Investigation on photorespiration with a sensitive C-assay

A leaf disk assay for photorespiration has been developed based on the rate of release of recently fixed ¹⁴CO₂ in light in a rapid stream of CO₂-free air at 30° to 35°. In tobacco leaves (Havana Seed) photorespiration with this assay is 3 to 5 times greater than the ¹⁴CO₂ output in the dark. In maize, photorespiration is only 2% of that in tobacco.

The importance of open leaf stomata, rapid flow rates of CO₂-free air, elevated temperatures, and oxygen in the atmosphere in order to obtain release into the air of a larger portion of the ¹⁴CO₂ evolved within the tissue in the light was established in tobacco. Photorespiration, but not dark respiration, was inhibited by α-hydroxy-2-pyridinemethanesulfonic acid, an inhibitor of glycolate oxidase, and by 3-(4-chlorophenyl)-1,1-dimethylurea (CMU), an inhibitor of photosynthetic electron transport, under conditions which did not affect the stomata. These experiments show that the substrates of photorespiration and dark respiration differ and also provide additional support for the role of glycolate as a major substrate of photorespiration. It was also shown that at 35° the quantity of ¹⁴CO₂ released in the assay may represent only 33% of the gross ¹⁴CO₂ evolved in the light, the remainder being recycled within the tissue.

It was concluded that maize does not evolve appreciable quantities of CO₂ in the light and that this largely accounts for the greater efficiency of net photosynthesis exhibited by maize. Hence low rates of photorespiration may be expected to be correlated with a high rate of CO₂ uptake at the normal concentrations of CO₂ found in air and at higher light intensities.

     

Metabolism of <Superscript>14</Superscript>C-glycine as a substrate for photorespiration of the leaf at different developmental stages of ephemeroides

The metabolism of ¹⁴C-glycine (a substrate for photorespiration) was studied in the light and in darkness under natural CO₂ concentration (0.03%) in the leaves of ephemeroides Scilla sibirica Haw. and Ficaria verna Huds. at different developmental stages. Using one and the same sample, potential photosynthesis (at 1% CO₂), true photosynthesis (at 0.03% CO₂), and leaf respiratory capacity were measured by the radiometric and manometric methods, respectively. All measurements were performed at 15°C, an average temperature during ephemer growth. It was found that, in the white zone of the Scilla leaf, the rate of CO₂ evolution resulting from metabolization of exogenous ¹⁴C-glycine was similar in the light and in darkness. In the green zone of the Scilla leaf and in the green leaf of Ficaria, both ¹⁴C-glycine absorption and ¹⁴CO₂ evolution were lower in the light as compared with darkness, which is explained by CO₂ reassimilation. In all treatments of both plant species, a specific inhibitor of glycine decarboxylase complex (GDC), aminoacetonitrile (5 mM) suppressed CO₂ evolution by 20–40%. It was concluded that in ephemeroides mitochondrial GDC, responsible for CO₂ evolution in photorespiration, is formed at the earliest stage of leaf development. This indicates that photorespiration can occur simultaneously with the development of the leaf photosynthetic activity. On the basis of the assumption that carbon losses in the form of CO₂ evolved during photorespiration comprise 25% of true photosynthesis, it was calculated that, in ephemer leaves, the highest rates of photorespiration and photosynthesis were attained during flowering when the leaf area was the largest and the rate of dark respiration was reduced by 1.5–2.0 times. The highest rates of dark respiration were observed in the beginning of growth. In senescing leaves by the end of the plant vegetation, potential photosynthesis and true photosynthesis were reduced, whereas dark respiration remained essentially unchanged. It is concluded that the high rates of potential and true photosynthesis are characteristic of ephemeroides when they complete their short developmental program in early spring (at 15°C); theoretically, photorespiration also occurs at a high rate during this period, when this process provides for a defense against the threat of photoinhibition at low temperature and high insolation.     

