA content. UV-B radiation significantly increased ZR content in two wheat cultivars and significantly decreased in five cultivars. ABA content of three wheat cultivars was increased significantly, while that of five cultivars was decreased significantly. UV-B radiation significantly increased the stomatal conductance of three cultivars, and significantly decreased that of four cultivars. Intercellular CO₂ concentrations were significantly increased in five cultivars and significantly decreased in one cultivar (Mianyang 20). Transpiration rate of three cultivars significantly increased, while that of three cultivars significantly decreased. UV-B radiation significantly decreased the net photosynthetic rate of six cultivars. Intraspecific differences were found for the different measured parameters. For seven measured parameters, UV-B radiation had significant effects on five wheat cultivars, while no effect on the others. Significant correlations were observed between net photosynthetic rate and stomatal conductance, intercellular CO₂ concentrations and transpiration rate in eight cultivars. UV-B radiation might change stomatal conductance, intercellular CO₂ concentrations and transpiration rate, thus resulting in changes in net photosynthetic rate.     

Effects of elevated [CO2] at the community level mediated by root symbionts

This review examines the effects of elevated [CO₂] on plant symbioses with mycorrhizal fungi and root nodule bacteria, with emphasis on community and ecosystem processes. The effects of elevated [CO₂] on the relationships between single plant species and root symbionts are considered first. There is some evidence that plant infection by and/or biomass of root symbionts are stimulated by elevated [CO₂], but growth enhancement of the host seemingly depends on its degree of dependence on symbiosis and on soil nutrient availability. Second, the effects of elevated [CO₂] on the relationships between plant multispecies assemblages and soil, and likely impacts on above-ground and belowground diversity, are analysed. Experimental and modelling work have suggested the existence of complex feedbacks in the responses of plants and the rhizosphere to CO₂ enrichment. By modifying C inputs from plants to soil, elevated [CO₂] may affect the biomass, the infectivity, and the species/isolate composition of root symbionts. This has the potential to alter community structure and ecosystem functioning. Finally, the incorporation of type and degree of symbiotic dependence into the definition of plant functional types, and into experimental work within the context of global change research, are discussed. More experimental work on the effects of elevated [CO₂] at the community/ecosystem level, explicitly considering the role of root symbioses, is urgently needed.     

CO(2) Enhancement of Growth and Photosynthesis in Rice (Oryza sativa) : Modification by Increased Ultraviolet-B Radiation

Two cultivars of rice (Oryza sativa L.) IR-36 and Fujiyama-5 were grown at ambient (360 microbars) and elevated CO₂ (660 microbars) from germination through reproduction in unshaded greenhouses at the Duke University Phytotron. Growth at elevated CO₂ resulted in significant decreases in nighttime respiration and increases in photosynthesis, total biomass, and yield for both cultivars. However, in plants exposed to simultaneous increases in CO₂ and ultraviolet-B (UV-B) radiation, CO₂ enhancement effects on respiration, photosynthesis, and biomass were eliminated in IR-36 and significantly reduced in Fujiyama-5. UV-B radiation simulated a 25% depletion in stratospheric ozone at Durham, North Carolina. Analysis of the response of CO₂ uptake to internal CO₂ concentration at light saturation suggested that, for IR-36, the predominant limitation to photosynthesis with increased UV-B radiation was the capacity for regeneration of ribulose bisphosphate (RuBP), whereas for Fujiyama-5 the primary photosynthetic decrease appeared to be related to a decline in apparent carboxylation efficiency. Changes in the RuBP regeneration limitation in IR-36 were consistent with damage to the photochemical efficiency of photosystem II as estimated from the ratio of variable to maximum chlorophyll fluorescence. Little change in RuBP regeneration and photochemistry was evident in cultivar Fujiyama-5, however. The degree of sensitivity of photochemical reactions with increased UV-B radiation appeared to be related to leaf production of UV-B-absorbing compounds. Fujiyama-5 had a higher concentration of these compounds than IR-36 in all environments, and the production of these compounds in Fujiyama-5 was stimulated by UV-B fluence. Results from this study suggest that in rice alterations in growth or photosynthesis as a result of enhanced CO₂ may be eliminated or reduced if UV-B radiation continues to increase.     

