Photosynthesis, Transpiration, Leaf Stomatal and Mesophyll Resistance Measurements by the Use of a Ventilated Diffusion Porometer

A ventilated diffusion porometer was modified and adapted for simultaneous measurements of leaf resistance and photosynthesis (using ¹⁴C). The system enables measurements to be made under field and laboratory conditions with different concentrations of CO₂ and vapor pressure gradients between the evaporating surfaces inside the leaf and the external atmosphere. The leaf is subjected to the porometer's atmosphere only for short periods (up to 30 seconds) and it is assumed that stomata are not affected. Establishing the linear regression of the effect of CO₂ concentration on net photosynthesis makes it possible to extrapolate for CO₂ compensation point, to calculate the overall resistance to CO₂ and the mesophyll resistance to CO₂.     

Photosynthetic Gas Exchange Characteristics of Wheat Flag Leaf Blades and Sheaths during Grain Filling: The Case of a Spring Crop Grown under Mediterranean Climate Conditions

The rate of net CO₂ assimilation (A), the stomatal (g_s) and residual (g_r) conductances to CO₂, the intercellular CO₂ concentration, the CO₂ compensation points at 21% O₂ (Γ₂₁) and at 2% O₂ (Γ₂), and the amounts of dry matter, nitrogen, and carbohydrates were determined, from anthesis through grain filling, in the flag leaf blade and sheath of spring wheat (Triticum aestivum L. cv Kolibri). The nitrogen content and the rate of net CO₂ assimilation declined slowly until the onset of senescence in both organs, about 3 weeks after anthesis. During senescence the reduction of A in both organs was not primarily caused by a decrease in g_s; the main factor is the decrease in g_r. From values of Γ₂₁ and Γ₂ it is suggested that the rate of respiration in the light contributing to the CO₂ compensation point is higher in sheaths than in blades irrespective of the O₂ level considered. The role of sheaths storing and later transporting assimilates to the developing grains seems to be more important for shoot yield than that of sheaths functioning as photosynthetic organs after the onset of senescence occurs. It is suggested that accumulation of carbohydrates in leaves might somehow trigger senescence in the flag leaf blade and sheath simultaneously.     

Changes in photosynthetic characteristics during leaf development in apple

A comprehensive developmental survey of leaf area, chlorophyll, photosynthetic rate, leaf resistance, transpiration ratio, CO₂ compensation point and photorespiration was conducted in apple. The largest changes in each of the photosynthetic characteristics studied took place during the earliest stages of leaf development, coinciding with the period of greatest leaf expansion and chlorophyll synthesis. During early development, photosynthesis increased 5-fold, reaching a maximum rate of 40 mg CO₂ dm^-2 hr^-1 at a leaf plastochron index (LPI) of 10. During this same period, leaf resistance, transpiration ratio, CO₂ compensation point and mesophyll resistance decreased, while carboxylation efficiency increased. Two especially interesting aspects of the data discussed are simultaneous changes that occur at a LPI of 10 and 12 in all of the photosynthetic characteristics examined and an apparent decrease in photorespiration as leaves age. From our results it is clear that stage of leaf development is an important factor affecting the rate of photosynthesis and photorespiration.Scientific Paper No. 5687, College of Agriculture, Washington State University, Pullman. This work is supported by the National Science Foundation Grant 80-10958 and the Columbia River Orchards Foundation.     

Genotypic variation in carboxylation of tomatoes

The gas exchange characteristics of 24 genotypes of Lycopersicon esculentum Mill. and one of L. minutum were measured with an infrared gas analyzer and dew point hygrometer in an open system. Net carbon exchange (NCE) and transpiration rate were measured at 50, 100, 150, and 300 μ1 1⁻¹ CO₂, and a regression of NCE versus internal lead [CO₂] estimates was calculated. The slope of the regression curve at the CO₂ compensation point was used as the measure of carboxylation efficiency (CE). Significant genotypic differences for CE were obtained. Differences in CE did not appear to be due to differences in diffusive resistance defined as the sum of the boundary layer resistance (r_a) and the stomatal plus cuticular resistance (r₁). There was no correlation (r = -0.07) between (r_a + r₁) and CE. Within groups with nonsignificantly different means for (r_a + r₁) there were genotypes with extremes for CE.     

Variables Affecting the CO(2) Compensation Point

Some factors influencing dark respiration, photorespiration, and photosynthesis were examined for their effect on the CO₂ compensation point (70 μl/l) of detached soybean (Glycine max) leaf discs. A higher compensation point in young leaves decreased to the constant value after leaf expansion and maturation, but increased again during senescence. The compensation point was 40 to 50% higher in plants grown in the summer than in the winter. The compensation point and dark respiration increased with temperatures above 17 C. Below 17 C dark respiration continued to decrease, but the compensation point did not decrease further. Increasing light intensities did not affect the compensation point.     

