Control of carbon partitioning and photosynthesis by the triose phosphate/phosphate translocator in transgenic tobacco plants (Nicotiana tabacum). II. Assessment of control coefficients of the triose phosphate/phosphate translocator

 Transgenic tobacco (Nicotiana tabacum L.) plants with decreased and increased transport capacities of the chloroplast triose phosphate/phosphate translocator (TPT) were used to study the control the TPT exerts on the flux of starch and sucrose biosynthesis, as well as CO₂ assimilation, respiration and photosynthetic electron transport. For this purpose, tobacco lines with an antisense repression of the endogenous TPT (αTPT) and tobacco lines overexpressing a TPT gene from Flaveria trinervia (FtTPT) were used. In ambient CO₂, there was no or little effect of altered TPT transport activities on either rates of photosynthetic electron transport and/or CO₂ assimilation. However, in elevated CO₂ (1500 μl · l⁻¹) and low O₂ (2%) the TPT exerted strong control on the rate of CO₂ assimilation (control coefficient for the wild type; C^JA _TPT=0.30) in saturating light. Similarly, the incorporation of ¹⁴C into starch in high CO₂ was increased in tobacco plants with decreased TPT activity, but was reduced in plants overexpressing the TPT from F. trinervia. Thus, the TPT exerted negative control on the rate of starch biosynthesis with a C^JStarch _TPT=−0.19 in the wild type estimated from a hyperbolic curve fitted to the data points. This was less than the positive control strength on the rate of sucrose biosynthesis (C^JSuc _TPT=0.35 in the wild type). Theoretically, the positive control exerted on sucrose biosynthesis should be numerically identical to the negative control on starch biosynthesis unless additional metabolic pathways are affected. The rate of dark respiration showed some correlation with the TPT activity in that it increased in FtTPT overexpressors, but decreased in αTPT plants with an apparent control coefficient of C^JRes _TPT=0.24. If the control on sucrose biosynthesis is referred to as “gain of carbon” (positive control) and the control on starch biosynthesis as well as dark respiration as a “loss of carbon” (negative control) for sucrose biosynthesis and subsequent export, the sum of the control coefficients on dark respiration and starch biosynthesis would be numerically similar to the control coefficient on the rate of sucrose biosynthesis. There was also some control on the rate of photosynthetic electron transport, but only at high light and in elevated CO₂ combined with low O₂. The control coefficient for the rate of photosynthetic electron transport was C^JETR _TPT=0.16 in the wild type. Control coefficients were also calculated for plants with elevated and lowered TPT activity. Furthermore, the extent to which starch degradation/glucose utilisation compensates for the lack of triose phosphate export was assessed. The TPT also exerted control on metabolite contents in air. Received: 26 March 1999 / Accepted: 21 August 1999     

δ13C of CO2 respired in the dark in relation to δ13C of leaf metabolites: comparison between Nicotiana sylvestris and Helianthus annuus under drought

The variations of δ¹³C in leaf metabolites (lipids, organic acids, starch and soluble sugars), leaf organic matter and CO₂ respired in the dark from leaves of Nicotiana sylvestris and Helianthus annuus were investigated during a progressive drought. Under well‐watered conditions, CO₂ respired in the dark was ¹³C‐enriched compared to sucrose by about 4‰ in N. sylvestris and by about 3‰ and 6‰ in two different sets of experiments in H. annuus plants. In a previous work on cotyledonary leaves of Phaseolus vulgaris, we observed a constant ¹³C‐enrichment by about 6‰ in respired CO₂ compared to sucrose, suggesting a constant fractionation during dark respiration, whatever the leaf age and relative water content. In contrast, the ¹³C‐enrichment in respired CO₂ increased in dehydrated N. sylvestris and decreased in dehydrated H. annuus in comparison with control plants. We conclude that (i) carbon isotope fractionation during dark respiration is a widespread phenomenon occurring in C₃ plants, but that (ii) this fractionation is not constant and varies among species and (iii) it also varies with environmental conditions (water deficit in the present work) but differently among species. We also conclude that (iv) a discrimination during dark respiration processes occurred, releasing CO₂ enriched in ¹³C compared to several major leaf reserves (carbohydrates, lipids and organic acids) and whole leaf organic matter.     