Variation in photorespiration. The effect of genetic differences in photorespiration on net photosynthesis in tobacco

The hypothesis that net photosynthesis is diminished in many plant species because of a high rate of CO₂ evolution in the light has been tested further. High rates of CO₂ output in CO₂-free air in comparison with dark respiration were found in Chlamydomonas reinhardi, wheat leaves, tomato leaves, and to a lesser extent in Chlorella pyrenoidosa by means of the ¹⁴C-photorespiration assay. In tobacco leaves high photorespiration was characteristic of a standard variety, Havana Seed, and a possibly still higher rate was found in a yellow heterozygous mutant, JWB Mutant. However, the dark homozygous sibling of the latter, JWB Wild, had a low photorespiration for the tobacco species. The relative rates of photorespiration were in the same sequence when measured by the ¹⁴CO₂ released in normal air from leaf disks supplied with glycolate-1-¹⁴C in the light.

As would be predicted by the hypothesis, the maximal net rate of photosynthesis at 300 ppm of CO₂ in the air in JWB Wild leaves was greater (24%) than in Havana Seed, while JWB Mutant had less CO₂ uptake than the standard variety (21%). At 550 ppm of CO₂ the differences in net photosynthesis were not as great between the 2 siblings as at 200 ppm. The relative leaf expansion rates of seedlings of the 3 tobacco varieties in a greenhouse had the same relationship as their rates of CO₂ assimilation.

Thus within the tobacco species, as in a comparison between tobacco and maize, low photorespiratory CO₂ evolution was correlated with higher photosynthetic efficiency. Therefore it seems that increased CO₂ uptake should be achieved by genetic interference with the process of photorespiration.

     

Influence of light intensity at different temperatures on rate of respiration of douglas-fir seedlings

The rate of photorespiration of Douglas-fir seedlings was measured under different light intensities by: (1) extrapolating the curve for CO₂ uptake in relation to atmospheric CO₂ content to zero CO₂ content, and (2) measuring CO₂ evolution of the plants into a CO₂-free airstream. Different results, obtained from these techniques, were believed to be caused by a severe restriction of the photosynthetic activity when the latter was used. With the first method, CO₂ evolution was lower than the dark respiration rate at low light intensity. For all temperatures studied (6°, 20°, 28°) a further increase in light intensity raised the CO₂ evolution above dark respiration before it leveled off. The rate of CO₂ evolution was stimulated by increase in temperature at all light intensities. With the CO₂-free air method, CO₂ evolution in the light was less than dark respiration at all light intensities.     

Oxygen and electron flow in C4 photosynthesis: Mehler reaction, photorespiration and CO2 concentration in the bundle sheath

The photosynthetic linear electron transport rate in excess of that used for CO₂ reduction was evaluated in Sorghum bicolor Moench. [NADP-malic enzyme (ME)-type C₄ plant], Amaranthus cruentus L. (NAD-ME-type C₄ plant) and Helianthus annuus L. (C₃ plant) leaves at different CO₂ and O₂ concentrations. The electron transport rate (J _F) was calculated from fluorescence using the light partitioning factor (relative PSII cross-section) determined under conditions where excess electron transport was assumed to be negligible: low light intensities, 500 μmol CO₂ · mol⁻¹ and 2% O₂. Under high light intensities there was a large excess of J _F/4 at 10–100% O₂ in the C₃ plant due to photorespiration, but very little in sorghum and somewhat more in amaranth, showing that photorespiration is suppressed, more in the NADP-ME- and less in the NAD-ME-type species. It is concluded that when C₄ photosynthesis is limited by supply of atmospheric CO₂ to the C₄ cycle, the C₃ cycle becomes limited by regeneration of ribulose 1,5-bisphosphate (RuBP) which in turn limits RuBP oxygenase activity and photorespiration. The rate of excess electron transport over that consumed for CO₂ fixation in C₄ plants was very sensitive to the presence of O₂ in the gas phase, rapidly increasing between 0.01 and 0.1% O₂, and at 2% O₂ it was about two-thirds of that at 21% O₂. This shows the importance of the Mehler O₂ reduction as an electron sink, compared with photorespiration in C₄ plants. However, the rate of the Mehler reaction is still too low to fully account for the extra ATP which is needed in C₄ photosynthesis. Received: 8 November 1997 / Accepted: 26 December 1997     

