The seasonal pattern of CO2 exchange of Festuca rubra L. in a montane meadow community in Northern Germany

Summary Completely climatized cuvettes were used to follow the CO₂ gas exchange of red fescue (Festuca rubra L.), growing on a fertilized and an unfertilized plot, during a growing season from May through October. Objective of the study was to determine the effect of environmental factors on the seasonal CO₂ gas exchange.Gas exchange rates were calculated on the basis of leaf dry weight, surface area and chlorophyll content. Photosynthetic rates differed between the fertilized and unfertilized plants when based on leaf dry weight or leaf surface area but were similar when based on chlorophyll.Multiple regression analysis was used to related photosynthetic rates to radiation, temperature, water vapor concentration difference, chlorophyll content and time. A cubic regression equation based on daily radiation alone explained 85% of the variation for the fertilized plants and 87% of the variation for the unfertilized plants.During the growing season the unfertilized plants had a continual decline in their photosynthetic rates. The fertilized plants had high photosynthetic rates in the spring and in the fall.Light response curves indicated greater photosynthetic rates at light saturation as well as in the light limited portion of the light response curve for the fertilized plants. Photosynthetic rates of the fertilized plants were generally depressed during periods of warm temperature and high light intensity in June and July.Photosynthetic rates declined at temperatures above 24°C. The decline was greater for the more mesomorphic fertilized plants. A similar response was noted to increasing water vapor difference, although it was difficult to separate from the temperature effect. Maximum photosynthetic rates were found between 14°C and 22°C, although there was considerable variation in the maximum rates.The effects of cutting (mowing) on the gas exchange were difficult to determine due to the interaction of the environmental factors.Chlorophyll content showed significant correlation with photosynthetic rates.     

Ecophysiological Significance of CO(2)-Recycling via Crassulacean Acid Metabolism in Talinum calycinum Engelm. (Portulacaceae)

High levels of variability in gas exchange characteristics and degree of CAM-cycling were found in the same and different individuals of Talinum calycinum Engelm. collected from rock outcrops in Missouri. Differences in CO₂ assimilation were mostly correlated with differences in shoot conductance to CO₂ not shoot internal CO₂ concentration. As found previously, CAM acid fluctuations were evident in well-watered plants exhibiting C₃ gas exchange patterns (CAM-cycling) and also in drought-stressed plants with stomata closed, or nearly so, day and night (CAM-idling). Drought stress also resulted in rapid stomatal closure, conserving water during droughts. Maximal CO₂ uptake rates occurred below 35°C; higher temperatures induced decreases in CO₂ assimilation and conductance while shoot internal CO₂ concentrations remained similar. Plant water-use-efficiency was severely curtailed at temperatures above 30°C. Tissue acid fluctuations were the result of changes in malic acid concentrations. Calculations of the amount of water potentially conserved by CAM-cycling yielded values of approximately 5 to 44% of daytime water loss. Thus, CAM-cycling may be an important adaptation minimizing water loss by perennial succulents growing in shallow soil on rock outcrops.     

Dynamics of Photosynthesis in Pine Stands

The CO₂ exchange of pine (Pinus sylvestris L.) shoots was continuously monitored over several years. The spatial and temporal variability of gas exchange was thoroughly investigated for three forest types. The net uptake of CO₂ by current-season shoots in a bilberry pine stand equaled 5.4 g CO₂ per g dry wt for the growing season. The net CO₂ assimilation by one-year-old shoots over a growing season constituted 9.9 and 2.4 g CO₂/(g dry wt year) in the upper and lower crown parts, respectively. The nighttime respiration of the current-season shoots released 0.7 g CO₂/(g dry wt year), and the respiration of one-year-old shoots in the upper and lower parts of the canopy released 0.45 and 0.36 g CO₂/(g dry wt year), respectively. Recalculation of three-year data per entire crown yielded an annual average carbon assimilation of 1.54 g C/(g dry wt year). Water relations markedly affected CO₂ fixation. Nevertheless, the daily average values of photosynthesis in the summer were similar for pine stands with bilberry, heather, and dwarf shrub–polytric as understory dominants. It is shown that the realization of changes in photosynthetic function is related in time to growth processes.     