Diffusional Contribution to Carbon Isotope Fractionation during Dark CO(2) Fixation in CAM Plants

A mathematical model is developed which can be used to predict in vivo carbon isotope fractionations associated with carbon fixation in plants in terms of diffusion, CO₂ hydration, and carboxylation components. This model also permits calculation of internal CO₂ concentration for comparison with results of gas-exchange experiments. The isotope fractionations associated with carbon fixation in Kalanchoë daigremontiana and Bryophyllum tubiflorum have been measured by isolation of malic acid following dark fixation and enzymic determination of the isotopic composition of carbon-4 of this material. Corrections are made for residual malic acid, fumarase activity, and respiration. Comparison of these data with calculations from the model indicates that the rate of carbon fixation is limited principally by diffusion, rather than by carboxylation. Processes subsequent to the initial carboxylation also contribute to the over-all isotopic composition of the plant.     

Process-based isotope partitioning of winter soil respiration in a subalpine ecosystem reveals importance of rhizospheric respiration

Deep snow in sub-alpine ecosystems may reduce or eliminate soil freezing, thus contributing to the potential for winter soil respiration to account for a significant fraction of annual CO₂ efflux to the atmosphere. Quantification of carbon loss from soils requires separation of respiration produced by roots and rhizosphere organisms from that produced by heterotrophic, decomposer organisms because the former does not result in a net loss of stored carbon. Our objective was to quantify winter soil respiration rates in a sub-alpine forest and meadow, and to partition that flux into its rhizosphere and heterotrophic components. We were particularly interested in comparing early winter soil respiration to late winter/early spring soil respiration of each component because previous work has shown a consistent increase in soil respiration of subalpine systems from early winter to late winter/spring. Field data on the total soil CO₂ flux and its carbon isotope composition were coupled with data from laboratory incubations using a novel process-based stable isotope mixing model implemented in a hierarchical Bayesian framework. We found that soil respiration generally increased from early to later winter and was greatest mid-summer. After correcting for the effect of wind on snowpack δ¹³C–CO₂, the δ¹³C of soil-respired CO₂ varied little over winter, and the contributions of rhizospheric (~35 %) and heterotrophic (~65 %) respiration were relatively constant. The significance of winter respiration from the rhizosphere and apparent coupling of increases in rhizospheric and heterotrophic respiration in late winter are likely to be important for predicting changes in soil carbon in sub-alpine ecosystems.     

The influence of increasing growth temperature and CO2 concentration on the ratio of respiration to photosynthesis in soybean seedlings

Using controlled environmental growth chambers, whole plants of soybean, cv. ‘Clark’, were examined during early development (7–20 days after sowing) at both ambient (≈ 350 μL L^–1) and elevated (≈ 700 μL L^–1) carbon dioxide and a range of air temperatures (20, 25, 30, and 35 °C) to determine if future climatic change (temperature or CO₂ concentration) could alter the ratio of carbon lost by dark respiration to that gained via photosynthesis. Although whole-plant respiration increased with short-term increases in the measurement temperature, respiration acclimated to increasing growth temperature. Respiration, on a dry weight basis, was either unchanged or lower for the elevated CO₂ grown plants, relative to ambient CO₂ concentration, over the range of growth temperatures. Levels of both starch and sucrose increased with elevated CO₂ concentration, but no interaction between CO₂ and growth temperature was observed. Relative growth rate increased with elevated CO₂ concentration up to a growth temperature of 35 °C. The ratio of respiration to photosynthesis rate over a 24-h period during early development was not altered over the growth temperatures (20–35 °C) and was consistently less at the elevated relative to the ambient CO₂ concentration. The current experiment does not support the proposition that global increases in carbon dioxide and temperature will increase the ratio of respiration to photosynthesis; rather, the data suggest that some plant species may continue to act as a sink for carbon even if carbon dioxide and temperature increase simultaneously.     