The effect on net photosynthesis of pedigree selection for low and high rates of photorespiration in tobacco

A normal appearing plant with a low rate of photorespiration (ratio of ¹⁴CO₂ released light/dark = 1.6) was found in an unselected tobacco (Nicotiana tabacum) cultivar. The plant was self-pollinated, and further selections were made on several successive generations. Excised leaves from the progeny of the selections were examined for photorespiration and net CO₂ assimilation in normal air during photosynthesis. Similar measurements were made of plants derived from selfed parents with high rates of photorespiration (ratio of ¹⁴CO₂ released light/dark = 3.0 or greater). Efficient photosynthetic plants (greater than 22.0 mg of CO₂ dm⁻² hr⁻¹) with low rates of photorespiration produced a larger proportion of efficient progeny (about 25%) than did selfing inefficient plants (about 6%), but this proportion did not increase in successive generations.     

Simultaneous gas exchange and fluorescence measurements indicate differences in the response of sunflower,bean and maize to water stress

Gas exchange and fluorescence measurements of attached leaves of water stressed bean, sunflower and maize plants were carried out at two light intensities (250 mol quanta m^-2s^-1 and 850 mol quanta m^-2s^-1). Besides the restriction of transpiration and CO₂ uptake, the dissipation of excess light energy was clearly reflected in the light and dark reactions of photosynthesis under stress conditions. Bean and maize plants preferentially use non-photochemical quenching for light energy dissipation. In sunflower plants, excess light energy gave rise to photochemical quenching. Autoradiography of leaves after photosynthesis in ¹⁴CO₂ demonstrated the occurrence of leaf patchiness in sunflower and maize but not in bean. The contribution of CO₂ recycling within the leaves to energy dissipation was investigated by studies in 2.5% oxygen to suppress photorespiration. The participation of different energy dissipating mechanisms to quanta comsumption on agriculturally relevant species is discussed.Abbreviations F_o minimal fluorescence - F_m maximal fluorescence - F_p peak fluorescence - g leaf conductance - P_N net CO₂ uptake - q_N coefficient of non-photochemical quenching - q_P coefficient of photochemical quenching     

Relationship between leaf development,carboxylase enzyme activities and photorespiration in the C4-plant Portulaca oleracea L.

Summary Ribulose diphosphate (RuDP) and (PEP) phosphoenolpyruvate carboxylase enzyme activities were studied in young, mature, and senescent Portulaca oleracea leaves. While the absolute amount of both the C₃ (RuDP) and C₄ (PEP) carboxylase is less in senescent leaves than in mature leaves, RuDP carboxylase activity is reduced to a lesser degree. In senescent leaves, PEP carboxylase activity equals 10% of that in mature tissue, but RuDP carboxylase is 27% of that in mature leaves. The same ontogenetic series was also used to determine photorespiration rates and responses to several gas treatments. Young and mature leaves were unaffected by changes in the light regime or oxygen concentrations, and exhibited typical C₄-plant light/dark ¹⁴CO₂ evolution ratios. Senescent leaves, on the other hand, have photorespiration ratios similar to C₃-plants. In addition, senescent leaves were affected by minus CO₂, 100% O₂ and N₂ in a manner expected of C₃-plants, but not C₄-plants. These results are discussed in terms of a relative increase in activity of the C₃ cycle in later developmental stages in this plant.Abbreviation RuDP ribulose diphosphate - PEP phosphoenolpyruvate - PGA phosphoglyceric acid     

Effect of carbon dioxide and temperature on photosynthetic CO2 uptake and photorespiratory CO2 evolution in sunflower leaves