No cumulative effect of 10 years of elevated [CO2] on perennial plant biomass components in the Mojave Desert

Elevated atmospheric CO₂ concentrations ([CO₂]) generally increase primary production of terrestrial ecosystems. Production responses to elevated [CO₂] may be particularly large in deserts, but information on their long‐term response is unknown. We evaluated the cumulative effects of elevated [CO₂] on primary production at the Nevada Desert FACE (free‐air carbon dioxide enrichment) Facility. Aboveground and belowground perennial plant biomass was harvested in an intact Mojave Desert ecosystem at the end of a 10‐year elevated [CO₂] experiment. We measured community standing biomass, biomass allocation, canopy cover, leaf area index (LAI), carbon and nitrogen content, and isotopic composition of plant tissues for five to eight dominant species. We provide the first long‐term results of elevated [CO₂] on biomass components of a desert ecosystem and offer information on understudied Mojave Desert species. In contrast to initial expectations, 10 years of elevated [CO₂] had no significant effect on standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground components. However, elevated [CO₂] increased short‐term responses, including leaf water‐use efficiency (WUE) as measured by carbon isotope discrimination and increased plot‐level LAI. Standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground pools significantly differed among dominant species, but responses to elevated [CO₂] did not vary among species, photosynthetic pathway (C₃ vs. C₄), or growth form (drought‐deciduous shrub vs. evergreen shrub vs. grass). Thus, even though previous and current results occasionally show increased leaf‐level photosynthetic rates, WUE, LAI, and plant growth under elevated [CO₂] during the 10‐year experiment, most responses were in wet years and did not lead to sustained increases in community biomass. We presume that the lack of sustained biomass responses to elevated [CO₂] is explained by inter‐annual differences in water availability. Therefore, the high frequency of low precipitation years may constrain cumulative biomass responses to elevated [CO₂] in desert environments.     

Contrasting crop species responses to CO2 and temperature: rice,soybean and citrus

The continuing increase in atmospheric carbon dioxide concentration ([CO₂]) and projections of possible future increases in global air temperatures have stimulated interest in the effects of these climate variables on plants and, in particular, on agriculturally important food crops. Mounting evidence from many different experiments suggests that the magnitude and even direction of crop responses to [CO₂] and temperature is almost certain to be species dependent and very likely, within a species, to be cultivar dependent. Over the last decade, [CO₂] and temperature experiments have been conducted on several crop species in the outdoor, naturally-sunlit, environmentally controlled, plant growth chambers by USDA-ARS and the University of Florida, at Gainesville, Florida, USA. The objectives for this paper are to summarize some of the major findings of these experiments and further to compare and contrast species responses to [CO₂] and temperature for three diverse crop species: rice (Oryza sativa, L.), soybean (Glycine max, L.) and citrus (various species). Citrus had the lowest growth and photosynthetic rates but under [CO₂] enrichment displayed the greatest percentage increases over ambient [CO₂] control treatments. In all three species the direct effect of [CO₂] enrichment was always an increase in photosynthetic rate. In soybean, photosynthetic rate depended on current [CO₂] regardless of the long-term [CO₂] history of the crop. In rice, photosynthetic rate measured at a common [CO₂], decreased with increasing long-term [CO₂] growth treatment due to a corresponding decline in RuBP carboxylase content and activity. Rice specific respiration decreased from subambient to ambient and superambient [CO₂] due to a decrease in plant tissue nitrogen content and a decline in specific maintenance respiration rate. In all three species, crop water use decreased with [CO₂] enrichment but increased with increases in temperature. For both rice and soybean, [CO₂] enrichment increased growth and grain yield. Rice grain yields declined by roughly 10 % per each 1 °C rise in day/night temperature above 28/21 °C.     