The effect of carbon dioxide concentration on respiration of growing and mature soybean leaves

Soybean plants were grown continuously at 350 and 700cm³m^?3 CO₂ at constant temperature. Respiration rates of third trifoliolate leaves were measured at the growth CO₂ concentration for the whole dark period from 5d before through to 5d after full area expansion. The short-term response of respiration rate to the measurement CO₂ concentration was also determined at each age. Respiration rates per unit of dry mass declined with age and were significantly less at a given age or RGR in leaves grown and measured at the elevated CO₂. The difference in respiration rate was largest in mature leaves and resulted from the different measurement CO₂ concentrations. The respiratory costs of the tissue synthesis, estimated from the elemental composition of the tissue, did not differ substantially between CO₂ treatments. The response of respiration rate to carbon dioxide concentration was not strongly affected by the form of nitrogen supplied. Maintenance respiration calculated by subtracting growth respiration from total respiration was negative in rapidly growing leaves for both CO₂ treatments. This indicates that CO₂ efflux in the dark does not accurately reflect the average 24 h rate of energy expenditure on growth and maintenance for soybean leaves.     

On the 13C/12C isotopic signal of day and night respiration at the mesocosm level

While there is currently intense effort to examine the ¹³C signal of CO₂ evolved in the dark, less is known on the isotope composition of day‐respired CO₂. This lack of knowledge stems from technical difficulties to measure the pure respiratory isotopic signal: day respiration is mixed up with photorespiration, and there is no obvious way to separate photosynthetic fractionation (pure c_i/c_a effect) from respiratory effect (production of CO₂ with a different δ¹³C value from that of net‐fixed CO₂) at the ecosystem level. Here, we took advantage of new simple equations, and applied them to sunflower canopies grown under low and high [CO₂]. We show that whole mesocosm‐respired CO₂ is slightly ¹³C depleted in the light at the mesocosm level (by 0.2–0.8‰), while it is slightly ¹³C enriched in darkness (by 1.5–3.2‰). The turnover of the respiratory carbon pool after labelling appears similar in the light and in the dark, and accordingly, a hierarchical clustering analysis shows a close correlation between the ¹³C abundance in day‐ and night‐evolved CO₂. We conclude that the carbon source for respiration is similar in the dark and in the light, but the metabolic pathways associated with CO₂ production may change, thereby explaining the different ¹²C/¹³C respiratory fractionations in the light and in the dark.     

Soil respiration in northern forests exposed to elevated atmospheric carbon dioxide and ozone

The aspen free-air CO₂ and O₃ enrichment (FACTS II–FACE) study in Rhinelander, Wisconsin, USA, is designed to understand the mechanisms by which young northern deciduous forest ecosystems respond to elevated atmospheric carbon dioxide (CO₂) and elevated tropospheric ozone (O₃) in a replicated, factorial, field experiment. Soil respiration is the second largest flux of carbon (C) in these ecosystems, and the objective of this study was to understand how soil respiration responded to the experimental treatments as these fast-growing stands of pure aspen and birch + aspen approached maximum leaf area. Rates of soil respiration were typically lowest in the elevated O₃ treatment. Elevated CO₂ significantly stimulated soil respiration (8–26%) compared to the control treatment in both community types over all three growing seasons. In years 6–7 of the experiment, the greatest rates of soil respiration occurred in the interaction treatment (CO₂ + O₃), and rates of soil respiration were 15–25% greater in this treatment than in the elevated CO₂ treatment, depending on year and community type. Two of the treatments, elevated CO₂ and elevated CO₂ + O₃, were fumigated with ¹³C-depleted CO₂, and in these two treatments we used standard isotope mixing models to understand the proportions of new and old C in soil respiration. During the peak of the growing season, C fixed since the initiation of the experiment in 1998 (new C) accounted for 60–80% of total soil respiration. The isotope measurements independently confirmed that more new C was respired from the interaction treatment compared to the elevated CO₂ treatment. A period of low soil moisture late in the 2003 growing season resulted in soil respiration with an isotopic signature 4–6‰ enriched in ¹³C compared to sample dates when the percentage soil moisture was higher. In 2004, an extended period of low soil moisture during August and early September, punctuated by a significant rainfall event, resulted in soil respiration that was temporarily 4–6‰ more depleted in ¹³C. Up to 50% of the Earth’s forests will see elevated concentrations of both CO₂ and O₃ in the coming decades and these interacting atmospheric trace gases stimulated soil respiration in this study.     

