Resistances to the Diffusion of Gas and Vapor in Leaves

Considerable error may be incurred when the component leaf resistances to gases are calculated from the rates of photosynthesis and transpiration measured for the entire leaf. This is due to the fact that there are two parallel pathways for diffusion — through the upper and the lower leaf surfaces — which in most leaves are asymmetrical. True values for the resistances, and predictions of the results of their modification, have been obtained from separate measurements of the photosynthesis and transpiration via the two leaf surfaces.     

CO2 and Water Vapor Exchange across Leaf Cuticle (Epidermis) at Various Water Potentials

Cuticular properties affect the gas exchange of leaves, but little is known about how much CO2 and water vapor cross the cuticular barrier or whether low water potentials affect the process. Therefore, we measured the cuticular conductances for CO2 and water vapor in grape (Vitis vinifera L.) leaves having various water potentials. The lower leaf surface was sealed to force all gas exchange through the upper surface, which was stoma-free. In this condition both gases passed through the cuticle, and the CO2 conductance could be directly determined from the internal mole fraction of CO2 near the compensation point, the external mole fraction of CO2, and the CO2 flux. The cuticle allowed small amounts of CO2 and water vapor to pass through, indicating that gas exchange occurs in grape leaves no matter how tightly the stomata are closed. However, the CO2 conductance was only 5.7% of that for water vapor. This discrimination against CO2 markedly affected calculations of the mole fraction of CO2 in leaves as stomatal apertures decreased. When the leaf dehydrated, the cuticular conductance to water vapor decreased, and transpiration and assimilation diminished. This dehydration effect was largest when turgor decreased, which suggests that cuticular gas exchange may have been influenced by epidermal stretching.     

Gas Exchange of Carrot Leaves in Response to Elevated CO2 Concentration

Short-term responses of four carrot (Daucus carota) cultivars: Cascade, Caro Choice (CC), Oranza, and Red Core Chantenay (RCC) to CO₂ concentrations (C _a) were studied in a controlled environment. Leaf net photosynthetic rate (P _N), intercellular CO₂ (C _i), stomatal conductance (g _s), and transpiration rate (E) were measured at C _a from 50 to 1 050 mol mol^–1. The cultivars responded similarly to C _a and did not differ in all the variables measured. The P _N increased with C _a until saturation at 650 mol mol^–1 (C _i= 350–400 mol mol^–1), thereafter P _N increased slightly. On average, increasing C _a from 350 to 650 and from 350 to 1 050 mol mol^–1 increased P _N by 43 and 52 %, respectively. The P _N vs. C _i curves were fitted to a non-rectangular hyperbola model. The cultivars did not differ in the parameters estimated from the model. Carboxylation efficiencies ranged from 68 to 91 mol m^–2 s^–1 and maximum P _N were 15.50, 13.52, 13.31, and 14.96 mol m^–2 s^–1 for Cascade, CC, Oranza, and RCC, respectively. Dark respiration rate varied from 2.80 mol m^–2 s^–1 for Oranza to 3.96 mol m^–2 s^–1 for Cascade and the CO₂ compensation concentration was between 42 and 46 mol mol^–1. The g _s and E increased to a peak at C _a= 350 mol mol^–1 and then decreased by 17 and 15 %, respectively when C _a was increased to 650 mol mol^–1. An increase from 350 to 1 050 mol mol^–1 reduced g _s and E by 53 and 47 %, respectively. Changes in g _s and P _N maintained the C _i:C _a ratio. The water use efficiency increased linearly with C _a due to increases in P _N in addition to the decline in E at high C _a. Hence CO₂ enrichment increases P _N and decreases g _s, and can improve carrot productivity and water conservation.     

Effect of Temperature, CO(2) Concentration, and Light Intensity on Oxygen Inhibition of Photosynthesis in Wheat Leaves

The effect of 21% O₂ and 3% O₂ on the CO₂ exchange of detached wheat leaves was measured in a closed system with an infrared carbon dioxide analyzer. Temperature was varied between 2° and 43°, CO₂ concentration between 0.000% and 0.050% and light intensity between 40 ft-c and 1000 ft-c. In most conditions, the apparent rate of photosynthesis was inhibited in 21% O₂ compared to 3% O₂. The degree of inhibition increased with increasing temperature and decreasing CO₂ concentration. Light intensity did not alter the effect of O₂ except at light intensities or CO₂ concentrations near the compensation point. At high CO₂ concentrations and low temperature, O₂ inhibition of apparent photosynthesis was absent. At 3% O₂, wheat resembled tropical grasses in possessing a high rate of photosynthesis, a temperature optimum for photosynthesis above 30°, and a CO₂ compensation point of less than 0.0005% CO₂. The effect of O₂ on apparent photosynthesis could be ascribed to a combination of stimulation of CO₂ production during photosynthesis, and inhibition of photosynthesis itself.     