Effects of enhanced ultraviolet radiation and carbon dioxide concentration on the moss Hylocomium splendens

In a laboratory experiment interaction effects of UV-B and CO₂ on photosynthesis and growth of the moss Hylocomium splendens were studied. The plants were exposed to two CO₂ levels (350 ppm and 600 ppm) and three UV-B levels (no UV-B, ambient UV-B and that corresponding to 20% ozone depletion) for 5 months. The effects were recorded by measuring the photosynthetic response and growth of the plants. There was a statistically significant change in photosynthetic efficiency and maximum photosynthetic rates due to time and to enhanced CO₂ concentration, whereas there was no effect due to UV-B. There was a decreased growth due to both UV-B and CO₂ and an interaction effect on growth (in length). The UV-B dose corresponding to the ambient level had a larger reducing effect on growth than the highest UV-B dose. This was a counter-intuitive result and the following tentative interpretation was made: differences in the measured UV-A/UV-B/PAR ratios between the treatments could explain the result provided there was a non-linear response to UV over the range of irradiance levels used.     

Effects of solar ultraviolet-B radiation,temperature and CO2 on growth and physiology of sunflower and maize seedlings

The effects of solar UV-B radiation, in combination with elevated temperature (4 °C ) and CO₂ (680 L L^-1 concentration, on sunflower and maize seedlings were studied from May to August in 1991 at the research station Quinta de São Pedro in Portugal (38.7°N). The ambient solar radiation of Portugal was reduced to levels of Central European latitudes by using the ozone filter technique. This radiation served as control, while the ambient solar radiation of Portugal was to simulate intense UV-B treatment (+30%). All plants were grown up to 18 days in 4 climate controlled growth chambers simulating a daily course of temperature with T_max=28 °C or 32 °C , resp., and ambient CO₂ concentrations (340 L L^-1); in one chamber the CO₂ concentration was twice as high (680 L L^-1). Under intense UV-B and at 28 °C (T_max) all growth parameters (height, leaf area, fresh and dry weight, stem elongation rate, relative growth rate) of sunflower and maize seedlings were reduced down to 35% as compared to controls. An increase in growing temperature by 4 °C , alone or in combination with doubled CO₂, compensated or even overcompensated the UV-B effect so that the treated plants were comparable to controls. Chlorophyll content, on a leaf area basis, increased under intense UV-B radiation. This increase was compensated by lower leaf areas, resulting in comparable chlorophyll contents. Similar to growth, also the net photosynthetic rates of sunflower and maize seedlings were reduced down to 29% by intense UV-B calculated on a chlorophyll basis. This reduction was compensated by an increased temperature. Doubling of CO₂ concentration had effects only on sunflower seedlings in which the photosynthetic rates were higher than in the controls. Dark respiration rates of the seedlings were not influenced by any experimental condition. Transpiration and water use efficiency (wue) were not influenced by intense UV-B. Higher temperatures led to higher transpiration rates and lower water use efficiencies, resp.. Doubling of CO₂ reduced the transpiration rate drastically while for wue maximum values were recorded.     

Effect of CO2 enrichment and nitrogen availability on resource acquisition and resource allocation in a grass,Bromus mollis

Summary The effects of CO₂ enrichment on the growth, biomass partitioning, photosynthetic rates, and leaf nitrogen concentration of a grass, Bromus mollis (C₃), were investigated at a favorable and a low level of nitrogen availability. Despite increases in root: shoot ratios, leaf nitrogen concentrations were decreased under CO₂ enrichment at both nitrogen levels. For the low-nitrogen treatment, this resulted in lower photosynthetic rates measured at 650 l/l for the CO₂-enriched plants, compared to photosynthetic rates measured at 350 l/l for the non-enriched plants. At higher nitrogen availability, photosynthetic rates of plants grown and measured at 650 l/l were greater than photosynthetic rates of the non-enriched plants measured at 350 l/l. Water use efficiency, however, was increased in enriched plants at both nitrogen levels. CO₂ enrichment stimulated vegetative growth at both high and low nitrogen during most of the vegetative growth phase but, at the end of the experiment, total biomass of the high and low CO₂ treatments did not differ for plants grown at low nitrogen availability. While not statistically significant, CO₂ tended to stimulate seed production at high nitrogen and to decrease it at low nitrogen.     