Deep Autotrophic Soil Respiration in Shrubland and Woodland Ecosystems in Central New Mexico

Quantifying the controls on soil respiration is important for understanding ecosystem physiology and for predicting the response of soil carbon reservoirs to climate change. The majority of soil respiration is typically considered to occur in the top 20–30 cm of soils. In desert soils, where organic matter concentrations tend to be low and plants are deeply rooted, deeper respiration might be expected. However, little is known about the depth distribution of respiration in dryland soils. Here we show that the average depth of soil respiration between pulse precipitation events is almost always greater than 20 cm and is frequently greater than 50 cm in two central New Mexico desert shrublands. The average depth of soil respiration in a pi?on-juniper woodland was shallower, between 5 and 40 cm. In the shrublands, 8‰ seasonal variations in the carbon isotope composition of soil-respired CO₂ (δ¹³C_r-soil) that correlate with vapor pressure deficit support root/rhizosphere respiration as the dominant source of soil CO₂. Such deep autotrophic respiration indicates that shrubs preferentially allocate photosynthate to deep roots when conditions near the surface are unfavorable. Therefore, respiration rates in these soils are not necessarily correlated with root biomass. The δ¹³C_r-soil values provide no evidence for CO₂ evolved from soil inorganic carbon. Our results also suggest that organic carbon cycling is rapid and efficient in these soils and that the δ¹³C value of CO₂ respired from soils in much of the southwestern US, and perhaps in other semiarid regions, varies seasonally by at least 4‰.     

Inferring biogenic and anthropogenic carbon dioxide sources across an urban to rural gradient

We continuously monitored CO₂ concentrations at three locations along an urban-to-rural gradient in the Salt Lake Valley, Utah from 2004 to 2006. The results showed a range of CO₂ concentrations from daily averages exceeding 500 p.p.m. at the city center to much lower concentrations in a non-urbanized, rural region of the valley. The highest values were measured in the wintertime and under stable atmospheric conditions. At all three sites, we utilized weekly measurements of the C and O isotope composition of CO₂ for a 1-year period to evaluate the CO₂ sources underlying spatial and temporal variability in CO₂ concentrations. The results of an inverse analysis of CO₂ sources and the O isotope composition of ecosystem respiration (δ¹⁸ O _R) showed large contributions (>50%) of natural gas combustion to atmospheric CO₂ in the wintertime, particularly at the city center, and large contributions (>60%) of biogenic respiration to atmospheric CO₂ during the growing season, particularly at the rural site. δ¹⁸ O _R was most enriched at the rural site and more isotopically depleted at the urban sites due to the effects of irrigation on ecosystem water pools at the urban sites. The results also suggested differences in the role of leaf versus soil respiration between the two urban sites, with seasonal variation in the contribution of leaf respiration at a residential site and relatively constant contributions of leaf respiration at the city center. These results illustrate that spatial and temporal patterns of urban CO₂ concentrations and isotopic composition can be used to infer patterns of energy use by urban residents as well as plant and soil processes in urban areas.     

Photosynthesis, Respiration and Post-illumination Fixation of CO2 by Corn Leaves as Influenced by Light and Oxygen Concentration