Über den Einfluß niedriger und hoher O2-Partialdrucke auf den Sauerstoff- und Kohlendioxidumsatz von Amaranthus und Phaseolus während der Lichtphase

Summary Carbon dioxide and oxygen gas exchange of illuminated Amaranthus and Phaseolus leaves was measured from 0–600 ppm of CO₂ in an open system.At low oxygen concentration (2% O₂) the ratio of CO₂ uptake to O₂ evolution came close to 1.At high oxygen partial pressure (42% O₂) the O₂ compensation point of an Amaranthus leaf was increased and oxygen evolution was depressed. Accordingly the CO₂/O₂ quotients were variable; the lowest value of 1,9 differed significantly from 1,0.The oxygen and carbon dioxide compensation points of a Phaseolus leaf were increased at high oxygen concentration (42% O₂) and oxygen evolution as well as carbon dioxide uptake were reduced. Therefore the ratios CO₂ over O₂ varied and differed greatly from 1,0.It was concluded that the nature of photosynthates is regulated by the gas composition around the leaves.     

Light-Dependent Oxygen Uptake, Glycolate, and Ammonia Release in l-Methionine Sulfoximine-Treated Chlamydomonas

Glycolate and ammonia excretion plus oxygen exchanges were measured in the light in l-methionine-dl-sulfoximine treated air-grown Chlamydomonas reinhardii. At saturating CO₂ (between 600 and 700 microliters per liter CO₂) neither glycolate nor ammonia were excreted, whereas at the CO₂ compensation concentration (<10 microliters per liter CO₂) treated algae excreted both glycolate and ammonia at rates of 37 and 59 nanomoles per minute per milligram chlorophyll, respectively. From the excretion values we calculate the amount of O₂ consumed through the glycolate pathway. The calculated value was not significantly different from the component of O₂ uptake sensitive to CO₂ obtained from the difference between O₂ uptake of the CO₂ compensation point and at saturating CO₂. This component was about 40% of stationary O₂ uptake measured at the CO₂ compensation point. From these data we conclude that glyoxylate decarboxylation in air-grown Chlamydomonas represents a minor pathway of metabolism even in conditions where amino donors are deficient and that processes other than glycolate pathway are responsible for the O₂ uptake insensitive to CO₂.     

Photosynthesis and Growth of Water Hyacinth under CO(2) Enrichment

Water hyacinth (Eichhornia crassipes [Mart.] Solms) plants were grown in environmental chambers at ambient and enriched CO₂ levels (330 and 600 microliters CO₂ per liter). Daughter plants (ramets) produced in the enriched CO₂ gained 39% greater dry weight than those at ambient CO₂, but the original mother plants did not. The CO₂ enrichment increased the number of leaves per ramet and leaf area index, but did not significantly increase leaf size or the number of ramets formed. Flower production was increased 147%. The elevated CO₂ increased the net photosynthetic rate of the mother plants by 40%, but this was not maintained as the plants acclimated to the higher CO₂ level. After 14 days at the elevated CO₂, leaf resistance increased and transpiration decreased, especially from the adaxial leaf surface. After 4 weeks in elevated as compared to ambient CO₂, ribulose bisphosphate carboxylase activity was 40% less, soluble protein content 49% less, and chlorophyll content 26% less; whereas starch content was 40% greater. Although at a given CO₂ level the enriched CO₂ plants had only half the net photosynthetic rate of their counterparts grown at ambient CO₂, they showed similar internal CO₂ concentrations. This suggested that the decreased supply of CO₂ to the mesophyll, as a result of the increased stomatal resistance, was counterbalanced by a decreased utilization of CO₂. Photorespiration and dark respiration were lower, such that the CO₂ compensation point was not altered. The photosynthetic light and CO₂ saturation points were not greatly changed, nor was the O₂ inhibition of photosynthesis (measured at 330 microliters CO₂ per liter). It appears that with CO₂ enrichment the temporary increase in net photosynthesis produced larger ramets. After acclimation, the greater total ramet leaf area more than compensated for the lower net photosynthetic rate on a unit leaf area basis, and resulted in a sustained improvement in dry weight gain.     