CO2 effects on apical dominance in Pisum sativum

Alaska pea plants (Pisum sativum L.) were grown at 0.10 vol% and 0.035 vol% CO₂ to determine the effects of high CO₂ concentration upon plant growth and apical dominance. The results showed that a 0.10 vol% CO₂ atmosphere significantly increased the rate of lateral branch, flower bud, flower and fruit development over an environment with 0.035 vol% CO₂. At plant maturity, however, there were no significant differences in the number of branches or fruits produced at the different CO₂ levels. Thus, no evidence was obtained for the loss of apical dominance at the CO₂ concentrations tested. Root dry weight was significantly greater in plants grown at 0.10 vol% CO₂ than in those grown at 0.035 vol% CO₂ and leaf dry weight was significantly lower. However, no significant differences were found in total plant dry weight production at plant maturity.     

Effect of elevated CO<Subscript>2</Subscript> levels on leaf starch,nitrogen and photosynthesis of plants growing at three natural CO<Subscript>2</Subscript> springs in Japan

Plant communities around natural CO₂ springs have been exposed to elevated CO₂ levels over many generations and give us a unique opportunity to investigate the effects of long-term elevated CO₂ levels on wild plants. We searched for natural CO₂ springs in cool temperate climate regions in Japan and found three springs that were suitable for studying long-term responses of plants to elevated levels of CO₂: Ryuzin-numa, Yuno-kawa and Nyuu. At these CO₂ springs, the surrounding air was at high CO₂ concentration with no toxic gas emissions throughout the growth season, and there was natural vegetation around the springs. At each site, high-CO₂ (HC) and low-CO₂ (LC) plots were established, and three dominant species at the shrub layers were used for physiological analyses. Although the microenvironments were different among the springs, dicotyledonous species growing at the HC plots tended to have more starch and less nitrogen per unit dry mass in the leaves than those growing at the LC plots. In contrast, monocotyledonous species growing in the HC and LC plots had similar starch and nitrogen concentrations. Photosynthetic rates at the mean growth CO₂ concentration were higher in HC plants than LC plants, but photosynthetic rates at a common CO₂ concentration were lower in HC plants. Efficiency of water and nitrogen use of leaves at growth CO₂ concentration was greatly increased in HC plants. These results suggest that natural plants growing in elevated CO₂ levels under cool temperate climate conditions have down-regulated their photosynthetic capacity but that they increased photosynthetic rates and resource use efficiencies due to the direct effect of elevated CO₂ concentration.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

Elevated atmospheric partial pressure of CO2 and plant growth

Cotton plants were grown in late spring under full sunlight in glasshouses containing normal ambient partial pressure of CO₂ (32±2Pa) and enriched partial pressure of CO₂ (64±1.5Pa) and at four levels of nitrogen nutrition. Thirty-five days after planting, the total dry weights of high CO₂-grown plants were 2- to 3.5-fold greater than plants grown in normal ambient CO₂ partial pressure. Depending on nitrogen nutrition level, non-structural carbohydrate content (mainly starch) in the leaves of plants grown in normal CO₂ was between 4 and 37% of the total leaf dry weight compared to 39 to 52% in the leaves of high CO₂-grown plants. Specific leaf weight calculated using total dry weight was 1.6- to 2-fold greater than that based on structural dry weight. In high CO₂-grown plants the amount of non-structural carbohydrate translocated from the leaves at night was between 10 and 20% of the level at the end of the photoperiod. This suggests that the plant was unable to utilize all the carbohydrate it assimilated in elevated CO₂ atmosphere. While there was a 1.5-fold enhancement in the rate of CO₂ assimilation in plants grown in 64 Pa CO₂, there was, however, some evidence to suggest that the activities of other metabolic pathways in the plants were not stimulated to the same extent by the enriched CO₂ atmosphere. This resulted in massive accumulation of non-structural carbohydrate, particularly at low level of nitrogen nutrition.Abbreviations A rate of CO₂ assimilation - PPFD photosynthetic photo flux density - NAR net assimilation rate - pCO₂ partial pressure of CO₂ - RGR relative growth rate     