The effect of oxygen concentration on the rate of CO₂-uptake in continuous and intermittent light was studied as well as the CO₂-fixation during a short dark period after light was turned off. In addition the dark respiration and the CO₂-compensation point of attached and detached corn leaves were determined. Leaves of 4 to 22-day old plants were used as experimental material. A closed circuit system of an infrared carbon dioxide analyzer was employed to measure the rate of CO₂-exchange. It was found that in an atmosphere consisting of 100 % oxygen, there was about 50 per cent inhibition of the rate of CO₂-uptake in continuous and intermittent light compared to that in an atmosphere consisting of 21% oxygen. The same was true of the rate of CO₂-fixation in darkness during a short period after the light was turned off. Since the response to oxygen concentration of the CO₂-uptake in light and of the CO₂-fixation in darkness after the light was turned off were similar, it is concluded that the fixation of CO₂ in the short dark period represents an over- shoot of photosynthesis. The rate of dark respiration was little affected by the oxygen concentration in the ranges used in the experiments. The carbon dioxide compensation point which has been observed in leaves of 4 to 14-day old plants was not influenced by either oxygen concentration or light intensity. Since the changes in the rate of CO₂-uptake due to changes in the concentration of oxygen and light intensity had no effect on the CO₂-compensation point, it is concluded that a reabsorption of respiratory CO₂ by photosynthesis could not account for the low value of this point. These results are interpreted as a further corroboration of the statement that the leaves of corn lack the process of photorespiration and that dark respiration is inhibited in light. It was observed that the rate of the CO₂-uptake gradually increased in plants which were from 4 to 22-days old. The inhibitory effect of high concentration of oxygen on the rate of CO₂-uptake was relatively higher in old plants than in young ones.     

The use of stable isotopes to study ecosystem gas exchange

Stable isotopes are a powerful research tool in environmental sciences and their use in ecosystem research is increasing. In this review we introduce and discuss the relevant details underlying the use of carbon and oxygen isotopic compositions in ecosystem gas exchange research. The current use and potential developments of stable isotope measurements together with concentration and flux measurements of CO₂ and water vapor are emphasized. For these applications it is critical to know the isotopic identity of specific ecosystem components such as the isotopic composition of CO₂, organic matter, liquid water, and water vapor, as well as the associated isotopic fractionations, in the soil-plant- atmosphere system. Combining stable isotopes and concentration measurements is very effective through the use of ”Keeling plots.” This approach allows the identification of the isotopic composition and the contribution of ecosystem, or ecosystem components, to the exchange fluxes with the atmosphere. It also allows the estimation of net ecosystem discrimination and soil disequilibrium effects. Recent modifications of the Keeling plot approach permit examination of CO₂ recycling in ecosystems. Combining stable isotopes with dynamic flux measurements requires precision in isotopic sampling and analysis, which is currently at the limit of detection. Combined with the micrometeorological gradient approach (applicable mostly in grasslands and crop fields), stable isotope measurements allow separation of net CO₂ exchange into photosynthetic and soil respiration components, and the evapotranspiration flux into soil evaporation and leaf transpiration. Similar applications in conjunction with eddy correlation techniques (applicable to forests, in addition to grasslands and crop fields) are more demanding, but can potentially be applied in combination with the Keeling plot relationship. The advance and potential in using stable isotope measurements should make their use a standard component in the limited arsenal of ecosystem-scale research tools. Received: 8 July 1999 / Accepted: 10 January 2000     

Effects of Prolonged Darkness on the Sensitivity of Leaf Respiration to Carbon Dioxide Concentration in C3and C4Species