Using an open gas-exchange system, apparent photosynthesis, true photosynthesis (TPS), photorespiration (PR) and dark respiration of sunflower (Helianthus annuus L.) leaves were determined at three temperatures and between 50 and 400 l/l external CO₂. The ratio of PR/TPS and the solubility ratio of O₂/CO₂ in the intercellular spaces both decreased with increasing CO₂. The rate of PR was not affected by the CO₂ concentration in the leaves and was independent of the solubility ratio of oxygen and CO₂ in the leaf cell. At photosynthesis-limiting concentrations of CO₂, the ratio of PR/TPS significantly increased from 18 to 30°C and the rate of PR increased from 4.3 mg CO₂ dm^-2 h^-1 at 18°C to 8.6 mg CO₂ dm^-2 h^-1 at 30°C. The specific activity of photorespired CO₂ was CO₂-dependent but temperature-independent, and the carbon traversing the glycolate pathway appeared to be derived both from recently fixed assimilate and from older reserve materials. It is concluded that PR as a percentage of TPS is affected by the concentrations of O₂ and CO₂ around the photosynthesizing cells, but the rate of PR may also be controlled by other factors.Abbreviations APS apparent photosynthesis (net CO₂ uptake) - PR photorespiration (CO₂ evolution in light) - RuBP ribulose-1,5-bisphosphate - TPS true photosynthesis (true CO₂ uptake)     

Photoinhibition of intact attached leaves of c(3) plants illuminated in the absence of both carbon dioxide and of photorespiration

When leaflets of bean and leaves of other species of C₃ plants are illuminated in the absence of CO₂ and at low O₂ partial pressure, the capacity for CO₂ assimilation at saturating light and its efficiency at low light intensities are inhibited. This photoinhibition is dependent on leaflet age and period of illumination. In young leaflets and following short exposure to these photoinhibitory conditions, some recovery of CO₂ assimilation capacity is observed immediately after treatment. Following substantial (70 to 80%) photoinhibition of CO₂ assimilation, recovery in fully expanded leaflets is observed only after 48 hours in normal air. The photoinhibition is largely prevented by providing CO₂ at partial pressures equivalent to the CO₂ compensation point, or by >210 millibars O₂ which permits internal CO₂ production by photorespiration. If leaflets are illuminated in 60 microbars CO₂ and 210 millibars O₂ (the CO₂ compensation point in air), no photoinhibition is observed. Electron transport processes and fluorescence emission associated with photosystem II are inhibited in chloroplast thylakoids isolated from leaflets after illumination in zero CO₂ and 10 millibars O₂. These studies support the hypothesis that CO₂ recycling through photorespiration is one means of effectively dissipating excess photochemical energy when CO₂ supply to illuminated leaves is limited.     

Advanced radiogasometric method for the determination of the rates of photorespiratory and respiratory decarboxylations of primary and stored photosynthates under steady-state photosynthesis

An advanced radiogasometric method for the study of plant leaf CO₂ exchange is presented. The method enables determination of the rates of CO₂ fixation, photorespiration and respiration in the light under steady‐state photosynthesis and discrimination between primary and stored photosynthates as substrates of photorespiratory and respiratory decarboxylations. The method is based on the analysis of the time curves of ¹⁴CO₂ evolution from labeled primary and stored photosynthates in leaves previously exposed to ¹⁴CO₂. The molar rates of different decarboxylation reactions are calculated from the initial slopes of the curves taking into account the specific radioactivity of CO₂ fed to leaves and/or evolved from leaves. To estimate the contribution of primary and stored photosynthates, the measurements of ¹⁴CO₂ evolution are performed after feeding plant leaves for different periods with ¹⁴CO₂. Photorespiration and respiration are distinguished on the basis of data obtained from measurements of ¹⁴CO₂ evolution under normal (210 ml l⁻¹) and low (15 ml l⁻¹) concentrations of oxygen. A principally new method for the determination of the rate of intracellular refixation of respiratory CO₂ has been developed. The method is based on the measurements of ¹⁴CO₂ evolution from leaves into the medium of very high concentrations (30 ml l⁻¹) of ¹²CO₂, where the probability of refixation of ¹⁴CO₂ evolved inside the cell is close to zero. The results obtained were comparable with the data derived from parallel refixation measurements by means of gasometric methods. As an example of application, the data on CO₂ exchange in leaves of two contrasting groups of C₃‐species, differing in the ability of starch accumulation, are presented.     