Oscillatory Transpiration and CO2 Exchange of Citrus Leaves at the CO2 Compensation Concentration

CO₂ and H₂O vapor exchange were measured by enclosing citrus (Citrus sinensis cv. Sour Orange) leaves in a temperature controlled transparent leaf chamber. Introduction of dry air into the closed circuit gas flow caused cyclic oscillation in CO₂ and H₂O vapor exchange. It is suggested that oscillation in the CO₂ exchange at the CO₂ compensation concentration is due to oscillation in non-stomatal resistance to CO₂. Three types of oscillation were observed: 3–6 min (peak to peak) in young leaves, 30 min in mature leaves, and 160 min in old leaves.     

Prolonged Growth of Young Spruce (<Emphasis Type="Italic">Picea koraiensis</Emphasis> Nakai) Plants at Double Atmospheric CO<Subscript>2</Subscript> Concentration Stimulates the Preferential Growth of Thick Roots

Two-year-old young spruce (Picea koraiensis Nakai) plants were grown in a climatic chamber during three summer months at double atmospheric CO₂ concentration, sufficient content of soil inorganic nitrogen, and diurnal variation of temperature and illuminance, which simulate natural growth conditions. The control plants were grown in another climatic chamber under the same conditions, but at atmospheric CO₂ concentration (350 ppm). CO₂ exchange was measured with a Li-Cor 6400 infra-red gas analyzer in attached leaves placed in a climatic chamber in the morning under growth conditions and saturating light 1200 μE/(m² s) in June, July, and August. In addition, dry weights of needles, leafless shoot parts of plant, fraction of thick (more than 0.5 mm in diameter) and thin (less than 0.5 mm in diameter) roots were recorded. The data were used to plot CO₂ exchange rates as a function of carbon dioxide concentration and to calculate the increment of shoot and root phytomass. The maximum gas exchange rates in the treated and control plants similarly depended on CO₂ concentration. The slope of the CO₂ dependence curve, which corresponded to the kinetic characteristic V _m /K _M of photosynthetic carboxylation, increased monotonically during the experiment. To the end of observation period, the proportion of thick roots in plant phytomass significantly increased in the plants grown at double atmospheric CO₂ concentration, as compared to the control plants. Thus, the increase in the rate of photosynthetic gas exchange in plants grown for three months at double atmospheric CO₂ concentration was only due to the increase in CO₂, the substrate of Rubisco carboxylation activity. We found no differences in the CO₂ characteristic for Rubisco between the treated and control plants. The ratio of needle to thin roots in the treated and control plants was similar and did not change during the experiment. The excess of photoassimilates in the treated, as compared to the control plants, was preferentially used for thick root growth. This result shows that photosynthesis in young spruce forests can deposit excess atmospheric CO₂ in the soil horizon in the form of thick root phytomass. __________ Translated from Fiziologiya Rastenii, Vol. 52, No. 5, 2005, pp. 741–746. Original Russian Text Copyright ? 2005 by Mao, Y.-J. Wang, Zhu, X.-W. Wang, Sun, Zhou, Voronin.     

Growth and Photosynthetic Characteristics of Solanum tuberosum Plantlets Cultivated in Vitro in Different Conditions of Aeration, Sucrose Supply, and CO(2) Enrichment

Growth characteristics, oxygen exchange, and carbohydrate and chlorophyll contents were determined 30 days after subculturing of single node-derived plantlets of Solanum tuberosum cv Haig cultivated in vitro. Cultivation conditions were: (a) photomixotrophy in closed vessel, (b) photomixotrophy in closed vessel on medium supplemented with silver thiosulfate, (c) photomixotrophy in aerated vessel, (d) photoautotrophy in air, (e) photoautotrophy in CO₂-enriched air. In photomixotrophic conditions, aeration of the vessel enhanced sucrose utilization and had a positive effect on plantlet growth. In photoautotrophic conditions, growth of the plantlets was slow in air and was strongly enhanced by CO₂ enrichment of the atmosphere. Starch to sucrose ratios were higher in plants grown photoautotrophically than in plants grown with sucrose in the medium. Oxygen exchange characteristics on a chlorophyll basis were similar between the plantlets when measured under moderate light, and resembled those of greenhouse plant leaves. In high light, however, plantlets grown photoautotrophically in a CO₂-enriched atmosphere had higher oxygen exchange rates. We concluded from these results that potato plantlets in vitro in conditions (c), (d), and (e) developed C3-plant photosynthetic characteristics, which were in photoautotrophically grown plantlets comparable to those of field-grown plants.     