Predicting responses of plant and global carbon balance to theincreasing concentration of carbon dioxide in the atmosphererequires an understanding of the response of plant respirationto carbon dioxide concentration ([CO₂]). Direct effects of thecarbon dioxide concentration at which rates of respiration ofplant tissue are measured are quite variable and their effectsremain controversial. One possible source of variation in responsivenessis the energy status of the tissue, which could influence thecontrol coefficients of enzymes, such as cytochrome-c oxidase,whose activity is sensitive to [CO₂]. In this study we comparedresponses of respiration rate to [CO₂] over the range of 60to 1000 µmol mol^-1in fully expanded leaves of four C₃andfour C₄herbaceous species. Responses were measured near themiddle of the normal 10 h dark period, and also after another24 h of darkness. On average, rates of respiration were reducedabout 70% by the prolonged dark period, and leaf dry mass perunit area decreased about 30%. In all species studied, the relativedecrease in respiration rate with increasing [CO₂] was largerafter prolonged darkness. In the C₃species, rates measured at1000 µmol mol^-1CO₂averaged 0.89 of those measured at 60µmol mol^-1in the middle of the normal dark period, and0.70-times when measured after prolonged darkness. In the C₄species,rates measured at 1000 µmol mol^-1CO₂averaged 0.79 of thoseat 60 µmol mol^-1CO₂in the middle of the normal dark period,and 0.51-times when measured after prolonged darkness. In threeof the C₃species and one of the C₄species, the decrease in theabsolute respiration rate between 60 and 1000 µmol mol^-1CO₂wasessentially the same in the middle of the normal night periodand after prolonged darkness. In the other species, the decreasein the absolute rate of respiration with increase in [CO₂] wassubstantially less after prolonged darkness than in the middleof the normal night period. These results indicated that increasingthe [CO₂] at the time of measurement decreased respiration inall species examined, and that this effect was relatively largerin tissues in which the respiration rate was substrate-limited.The larger relative effect of [CO₂] on respiration in tissuesafter prolonged darkness is evidence against a controlling roleof cytochrome-c oxidase in the direct effects of [CO₂] on respiration.Copyright 2001 Annals of Botany Company Carbon dioxide, respiration, Abutilon theophrasti(L.), Amaranthus retroflexus(L.),Amaranthus hypochondriacus (L.), Datura stramonium(L.), Helianthus annuus(L.), Solanum melongena(L.), Sorghum bicolor(L. Moench), Zea mays     

Influence of High Temperature on End-of-Season Tundra CO2 Exchange

The high-arctic terrestrial environment is generally recognized as one of the world's most sensitive areas with regard to global warming. In this study, we examined the influence of an isolated warm period on net ecosystem carbon dioxide (CO₂) exchange at high latitude during autumn. Using the Free Air Temperature Increase (FATI) technique, we manipulated air, soil, and vegetation temperatures in late August in a tundra site at Zackenberg in the National Park of North and East Greenland (74°N 21°W). The consequences for gross canopy photosynthesis, canopy respiration, and belowground respiration of increasing these temperatures by approximately 2.5°C were determined with closed dynamic CO₂ exchange systems. Under current temperatures, the ecosystem acted as a net CO₂ source, releasing 19 g CO₂-C m⁻² over the 14-day study period. Warm soils and senescing vegetation in autumn were unequivocally responsible for this efflux. Heating enhanced CO₂ efflux to 29 g CO₂-C m⁻². This effect was attributed to a 39% increase in belowground respiration, which was the main component of the carbon (C) budget. Gross photosynthesis, on the other hand, was not affected significantly by the simulated warming. Although the aftereffects of an isolated warm period on the C balance in early winter could be significant, simulations with a simple C budget model suggest that soil carbon pools are not affected to a great extent by such a climatic disturbance. The influence on atmospheric carbon, however, appears to be significant. Received 9 June 2000; accepted 20 December 2000.     

About effect of processes in the earth crust on evolution of photosynthesis (as indicated by data on carbon isotopic composition)

A probable mechanism of effect of processes occurring in the Earth crust on evolution of photosynthesis is considered. According to the hypothesis, this effect is realized through entrance to the Earth atmosphere of carbon dioxide that stimulates photosynthesis. Supply of CO₂ is irregular and is due to irregular movements of the Earth crust plates. This is accompanied by destruction of carbonates and conversion of carbon of the organic matter to CO₂ due to processes of reduction of sulfates. The CO₂ content in atmosphere rises for relatively short orogenic periods, due to intensive crust plate movement, while for the subsequent long periods, called the geosynclinal ones, of the relatively slow plate movement, the CO₂ content falls due to the higher rate of its consumption for photosynthesis. Owing to the carbon isotopic fractionation accompanying photosynthesis, regular isotopic differences appear between the atmospheric CO₂ and the “living” matter (Relay’s effect); these differences are then transformed to isotope differences of the carbonate and organic carbon. At the appearance in atmosphere of free oxygen-product of photosynthesis-in organisms there appears photorespiration that also is accompanied by fractionation of carbon isotopes, but with effect of opposite sign. This leads to enrichment of the photosynthesizing biomass with ¹³C isotope at the orogenic periods. As a result, the initially pronounced isotope differences of the carbonate and organic carbon decrease by the end of the geosynclinal periods. According to the proposed model, concentrations of CO₂ and O₂ are exchanged in the antiphase. They lead to alternation of periods of warning up and cooling off on the Earth. The former coincide with the orogenic periods, the latter appear at the end of geosynclinal periods when in atmosphere there is accumulated oxygen, while in sediments-organic substance. The accumulation of organic substance leads to formation of petroleum-maternal masses. To substantiate the model, data on isotope composition of carbon of carbonate and organic substance of rocks are used and its ability to explain several known natural regularities and empirical correlations. The model is used for analysis of some key stages of evolution of photosynthesis.     