香樟凋落叶分解过程对两种间作作物生长和光合特性的影响

采用盆栽试验方法,探讨了香樟(Cinnamomum camphora)凋落叶不同土壤添加水平(0 g/盆为CK、25 g/盆、50 g/盆,100 g/盆)对受体作物小白菜(Brassica chinensis)、莴笋(Lactuca sativa)幼苗生长和光合特性的影响。结果显示:(1)香樟凋落叶分解对两种作物的地径、株高、生物量、叶片数和叶面积均有明显的抑制作用,且抑制效应随凋落叶添加量的增加而增强,但随着分解时间的延长其抑制作用逐渐减弱甚至表现为促进作用。(2)香樟凋落叶分解对两种作物的光合色素含量均有明显的抑制效应,并随凋落叶添加量的增加抑制作用增强,且持续时间延长。(3)经凋落叶处理的两种作物叶片净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、水分利用率(WUE)总体上均低于CK,而胞间CO₂浓度(C_i)在各凋落叶处理下均高于CK。(4)随着土壤中凋落叶量的增加,两种作物在光饱和以及CO₂饱和状态下的最大净光合速率(P_{n max})、表观量子效率(AQY)、羧化速率(CE)、光呼吸速率(R_p)和暗呼吸速率(R_d)均不断下降,而光补偿点(LCP)、光饱和点(LSP)、CO₂饱和点(CSP)、CO₂补偿点(CCP)因受体作物的不同,表现出不同的变化趋势。研究表明,土壤中香樟凋落叶分解释放的化感物质,能通过降低受体作物的光合色素合成和光合能力,限制其营养生长,最终影响生物量积累;相对于莴笋,小白菜对香樟凋落叶分解产生的化感胁迫效应具有更强的耐受性,可能更适宜在香樟林间种植。     

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.     

CO(2) and O(2) Exchange in Two Mosses, Hypnum cupressiforme and Dicranum scoparium

Photosynthetic CO₂ and O₂ exchange was studied in two moss species, Hypnum cupressiforme Hedw. and Dicranum scoparium Hedw. Most experiments were made during steady state of photosynthesis, using ¹⁸O₂ to trace O₂ uptake. In standard experimental conditions (photoperiod 12 h, 135 micromoles photons per square meter per second, 18°C, 330 microliters per liter CO₂, 21% O₂) the net photosynthetic rate was around 40 micromoles CO₂ per gram dry weight per hour in H. cupressiforme and 50 micromoles CO₂ per gram dry weight per hour in D. scoparium. The CO₂ compensation point lay between 45 and 55 microliters per liter CO₂ and the enhancement of net photosynthesis by 3% O₂versus 21% O₂ was 40 to 45%. The ratio of O₂ uptake to net photosynthesis was 0.8 to 0.9 irrespective of the light intensity. The response of net photosynthesis to CO₂ showed a high apparent K_m (CO₂) even in nonsaturating light. On the other hand, O₂ uptake in standard conditions was not far from saturation. It could be enhanced by only 25% by increasing the O₂ concentration (saturating level as low as 30% O₂), and by 65% by decreasing the CO₂ concentration to the compensation point. Although O₂ is a competitive inhibitor of CO₂ uptake it could not replace CO₂ completely as an electron acceptor, and electron flow, expressed as gross O₂ production, was inhibited by both high O₂ and low CO₂ levels. At high CO₂, O₂ uptake was 70% lower than the maximum at the CO₂ compensation point. The remaining activity (30%) can be attributed to dark respiration and the Mehler reaction.     

Effect of Oxygen on Photosynthesis, Photorespiration and Respiration in Detached Leaves. II. Corn and other Monocotyledons

The effect of O₂ on the CO₂ exchange of detached leaves of corn (Zea mays), wheat (Triticum vulgare), oats (Avena sativa), barley (Hordeum vulgare), timothy (Phleum pratense) and cat-tail (Typha angustifolia) was measured with a Clark oxygen electrode and infrared carbon dioxide analysers in both open and closed systems.

Corn leaves did not produce CO₂ in the light at any O₂ concentration, as was shown by the zero CO₂ compensation point and the absence of a CO₂ burst in the first minute of darkness. The rate of photosynthesis was inhibited by O₂ and the inhibition was not completely reversible. On the other hand, the steady rate of respiration after a few minutes in the dark was not affected by O₂.

These results were interpreted as indicating the absence of any measurable respiration during photosynthesis. Twelve different varieties of corn studied all responded to O₂ in the same way.

The other 5 monocotyledons studied did produce CO₂ in the light. Moreover, the CO₂ compensation point increased linearly with O₂ indicating a stimulation of photorespiration.

The implications of the lack of photorespiration in studies of primary productivity are discussed.