Leaf Photosynthesis and Respiration of High CO(2)-Grown Tobacco Plants Selected for Survival under CO(2) Compensation Point Conditions

Four self-pollinated, doubled-haploid tobacco, (Nicotiana tabacum L.) lines (SP422, SP432, SP435, and SP451), selected as haploids by survival in a low CO₂ atmosphere, and the parental cv Wisconsin-38 were grown from seed in a growth room kept at high CO₂ levels (600-700 parts per million). The selected plants were much larger (especially SP422, SP432, and SP451) than Wisconsin-38 nine weeks after planting. The specific leaf dry weight and the carbon (but not nitrogen and sulfur) content per unit area were also higher in the selected plants. However, the chlorophyll, carotenoid, and alkaloid contents and the chlorophyll a/b ratio varied little. The net CO₂ assimilation rate per unit area measured in the growth room at high CO₂ was not higher in the selected plants. The CO₂ assimilation rate versus intercellular CO₂ curve and the CO₂ compensation point showed no substantial differences among the different lines, even though these plants were selected for survival under CO₂ compensation point conditions. Adult leaf respiration rates were similar when expressed per unit area but were lower in the selected lines when expressed per unit dry weight. Leaf respiration rates were negatively correlated with specific leaf dry weight and with the carbon content per unit area and were positively correlated with nitrogen and sulfur content of the dry matter. The alternative pathway was not involved in respiration in the dark in these leaves. The better carbon economy of tobacco lines selected for low CO₂ survival was not apparently related to an improvement of photosynthesis rate but could be related, at least partially, to a significantly reduced respiration (mainly cytochrome pathway) rate per unit carbon.     

The influences of increased CO2 and water supply on growth,biomass allocation and water use efficiency of Sinapis alba L. grown under different wind speeds

We examined how independent and interactive effects of CO₂ concentrations, water supply and wind speed affect growth rates, biomass partitioning, water use efficiency, diffusive conductance and stomatal density of plants. To test the prediction that wind stress will be ameliorated by increased CO₂ and/or by unrestricted water supply we grew Sinapis alba L. plants in controlled chambers under combinations of two levels of CO₂ (350 ppmv, 700 ppmv), two water regimes and two wind speeds (0.3 ms^–1, 3.7 ms^–1). We harvested at ten different dates over a period of 60 days. A growth analysis was carried out to evaluate treatment effects on plant responses. Plants grown both in increased CO₂ and in low wind conditions had significantly greater stem length, leaf area and dry weights of plant parts. Water supply significantly affected stem diameter, root weight and leaf area. CO₂ enrichment significantly increased the rate of biomass accumulation and the relative ratio of biomass increase to leaf area expansion. High wind speed significantly reduced plant growth rates and the rate of leaf area expansion was reduced more than the rate of biomass accumulation. Regression analysis showed significant CO₂ effects on the proportion of leaf and stem dry weight to total dry weight. A marked plant-age effect was dependent on water supply, wind speed and CO₂ concentration. A reduced water supply significantly decreased the stomatal conductance, and water use efficiency significantly increased with a limited water supply, low wind and increased CO₂. We found significant CO₂ x wind effects for water diffusion resistance, adaxial number of stomata and water use efficiencies and significant wind x water effect for water use efficiency. In conclusion, wind stress was ameliorated by growing in unrestricted water but not by growing in increased CO₂.     