Effect of light intensity on partitioning of photosynthetic electron transport to photorespiration in four subtropical forest plants

Photosynthetic rate (P_n) and the partitioning of noncyclic photosynthetic electron transport to photorespiration (J_O) in seedlings of four subtropical woody plants growing at three light intensities were studied in the summer time by measurements of chlorophyll fluorescence and CO₂ exchange. ExceptSchima superba, an upper canopy tree species, the tree speciesCastanopsis fissa and two understory shrubsPsychotria rubra, Ardisia quinquegona had the highestP _n at 36% of sunlight intensity. The total photosynthetic electron transport rate (J_F) and the ratio ofJ _O/J_F were elevated in leaves under full sunlight.J _O/J_F ratio reached 0.5–0.6 and coincided with the increasing of oxygenation rate of Rubisco (V_O), the activity of glycolate oxidase and photorespiration rate at full sunlight. It is suggested that an increasing partitioning proportion of photosynthetic electron transport to photorespiration might be one of the protective regulation mechanisms in forest plant under strong summer light and high temperature conditions.     

Variations in light energy dissipation in <Emphasis Type="Italic">Woodfordia fruticosa</Emphasis> leaves during expansion

Young leaves of tropical trees frequently appear red in color, with the redness disappearing as the leaves mature. During leaf expansion, plants may employ photoprotective mechanisms to cope with high light intensities; however, the variations in anthocyanin contents, nonphotochemical quenching (NPQ), and photorespiration during leaf expansion are poorly understood. Here, we investigated pigment contents, gas exchange, and chlorophyll (Chl) fluorescence in Woodfordia fruticosa leaves during their expansion. Young red leaves had significantly lower Chl content than that of expanding or mature leaves, but they accumulated significantly higher anthocyanins and dissipated more excited light energy through NPQ. As the leaves matured, net photosynthetic rate, total electron flow through PSII, and electron flow for ribulose-1,5-bisphosphate oxygenation gradually increased. Our results provided evidence that photorespiration is of fundamental importance in regulating the photosynthetic electron flow and CO₂ assimilation during leaf expansion.     

Photosynthesis and photorespiration in algae

The CO₂ exchange of several species of fresh water and marine algae was measured in the laboratory to determine whether photorespiration occurs in these organisms. The algae were positioned as thin layers on filter paper and the CO₂ exchange determined in an open gas exchange system. In either 21 or 1% O₂ there was little difference between ¹⁴CO₂ and ¹²CO₂ uptake. Apparent photosynthesis was the same in 2, 21, or 50% O₂. The compensation points of all algae were less than 10 μl 1⁻¹. CO₂ or ¹⁴CO₂ evolution into CO₂-free air in the light was always less than the corresponding evolution in darkness. These observations are inconsistent with the proposal that photorespiration exists in these algae.     

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.     

Adaptations of the Photosynthetic Mechanism of Sugar Maple (Acer saccharum) Seedlings Grown in Various Light Intensities

Sugar maple (Acer saccharum Marsh.) seedlings were grown in a nursery for three years in 13, 25, 45 and 100 per cent of full daylight. During the third year of growth, the rates of their apparent photosynthesis and respiration were measured periodically with an infra-red gas analyzer at various light intensities and normal CO₂ concentration. In addition, the rates of apparent photosynthesis of a single attached leaf of the same seedlings were measured at saturating light intensity, hut varying CO₂ concentrations. An increase in the light intensity in which seedlings were grown had no effect on their height or mean leaf area, hut resulted in thicker leaves, an increase in the total leaf area per seedling due to an increase in the number of leaves, an increase in the dry weight especially of roots and a decrease in the chlorophyll content of leaves. Throughout the growing season seedlings grown in full daylight, as compared with those grown in lower light intensities, had the lowest rates of apparent photosynthesis measured at standard conditions (21,600 lux light intensity and 300 ul/l of CO₂), when this was expressed per unit leaf area, hut the highest rates on a per seedling basis. Thus dry matter production attained at the end of the growing season correlated positively with the photosynthetic rate per seedling, but not per unit leaf area. The rates of apparent photosynthesis of seedlings grown at lower light intensities were more responsive to changes in light intensity or CO₂ concentration than those of seedlings grown in full daylight intensity.     