     

Effects of Chemical Treatments upon Photosynthetic Parameters in Soybean Seedlings

The effects of various chemical treatments upon photosynthesis, soluble leaf protein, CO₂ compensation point, and leaf light transmission in soybean, Glycine max (L.) Merr., seedlings were examined following varying response periods after application at 14 to 17 days postemergence. The compounds N⁶-benzyladenine (BA), 2-(4-chlorophenoxy)-2-methylpropanoic acid (CPMP), (4-chlorophenoxy)acetic acid (CPA), rhodanine-N-acetic acid (RAA), and 2,3,5-triiodobenzoic acid (TIBA) significantly increased soluble protein and decreased senescence, measured by leaf light transmission, at CO₂ concentrations below the compensation point in a survival chamber. All compounds except BA significantly decreased transmission values under ambient atmospheric conditions. In statistically significant experiments, applications of 3.49 millimolar CPMP increased net photosynthesis on a leaf area basis by an average of 14.4% at all trifoliolate positions with increases generally requiring response periods of 12 days or longer. RAA at 1.31 and 2.61 millimolar increased net photosynthesis by 19 to 36% following 13-day response periods. CPMP and other compounds tested had no effect upon the CO₂ compensation point after 4- to 8-day response periods. The effects of CPMP and RAA upon net photosynthesis and soluble protein appeared to involve a combined stimulation of protein synthesis and an antisenescent effect. There were no indications that any of the photosynthetic changes observed resulted from direct differential effects upon ribulose bisphosphate carboxylase-oxygenase. The assays for soluble protein and light transmission responded more consistently to the chemicals than did photosynthesis.     

Effect of oxygen and temperature on the efficiency of photosynthetic carbon assimilation in two microscopic algae

The CO₂ compensation points of Coccochloris peniocystis, a blue-green alga and Chlamydomonas reinhardtii, a green alga, were determined at pH 8.0 in a closed system by a gas chromatographic technique. The compensation point of Chlamydomonas increased markedly with temperature, rising from 0.79 microliter per liter CO₂ at 15 C to 2.5 microliters per liter CO₂ at 35 C. In contrast, the compensation point of Coccochloris at 20 C was 0.71 microliter per liter CO₂ and rose to only 0.95 microliter per liter CO₂ at 40 C.     

Manipulation of Catalase Levels Produces Altered Photosynthesis in Transgenic Tobacco Plants

Constructs containing the cDNAs encoding the primary leaf catalase in Nicotiana or subunit 1 of cottonseed (Gossypium hirsutum) catalase were introduced in the sense and antisense orientation into the Nicotiana tabacum genome. The N. tabacum leaf cDNA specifically overexpressed CAT-1, the high catalytic form, activity. Antisense constructs reduced leaf catalase specific activities from 0.20 to 0.75 times those of wild type (WT), and overexpression constructs increased catalase specific activities from 1.25 to more than 2.0 times those of WT. The NADH-hydroxypyruvate reductase specific activity in transgenic plants was similar to that in WT. The effect of antisense constructs on photorespiration was studied in transgenic plants by measuring the CO₂ compensation point (Γ) at a leaf temperature of 38°C. A significant linear increase was observed in Γ with decreasing catalase (at 50% lower catalase activity Γ increased 39%). There was a significant temperature-dependent linear decrease in Γ in transgenic leaves with elevated catalase compared with WT leaves (at 50% higher catalase Γ decreased 17%). At 29°C, Γ also decreased with increasing catalase in transgenic leaves compared with WT leaves, but the trend was not statistically significant. Rates of dark respiration were the same in WT and transgenic leaves. Thus, photorespiratory losses of CO₂ were significantly reduced with increasing catalase activities at 38°C, indicating that the stoichiometry of photorespiratory CO₂ formation per glycolate oxidized normally increases at higher temperatures because of enhanced peroxidation.     

Limiting Factors in Photosynthesis: IV. Iron Stress-Mediated Changes in Light-Harvesting and Electron Transport Capacity and its Effects on Photosynthesis in Vivo

Using iron stress to reduce the total amount of light-harvesting and electron transport components per unit leaf area, the influence of light-harvesting and electron transport capacity on photosynthesis in sugar beet (Beta vulgaris L. cv F58-554H1) leaves was explored by monitoring net CO₂ exchange rate (P) in relation to changes in the content of Chl.