A Direct Confirmation of the Standard Method of Estimating Intercellular Partial Pressure of CO(2)

The partial pressure of CO₂ inside leaves of several species was measured directly. Small gas exchange chambers were clamped above and below the same section of an amphistomatous leaf. A flowing gas stream through one chamber allowed normal CO₂ and water vapor exchange. The other chamber was in a closed circuit consisting of the chamber, an infrared gas analyzer, and a peristaltic pump. The CO₂ in the closed system rapidly reached a steady pressure which it is believed was identical to the CO₂ pressure inside the leaf, because there was no flux of CO₂ across the epidermis. This measured partial pressure was in close agreement with that estimated from a consideration of the fluxes of CO₂ and vapor at the other surface.     

Regulation of hormonal responses of sweet pepper as affected by salinity and elevated CO2 concentration

This study examines the extent to which the predicted CO₂‐protective effects on the inhibition of growth, impairment of photosynthesis and nutrient imbalance caused by saline stress are mediated by an effective adaptation of the endogenous plant hormonal balance. Therefore, sweet pepper plants (Capsicum annuum, cv. Ciclón) were grown at ambient or elevated [CO₂] (400 or 800 µmol mol^–1) with a nutrient solution containing 0 or 80 mM NaCl. The results show that, under saline conditions, elevated [CO₂] increased plant dry weight, leaf area, leaf relative water content and net photosynthesis compared with ambient [CO₂], whilst the maximum potential quantum efficiency of photosystem II was not modified. In salt‐stressed plants, elevated [CO₂] increased leaf NO₃^– concentration and reduced Cl^– concentration. Salinity stress induced ABA accumulation in the leaves but it was reduced in the roots at high [CO₂], being correlated with the stomatal response. Under non‐stressed conditions, IAA was dramatically reduced in the roots when high [CO₂] was applied, which resulted in greater root DW and root respiration. Additionally, the observed high CK concentration in the roots (especially tZR) could prevent downregulation of photosynthesis at high [CO₂], as the N level in the leaves was increased compared with the ambient [CO₂], under salt‐stress conditions. These results demonstrate that the hormonal balance was altered by the [CO₂], which resulted in significant changes at the growth, gas exchange and nutritional levels.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Winter soil frost conditions in boreal forests control growing season soil CO2 concentration and its atmospheric exchange

The impact of changes in winter soil frost regime on soil CO₂ concentration and its atmospheric exchange in a boreal Norway spruce forest was investigated using a field‐scale soil frost manipulation experiment. The experiment comprised three treatments: deep soil frost, shallow soil frost and control plots (n= 3). Winter soil temperatures and soil frost distribution were significantly altered by the different treatments. The average soil CO₂ concentrations during the growing season were significantly lower in plots with deep soil frost than in plots with shallow soil frost. The average CO₂ soil–atmosphere exchange rate exhibited the same pattern, and differences in soil respiration rates among the treatments were statistically significant. Both the variation in soil CO₂ concentration and the CO₂ soil–atmosphere exchange rate could statistically be explained by the differences in the maximum soil frost depth during the previous winter. A response model for growing season soil respiration rates suggests that every 1 cm change in winter soil frost depth will change the emission rates by ca. 0.01 g CO₂ m^?2 day^?1, corresponding to 0.2–0.5% of the estimated net ecosystem productivity (NEP). This suggests that the soil frost regime has a significant influence on the C balance of the system, because interannual variations in soil frost up to 60 cm have been recorded at the site. We conclude that winter climate conditions can be important in controlling C balances in northern terrestrial ecosystems, and also that indirect effects of the winter season must be taken into account, because these can affect the prevailing conditions during the growing season.     