Comparison of the Oxygen Exchange between Photosynthetic Cell Suspensions and Detached Leaves of Euphorbia characias L

Using a mass-spectrometric ¹⁶O₂/¹⁸O₂-isotope technique, we compared the nature and the relative importance of oxygen exchange in photomixotrophic (PM) and photoautotrophic (PA) suspensions of Euphorbia characias L. with those in intact leaves of the same species. Young and mature leaves, dividing and nondividing cell suspensions were characterized in short-term experiments. On chlorophyll basis, the gross photosynthetic activities at CO₂ saturating concentration of PA and PM suspensions varied little from those of leaves. On dry weight basis, gross photosynthesis of PA suspensions was equal to that of leaves because of their similar chlorophyll content. This was not the case in PM suspensions where gross photosynthesis was lower and largely varied during the growth cycle. The CO₂ compensation point of PA cells (155-265 parts per million) was much higher than that of leaves (50-80 ppm). Oxygen uptakes were analyzed in terms of mitochondrial respiration, photorespiration and light stimulation of oxygen uptake (LSOU), often identified to Mehlertype reactions. In PA and PM suspensions, mitochondrial respiration rates were higher than in leaves by a factor of 1.5 to 4.5. In PM suspensions, photorespiration and LSOU were observed only in nondividing cells. Photorespiration and LSOU rates were comparable in PA suspensions and leaves. Our results demonstrate that photorespiration of PA suspensions has not been affected by the 2% CO₂ concentration imposed during 2 years of culture.     

A Transient Burst of CO(2) from Geranium Leaves during Illumination at Various Light Intensities as a Measure of Photorespiration

A transient CO₂ burst is exhibited by irradiated leaves of the C₃ plant geranium (Pelargonium X hortorum, Bailey) after the irradiance is quickly lowered. The light CO₂ burst appears to be related to photorespiration because of its irradiance dependency and its sensitivity to other environmental components such as CO₂ and O₂ concentration. The term post-lower-irradiance CO₂ burst or PLIB is used to describe the phenomenon. The PLIB appears to be a quantitative measurement of photorespiration with intact geranium leaves. The PLIB has been observed with intact leaves of other C₃ plants but not with C₄ leaves. Therefore, it is proposed that, after maximizing intact leaf photosynthetic rates and leaf chamber gas measuring conditions, photorespiration can be measured with intact C₃ leaves such as geranium as a transient post-lower-irradiance CO₂ burst.     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

Photosynthesis, Photorespiration and Respiration of Detached Spruce Twigs as Influenced by Oxygen Concentration and Light Intensity

The effects of oxygen concentration and light intensity on the rates of apparent photosynthesis, true photosynthesis, photorespiration and dark respiration of detached spruce twigs were determined by means of an infra-red carbon dioxide analyzer (IRCA). A closed circuit system IRCA was filled with either 1 per cent of oxygen in nitrogen, air (21 % O₂) or pure oxygen (100 % O₂). Two light intensities 30 × 10³ erg · cm ^?2· s^?1 and 120 × 10³ erg · cm^?2· s^?1 were applied. It has been found that the inhibitory effect of high concentration of oxygen on the apparent photosynthesis was mainly a result of a stimulation of the rate of CO₂ production in light (photorespiration). In the atmosphere of 100 % O₂, photorespiration accounts for 66–80 per cent of total CO₂ uptake (true photosynthesis). Owing to a strong acceleration of photorespiration by high oxygen concentrations, the rate of true photosynthesis calculated as the sum of apparent photosynthesis and photorespiration was by several times less inhibited by oxygen than the rate of apparent photosynthesis. The rates of dark respiration were essentially unaffected by the oxygen concentrations used in the experiments. An increase in the intensity of light from 30 × 10³ erg · cm^?3· s^?1 to 120 · 10³ erg · cm^?2· s^?1 enhanced the rate of photorespiration in the atmospheres of 21 and 100 % oxygen but not in 1 % O₂. The rate of apparent photosynthesis, however, was little affected by light intensity in an atmosphere of 1 % oxygen.