Carbon Dioxide Exchanges in Leaves. I. Discrimination Between CO(2) and CO(2) in Photosynthesis

In order to measure CO₂ exchange reactions by leaves using isotopes of CO₂, it is necessary to know precisely the discrimination against ¹⁴CO₂ by leaves. Earlier determinations of discrimination are at variance, and may be inaccurate because of assumptions made about the rate of photorespiration. Maize leaves evolve little or no CO₂ in light, and so provide suitable material for this measurement. Discrimination against ¹⁴CO₂ in photosynthesis by maize leaves is almost precisely the same as in CO₂ absorption by NaOH solution, amounting to 2.1 and 2.0% respectively. The agreement between these values and their close approximation to the relative rates of diffusion of ¹²CO₂ and ¹⁴CO₂, calculated from Graham's law, shows that diffusion into the leaf is primarily responsible for discrimination against ¹⁴CO₂ in photosynthesis.     

Influence of Leaf Starch Concentration on CO(2) Assimilation in Soybean

Net photosynthetic rate, CO₂ compensation concentration, and starch and soluble sugar concentrations were measured in soybean (Glycine max [L.] Merrill) leaves in an attempt to evaluate the effect of carbohydrate concentration on rate of CO₂ assimilation.     

Effect of Elevated CO2 Concentration on Leaf Structure of Brassica Juncea under Water Stress

Study on the effect of elevated CO₂ concentration on leaf structure of Brassica juncea L. cv. Bio-141 (95) under moisture stress revealed, that CO₂ elevated to 600 mol mol^–1 increased the length of epidermal cel and length of palisade parenchyma cells, and induced larger chloroplasts and more oval shaped starch granules in comparison with plants grown at ambient CO₂ concentration. This increase in structural sink size helped in check feedback inhibition by excessive photoassimilate which was subsequently used to compensate the adverse moisture stress effect in B. juncea leaves.     

Relationships of Starch Concentration with Specific Leaf Weight and Mineral Concentrations in Potato Leaves Under Varied CO2 and Temperature

Foliar concentrations of starch and major nutrients N, P, K, Ca, and Mg along with specific leaf weight (SLW) were determined in the potato (Solaruan tuberosum L. ) cvs "Denali", "Norland” and "Russet Burbank" grown for 35 days under the CO2 concentrations of 500, 1 000, 1 500 and 2 000 μmol' mol-l at both 16 and 20 ℃ air temperature. The starch concentration, pooled from the three cuhivars, increased with increasing CO2 concentration at both 16 and 20 ℃, and was consistently higher at 16 ℃ than at 20 ℃. The SLW (g·m-2) was positively related to the foliar starch concentration on the basis of leaf area or dry weight. The concentrations of N, P, Ca, and Mg in leaves were negatively related to starch concentration under 14% starch on a dry weight basis. Above 14%, there was no significant relationship between nutrient and starch concentrations. The similar patterns were seen when the SLW and nutrient concentrations were expressed on a starch-free basis. In contrast, the leaf concentration of K was not closely related to the starch concentration. The results indicated that the changes in SLW and concentrations of N, P, Ca, and Mg in potato leaves only partially resulted from the changed starch concentration.     

An Apparatus to Produce Gas Mixtures with Controlled CO(2), O(2), and Water Vapor Concentrations

An apparatus to produce continuous gas mixtures for use in measurements of plant gas exchange is described. A wide range of CO₂ and water vapor concentrations can be provided and O₂ concentration can be varied from 0 to 21%. Changes in the concentrations of the components are accomplished conveniently, rapidly, and independently. With occasional adjustments, CO₂ and O₂ concentrations can be maintained to within ± 1 μl/l and ± 0.1%, respectively. Dew point of the gas mixture can be maintained to within ± 0.05 C.     

Dark Release of CO(2) from Higher Plant Leaves

A study was conducted with 48 species of the amount of ¹⁴CO₂ released during the first minute of dark following fixation of ¹⁴CO₂ in the light. Light fixation periods varied from 5 to 60 seconds. The species examined included both monocots and dicots and represented C₄, C₃, and Crassulacean acid metabolism (CAM) photosynthetic types.     