Land plants equilibrate O2 and CO2 concentrations in the atmosphere

The role of land plants in establishing our present day atmosphere is analysed. Before the evolution of land plants, photosynthesis by marine and fresh water organisms was not intensive enough to deplete CO₂ from the atmosphere, the concentration of which was more than the order of magnitude higher than present. With the appearance of land plants, the exudation of organic acids by roots, following respiratory and photorespiratory metabolism, led to phosphate weathering from rocks thus increasing aquatic productivity. Weathering also replaced silicates by carbonates, thus decreasing the atmospheric CO₂ concentration. As a result of both intensive photosynthesis and weathering, CO₂ was depleted from the atmosphere down to low values approaching the compensation point of land plants. During the same time period, the atmospheric O₂ concentration increased to maximum levels about 300 million years ago (Permo-Carboniferous boundary), establishing an O₂/CO₂ ratio above 1000. At this point, land plant productivity and weathering strongly decreased, exerting negative feedback on aquatic productivity. Increased CO₂ concentrations were triggered by asteroid impacts and volcanic activity and in the Mesozoic era could be related to the gymnosperm flora with lower metabolic and weathering rates. A high O₂/CO₂ ratio is metabolically linked to the formation of citrate and oxalate, the main factors causing weathering, and to the production of reactive oxygen species, which triggered mutations and stimulated the evolution of land plants. The development of angiosperms resulted in a decrease in CO₂ concentration during the Cenozoic era, which finally led to the glacial-interglacial oscillations in the Pleistocene epoch. Photorespiration, the rate of which is directly related to the O₂/CO₂ ratio, due to the dual function of Rubisco, may be an important mechanism in maintaining the limits of O₂ and CO₂ concentrations by restricting land plant productivity and weathering.     

Impact of rising CO2 on emissions of volatile organic compounds: isoprene emission from Phragmites australis growing at elevated CO2 in a natural carbon dioxide spring†

Isoprene basal emission (the emission of isoprene from leaves exposed to a light intensity of 1000 µmol m^?2 s^?1 and maintained at a temperature of 30 °C) was measured in Phragmites australis plants growing under elevated CO₂ in the Bossoleto CO₂ spring at Rapolano Terme, Italy, and under ambient CO₂ at a nearby control site. Gas exchange and biochemical measurements were concurrently taken. Isoprene emission was lower in the plants growing at elevated CO₂ than in those growing at ambient CO₂. Isoprene emission and isoprene synthase activity (IsoS) were very low in plants growing at the bottom of the spring under very rich CO₂ and increased at increasing distance from the spring (and decreasing CO₂ concentration). Distance from the spring did not significantly affect photosynthesis making it therefore unlikely that there is carbon limitation to isoprene formation. The isoprene emission rate was very quickly reduced after rapid switches from elevated to ambient CO₂ in the gas‐exchange cuvette, whereas it increased when switching from ambient to elevated CO₂. The rapidity of the response may be consistent with post‐translational modifications of enzymes in the biosynthetic pathway of isoprene formation. Reduction of IsoS activity is interpreted as a long‐term response. Basal emission of isoprene was not constant over the day but showed a diurnal course opposite to photosynthesis, with a peak during the hottest hours of the day, independent of stomatal conductance and probably dependent on external air temperature or temporary reduction of CO₂ concentration. The present experiments show that basal emission rate of isoprene is likely to be reduced under future elevated CO₂ levels and allow improvement in the modelling of future isoprene emission rates.     

Relationship between Net CO(2) Assimilation and Dry Weight Accumulation in Field-Grown Tobacco

To assess the variability of net photosynthetic CO₂ exchange per unit leaf area and to construct budgets for stands of field-grown tobacco (Nicotiana tabacum, Connecticut Broadleaf), a number of short-time measurements were made on all available leaf positions on two varieties using a hand-held transparent chamber for conducting gas exchange measurements on leaves. Measurements of net CO₂ exchange were carried out on 18 separate days during a 35-day period, beginning 22 days after the seedlings were transplanted to the field. Gas exchange assays on leaves were conducted under ambient conditions of temperature and light intensity at all times of day. Solar radiation was monitored throughout the period, and losses of respiratory CO₂ from stems, roots, and leaves (in the dark) were estimated. A simple model was proposed to relate daily total CO₂ input to irradiance and total leaf area. The total leaf area was assumed to be a function of day number. Dark respiratory losses accounted for 41% to 47% of total CO₂ assimilation. Analysis of variance indicated that the two varieties were not significantly different in whole plant rate of CO₂ fixation per unit of leaf area. CO₂ input was closely associated with leaf area within each variety. Throughout the experiment, the difference between the two varieties in total leaf area per plant was the largest single factor in determining net CO₂ inputs. The cumulative dry weight increase for each variety was similar to the prediction of net dry matter input obtained by gas exchange measurements, thus confirming the close relationship between total plant net CO₂ assimilation and dry weight yield.