In most light/CO₂ environments, and especially those with high light (≥1000 microeinsteins photosynthetically active radiation per square meter per second) and high CO₂ (≥300 microliters CO₂ per liter air), P per area was positively correlated with changes in Chl (a + b) content (used here as an index of the total amount of light-harvesting and electron transport components). This positive correlation of P per area with Chl per area was obtained not only with Fe-deficient plants, but also over the normal range of variation in Chl contents found in healthy, Fe-sufficient plants. For example, light-saturated P per area at an ambient CO₂ concentration close to normal atmospheric levels (300 microliters CO₂ per liter air) increased by 36% with increase in Chl over the normal range, i.e. from 40 to 65 micrograms Chl per square centimeter. Iron deficiency-mediated changes in Chl content did not affect dark respiration rate or the CO₂ compensation point. The results suggest that P per area of sugar beet may be colimited by light-harvesting and electron transport capacity (per leaf area) even when CO₂ is limiting photosynthesis as occurs under field conditions.

     

Effects of water stress and differential hardening treatments on photosynthetic characteristics of a xeromorphic shrub,Eucalyptus socialis,F. Muell.

Summary The influence of water stress on photosynthesis of drought hardened, and non-hardened,Eucalyptus socialis plants was examined. Particular attention was given to the effects of low leaf water potential on stomatal and intracellular resistance to CO₂ transport and on the CO₂ compensation point. Though the hardening treatment had a pronounced influence on leaf morphology, there was no apparent difference in the photosynthetic response to drought stress between hardened and non-hardened treatments, or with repeated drought cycles. These results suggest a high degree of genetic preconditioning to drought in this species.     

Influence of certain environmental factors on photosynthesis and photorespiration in Simmondsia chinensis

The response of net photosynthesis and apparent light respiration to changes in [O₂], light intensity, and drought stress was determined by analysis of net photosynthetic CO₂ response curves. Low [O₂] treatment resulted in a large reduction in the rate of photorespiratory CO₂ evolution. Lightintensity levels influenced the maximum net photosynthetic rate at saturating [CO₂]. These results indicate that [CO₂], [O₂] and light intensity affect the levels of substrates involved in the enzymatic reactions of photosynthesis and photorespiration. Intracellular resistance to CO₂ uptake decreased in low [O₂] and increased at low leaf water potentials. This response reflects changes in the efficiency with which photosynthetic and photorespiratory substrates are formed and utilized. Water stress had no effect on the CO₂ compensation point or the [CO₂] at which net photosynthesis began to saturate at high light intensity. The relationship between these data and recently published in-vitro kinetic measurements with ribulose-diphosphate carboxylase is discussed.Abbreviations C _w intracellular CO₂ concentration - F _gross gross photosynthesis - F _net net photosynthesis - I light intensity - R _L light respiration rate - r _c carboxylation resistance - r ₈ leaf gas-phase resistance - r _i intracellular resistance; to CO₂ uptake - r _t resistance to CO₂ flux between the intercellular spaces and the carboxylation sites - T _L leaf temperature - _t leaf water potential - CO₂ compensation point     

Relation of CO(2) Compensation Concentration to Apparent Photosynthesis in Maize

Significant differences in CO₂ compensation concentration measured in the field among varieties of the species Zea mays L. are reported for the first time. CO₂ compensation concentrations were significantly (P≤ 0.01) and negatively correlated with apparent photosynthesis at 300 μl CO₂/liter air. The Michaelis constant (as defined) for a leaf was significantly (P≤ 0.01) and positively correlated with apparent photosynthesis among varieties. While the first correlation is similar to behavior of CO₂ compensation among species of different photosynthetic efficiency, the latter correlation is the converse of the behavior of Km among species.     

Photosynthesis at low water potentials in sunflower: lack of photoinhibitory effects

The losses in chloroplast capacity to fix CO₂ when photosynthesis is reduced at low leaf water potential (ψ₁) have been proposed to result from photoinhibition. We investigated this possibility in soil-grown sunflower (Helianthus annuus L. cv IS894) using gas exchange techniques to measure directly the influence of light during dehydration on the in situ chloroplast capacity to fix CO₂. The quantum yield for CO₂ fixation as well as the rate of light- and CO₂-saturated photosynthesis were strongly inhibited at low ψ₁. The extent of inhibition was the same whether the leaves were exposed to high or to low light during dehydration. When intercellular partial pressures of CO₂ were decreased to the compensation point, which was lower than the partial pressures resulting from stomatal closure, the inhibition of the quantum yield was also unaffected. Photoinhibition could be observed only after high light exposures were imposed under nonphysiological low CO₂ and O₂ where both photosynthesis and photorespiration were suppressed. The experiments are the first to test whether gas exchange at low ψ₁ is affected by potentially photoinhibitory conditions and show that the loss in chloroplast capacity to fix CO₂ was entirely the result of a direct effect of water availability on chloroplast function and not photoinhibition.