Compensatory responses of CO2 exchange and biomass allocation and their effects on the relative growth rate of ponderosa pine in different CO2 and temperature regimes

Increases in the concentration of atmospheric carbon dioxide may have a fertilizing effect on plant growth by increasing photosynthetic rates and therefore may offset potential growth decreases caused by the stress associated with higher temperatures and lower precipitation. However, plant growth is determined both by rates of net photosynthesis and by proportional allocation of fixed carbon to autotrophic tissue and heterotrophic tissue. Although CO₂ fertilization may enhance growth by increasing leaf-level assimilation rates, reallocation of biomass from leaves to stems and roots in response to higher concentrations of CO₂ and higher temperatures may reduce whole-plant assimilation and offset photosynthetic gains. We measured growth parameters, photosynthesis, respiration, and biomass allocation of Pinus ponderosa seedlings grown for 2 months in 2×2 factorial treatments of 350 or 650 bar CO₂ and 10/25° C or 15/30° C night/day temperatures. After 1 month in treatment conditions, total seedling biomass was higher in elevated CO₂, and temperature significantly enhanced the positive CO₂ effect. However, after 2 months the effect of CO₂ on total biomass decreased and relative growth rates did not differ among CO₂ and temperature treatments over the 2-month growth period even though photosynthetic rates increased 7% in high CO₂ treatments and decreased 10% in high temperature treatments. Additionally, CO₂ enhancement decreased root respiration and high temperatures increased shoot respiration. Based on CO₂ exchange rates, CO₂ fertilization should have increased relative growth rates (RGR) and high temperatures should have decreased RGR. Higher photosynthetic rates caused by CO₂ fertilization appear to have been mitigated during the second month of exposure to treatment conditions by a 3% decrease in allocation of biomass to leaves and a 9% increase in root:shoot ratio. It was not clear why diminished photosynthetic rates and increased respiration rates at high temperatures did not result in lower RGR. Significant diametrical and potentially compensatory responses of CO₂ exchange and biomass allocation and the lack of differences in RGR of ponderosa pine after 2 months of exposure of high CO₂ indicate that the effects of CO₂ fertilization and temperature on whole-plant growth are determined by complex shifts in biomass allocation and gas exchange that may, for some species, maintain constant growth rates as climate and atmospheric CO₂ concentrations change. These complex responses must be considered together to predict plant growth reactions to global atmospheric change, and the potential of forest ecosystems to sequester larger amounts of carbon in the future.     

Influence of ultraviolet-B (UV-B) radiation on photosynthetic and growth characteristics in field-grown cassava (Manihot esculentum Crantz)

The effects of ultraviolet-B (UV-B between 290 and 320 nm) on photosynthesis and growth characteristics were investigated in field grown cassava (Manihot esculentum Crantz). Plants were grown at ambient and ambient plus a 5.5kJ m^?2 d^?1 supplementation of UV-B radiation for 95 d. The supplemental UV-B fluence used in this experiment simulated a 15% depletion in stratospheric ozone at the equator (0°N). Carbon dioxide exchange, oxygen evolution, and the ratio of variable to maximum fluorescence (F_v/F_m) were determined for fully expanded leaves after 64–76 d of UV-B exposure. AH plants were harvested after 95 d of UV-B exposure, assayed for chlorophyll and UV-B absorbing compounds, and separated into leaves, petioles, stems and roots. Exposure to UV-B radiation had no effect on in situ rates of photosynthesis or dark respiration. No difference in the concentration of UV-B absorbing compounds was observed between treatments. A 2-d daytime diurnal comparison of F_v to F_m ratios indicated a significant decline in F_v/F_m ratios and a subsequent increase in photoinhibition under enhanced UV-B radiation if temperature or PPF exceeded 35°C or 1800μmol m^?2 s^?1, respectively. However, UV-B effects on fluorescence kinetics appeared to be temporal since maximal photosynthetic rates as determined by oxygen evolution at saturated CO₂ and PPF remained unchanged. Although total biomass was unaltered with UV-B exposure, alterations in the growth characteristics of cassava grown with supplemental UV-B radiation are consistent with auxin destruction and reduced apical dominance. Changes in growth included an alteration of biomass partitioning with a significant increase in shoot/root ratio noted for plants receiving supplemental UV-B radiation. The increase in shoot/root ratio was due primarily to a significant decrease in root weight (–32%) with UV-B exposure. Because root production determines the harvest-able portion of cassava, UV-B radiation may still influence the yield of an important tropical agronomic species, even though photosynthesis and total dry biomass may not be directly affected.