Acclimation of CO(2) Assimilation in Cotton Leaves to Water Stress and Salinity

Cotton (Gossypium hirsutum L. cv Acala SJ2) plants were exposed to three levels of osmotic or matric potentials. The first was obtained by salt and the latter by withholding irrigation water. Plants were acclimated to the two stress types by reducing the rate of stress development by a factor of 4 to 7. CO₂ assimilation was then determined on acclimated and nonacclimated plants. The decrease of CO₂ assimilation in salinity-exposed plants was significantly less in acclimated as compared with nonacclimated plants. Such a difference was not found under water stress at ambient CO₂ partial pressure. The slopes of net CO₂ assimilation versus intercellular CO₂ partial pressure, for the initial linear portion of this relationship, were increased in plants acclimated to salinity of −0.3 and −0.6 megapascal but not in nonacclimated plants. In plants acclimated to water stress, this change in slopes was not significant. Leaf osmotic potential was reduced much more in acclimated than in nonacclimated plants, resulting in turgor maintenance even at −0.9 megapascal. In nonacclimated plants, turgor pressure reached zero at approximately −0.5 megapascal. The accumulation of Cl⁻ and Na⁺ in the salinity-acclimated plants fully accounted for the decrease in leaf osmotic potential. The rise in concentration of organic solutes comprised only 5% of the total increase in solutes in salinity-acclimated and 10 to 20% in water-stress-acclimated plants. This acclimation was interpreted in light of the higher protein content per unit leaf area and the enhanced ribulose bisphosphate carboxylase activity. At saturating CO₂ partial pressure, the declined inhibition in CO₂ assimilation of stress-acclimated plants was found for both salinity and water stress.     

Effect of Doubled CO2 Concentration on Leaf Chlorophyll-protein Complexes in Several Plants

The effect of doubled CO2 on the chlorophyll-protein complexes of the leaves of soybean ( Glycine max L., Ca plants), cucumber ( Cucumis sativus L., C3 plant), millet ( Setaria italica (L.) Beauv., not a very typical C4 plant) and corn (Zea mays L. ,C4 plant) was studied. Experi- mental plants were pot-cultured in polyethylene membrane (or glass) open top cultured chambers. After sowing, C02 was kept immediately either at ambient ( (350 ± 10) x 10-6) concentration for the control or at doubled CO2 ((700 ± 10) x 10-6) concentration for the treatment chambers. The chlorophyll-protein complexes of the thylakoid membrane of the plants were resolved by disk SDS- PAGE. The results showed that after doubled CO2 treatment,either in the soybean and cucmnber,or in the millet, the quantity of polymer state of PS Ⅱ light-harvesting chlorophyll a/b-protein complex (LHC Ⅱ ) had increased as the monomer state of LHC Ⅱ decreased. But such response to doubled CO2 was not found in corn, the C4 plant. The change of the state of LHC Ⅱ in soybean etc. might be an adaptive effect of plant photosynthetic mechanism to the long term elevated CO2. Thus it could increase the efficiency of the absorption, transfer and conversion of light energy in plant photosynthesis, and support the high efficiency of photosynthetic carbon assimilation.     

Internal CO(2) Measured Directly in Leaves : Abscisic Acid and Low Leaf Water Potential Cause Opposing Effects

Observations of nonuniform photosynthesis across leaves cast doubt on internal CO₂ partial pressures (p_i) calculated on the assumption of uniformity and can lead to incorrect conclusions about the stomatal control of photosynthesis. The problem can be avoided by measuring p_i directly because the assumptions of uniformity are not necessary. We therefore developed a method that allowed p_i to be measured continuously in situ for days at a time under growth conditions and used it to investigate intact leaves of sunflower (Helianthus annuus L.), soybean (Glycine max L. Merr.), and bush bean (Phaseolus vulgaris L.) subjected to high or low leaf water potentials (ψ_w) or high concentrations of abscisic acid (ABA). The leaves maintained a relatively constant differential (Δp) between ambient CO₂ and measured p_i throughout the light period when water was supplied. When water was withheld, ψ_w decreased and the stomata began to close, but measured p_i increased until the leaf reached a ψ_w of −1.76 (bush bean), −2.12 (sunflower) or −3.10 (soybean) megapascals, at which point Δp = 0. The increasing p_i indicated that stomata did not inhibit CO₂ uptake and a Δp of zero indicated that CO₂ uptake became zero despite the high availability of CO₂ inside the leaf. In contrast, when sunflower leaves at high ψ_w were treated with ABA, p_i did not increase and instead decreased rapidly and steadily for up to 8 hours even as ψ_w increased, as expected if ABA treatment primarily affected stomatal conductance. The accumulating CO₂ at low ψ_w and contrasting response to ABA indicates that photosynthetic biochemistry limited photosynthesis at low ψ_w but not at high ABA.     

Measurement of CO(2) and H(2)O Vapor Exchange in Spinach Leaf Discs : Effects of Orthophosphate

A leaf chamber has been designed which allows the measurement of both CO₂ and water vapor exchange in Spinacia oleracea leaf discs. The center of the disc lies within a cylindrical gas chamber and its margins are enclosed within a cavity through which water or various metabolites can be pumped. In saturating light and normal atmospheres, the leaf discs have a relatively low resistance to H₂O vapor transfer (r_w = 1.87 seconds per centimeter) and can support high rates of photosynthesis for several hours. The abaxial surface of a disc had a higher resistance to water vapor transfer (r_w = 3.22 seconds per centimeter) than the adaxial (r_w = 2.45 seconds per centimeter) despite having a higher stomatal frequency (abaxial, 105/square millimeter; adaxial, 58/square millimeter). In 2% O₂, the discs required an internal concentration of CO₂ of 115 microliters per liter to support one-half of the maximal velocity of apparent photosynthesis (average value, 66 milligrams CO₂ per square decimeter per hour). In 20% O₂, the comparable values are 156 microliters per liter and 56 milligrams CO₂ per square decimeter per hour. In air, apparent photosynthesis saturated at intensities (750 microeinsteins per square meter per second) well below that of daylight but, when the internal CO₂ was raised to 700 to 900 microliters per liter, photosynthesis was not saturated even at daylight intensities (2025 microeinsteins per square meter per second). The distribution of Prussian blue crystals, formed after ferrocyanide feeding, showed that water entered the disc via the vasculature. When 25-minute pulses of orthophosphate were provided in the feeding solution, there were concentration-dependent increases in both r_w and r_m leading to inhibition of photosynthesis. The orthophosphate-dependent inhibitions were reversible.     

Effects of CO(2) Concentration on Rubisco Activity, Amount, and Photosynthesis in Soybean Leaves

Growth at an elevated CO₂ concentration resulted in an enhanced capacity for soybean (Glycine max L. Merr. cv Bragg) leaflet photosynthesis. Plants were grown from seed in outdoor controlled-environment chambers under natural solar irradiance. Photosynthetic rates, measured during the seed filling stage, were up to 150% greater with leaflets grown at 660 compared to 330 microliters of CO₂ per liter when measured across a range of intercellular CO₂ concentrations and irradiance. Soybean plants grown at elevated CO₂ concentrations had heavier pod weights per plant, 44% heavier with 660 compared to 330 microliters of CO₂ per liter grown plants, and also greater specific leaf weights. Ribulose 1,5-bisphosphate carboxylase/oxygenase (rubisco) activity showed no response (mean activity of 96 micromoles of CO₂ per square meter per second expressed on a leaflet area basis) to short-term (~1 hour) exposures to a range of CO₂ concentrations (110-880 microliters per liter), nor was a response of activity (mean activity of 1.01 micromoles of CO₂ per minute per milligram of protein) to growth CO₂ concentration (160-990 microliters per liter) observed. The amount of rubisco protein was constant, as growth CO₂ concentration was varied, and averaged 55% of the total leaflet soluble protein. Although CO₂ is required for activation of rubisco, results indicated that within the range of CO₂ concentrations used (110-990 microliters per liter), rubisco activity in soybean leaflets, in the light, was not regulated by CO₂.     

The Effect of Leaf Water Potential, Leaf Temperature and Light Intensity on Leaf Diffusion Resistance and the Transpiration of Leaves of Malus sylvestris

The relationship between leaf resistance to water vapour diffusion and each of the factors leaf water potential, light intensity and leaf temperature was determined for leaves on seedling apple trees (Malus sylvestris Mill. cv. Granny Smith) in the laboratory. Leaf cuticular resistance was also determined and transpiration was measured on attached leaves for a range of conditions. Leaf resistance was shown to be independent of water potential until potential fell below — 19 bars after which leaf resistance increased rapidly. Exposure of leaves to CO₂-free air extended the range for which resistance was independent of water potential to — 30 bars. The light requirement for minimum leaf resistance was 10 to 20 W m^?2 and at light intensities exceeding these, leaf resistance was unaffected by light intensity. Optimum leaf temperature for minimum diffusion resistance was 23 ± 2°C. The rate of change measured in leaf resistance in leaves given a sudden change in leaf temperature increased as the magnitude of the temperature change increased. For a sudden change of 1°C in leaf temperature, diffusion resistance changed at a rate of 0.01 s cm^?1 min^?1 whilst for a 9°C leaf temperature change, diffusion resistance changed at a rate of 0.1 s cm^?1 min^?1. Cuticular resistance of these leaves was 125 s cm^?1 which is very high compared with resistances for open stomata of 1.5 to 4 s cm^?1 and 30 to 35 s cm^?1 for stomata closed in the dark. Transpiration was measured in attached apple leaves enclosed in a leaf chamber and exposed to a range of conditions of leaf temperature and ambient water vapour density. Peak transpiration of approximately 5 × 10^?6 g cm^?2 s^?1 occurred at a vapour density gradient from the leaf to the air of 12 to 14 g m^?3 after which transpiration declined due presumably to increased stomatal resistance. Leaves in CO₂-free air attained a peak transpiration of 11 × 10^?6 g cm^?2 s^?1 due to lower values of leaf resistance in CO₂ free air. Transpiration then declined in these leaves due to development of an internal leaf resistance (of up to 2 s cm^?1). The internal resistance was masked in leaves at normal CO₂ concentrations by the increase in stomatal resistance.     

Effects of Temperature on Photosynthesis and CO(2) Evolution in Light and Darkness by Green Leaves

Using an open and a closed system of gas analysis, it was found that CO₂ evolution in light and in darkness from plant leaves (sunflower, soybean, watermelon, eggplant, and jackbean) have a different response to temperature. While the rate of CO₂ evolution in light increased with increasing temperature from 17 to 35° and then declined, the rate of CO₂ evolution in darkness increased continuously up to 40°. The rate of CO₂ evolution in light was affected by light intensity. At 1800 ft-c and below 35° the rate of CO₂ evolution in light was greater than in darkness, but above 35° it became lower than in darkness. The Q₁₀ for CO₂ evolution in light was consistently lower than that in darkness.     

Effect of CO(2) Concentration on Glycine and Serine Formation during Photorespiration

Amount and products of photosynthesis during 10 minutes were measured at different ¹⁴CO₂ concentrations in air. With tobacco (Nicotiana tabacum L. cv. Maryland Mammoth) leaves the percentage of ¹⁴C in glycine plus serine was highest (42%) at 0.005% CO₂, and decreased with increasing CO₂ concentration to 7% of the total at 1% CO₂ in air. However, above 0.03% CO₂ the total amount of ¹⁴C incorporated into the glycine and serine pool was about constant. At 0.005% or 0.03% CO₂ the percentage and amount of ¹⁴C in sucrose was small but increased greatly at higher CO₂ levels as sucrose accumulated as an end product. Relatively similar data were obtained with sugar beet (Beta vulgaris L. cv. US H20) leaves. The results suggest that photorespiration at high CO₂ concentration is not inhibited but that CO₂ loss from it becomes less significant.     

Effect of CO2 Concentration Doubling on the Leaf Morphology and Structure of 10 Species in Gramineae

The effects of CO2 concentration on leaf thickness, chloroplast manbers in the bundle sheath cell, epidermal cell density, stomatal density, stomatal index, stomatal size were compared in 10 species in Gramineae: Triticum aestivum L., T. aestivum ssp. tibeticum, Hordeum vulgare L., H. brevisubulatum ( Trin. ) Link, Oryza sativa L., O. meyeriana ssp. granulata, Setaria italica (L.) Beauv, S. viridis (L.) Beauv, Sorghum vulgare Pers., Zea mays L. following their exposure to doubled carbon dioxide (700μL/L) and ambient carbon dioxide concentration (350μL/L). The results indicated that different species of plants might vary in their response to doubled CO2. In general, the leaves became thicker under the elevated CO2 condition. The mean stomatal density of the C3 species was decreased in doubled CO2, whereas the results of C4 species showed an inverse trend. The epidermal cell density and the chloroplast numbers of the bundle sheath cell in the wild plant species were less than those in the control under CO2 enrichment. The stomatal density was positively correlated with the stomatal index. Finally, the general pattern of structural variation under different CO2 concentrations was proposed, and their implication to the research of global change was discussed as well.