Two distinct plant respiratory physiotypes might exist which correspond to fast-growing and slow-growing species

The origin of the carbon atoms in CO₂ respired by leaves in the dark of several plant species has been studied using ¹³C/¹²C stable isotopes. This study was conducted using an open gas exchange system for isotope labeling that was coupled to an elemental analyzer and further linked to an isotope ratio mass spectrometer (EA–IRMS) or coupled to a gas chromatography–combustion-isotope ratio mass spectrometer (GC–C-IRMS). We demonstrate here that the carbon, which is recently assimilated during photosynthesis, accounts for nearly ca. 50% of the carbon in the CO₂ lost through dark respiration (R_d) after illumination in fast-growing and cultivated plants and trees and, accounts for only ca. 10% in slow-growing plants. Moreover, our study shows that fast-growing plants, which had the largest percentages of newly fixed carbon of leaf-respired CO₂, were also those with the largest shoot/root ratios, whereas slow-growing plants showed the lowest shoot/root values.     

Relationship between Photosynthesis and Respiration: The Effect of Carbohydrate Status on the Rate of CO(2) Production by Respiration in Darkened and Illuminated Wheat Leaves

The rate of dark CO₂ efflux from mature wheat (Triticum aestivum cv Gabo) leaves at the end of the night is less than that found after a period of photosynthesis. After photosynthesis, the dark CO₂ efflux shows complex dependence on time and temperature. For about 30 minutes after darkening, CO₂ efflux includes a large component which can be abolished by transferring illuminated leaves to 3% O₂ and 330 microbar CO₂ before darkening. After 30 minutes of darkness, a relatively steady rate of CO₂ efflux was obtained. The temperature dependence of steady-state dark CO₂ efflux at the end of the night differs from that after a period of photosynthesis. The higher rate of dark CO₂ efflux following photosynthesis is correlated with accumulated net CO₂ assimilation and with an increase in several carbohydrate fractions in the leaf. It is also correlated with an increase in the CO₂ compensation point in 21% O₂, and an increase in the light compensation point. The interactions between CO₂ efflux from carbohydrate oxidation and photorespiration are discussed. It is concluded that the rate of CO₂ efflux by respiration is comparable in darkened and illuminated wheat leaves.     

Enhanced Dark CO(2) Fixation by Preilluminated Chlorella pyrenoidosa and Anacystis nidulans

The products of short time photosynthesis and of enhanced dark ¹⁴CO₂ fixation (illumination in helium prior to addition of ¹⁴CO₂ in dark) by Chlorella pyrenoidosa and Anacystis nidulans were compared. Glycerate 3-phosphate, phosphoenolpyruvate, alanine, and aspartate accounted for the bulk of the ¹⁴C assimilated during enhanced dark fixation while hexose and pentose phosphates accounted for the largest fraction of isotope assimilated during photosynthesis. During the enhanced dark fixation period, glycerate 3-phosphate is carboxyl labeled and glucose 6-phosphate is predominantly labeled in carbon atom 4 with lesser amounts in the upper half of the C₆ chain and traces in carbon atoms 5 and 6. Tracer spread throughout all the carbon atoms of photosynthetically synthesized glycerate 3-phosphate and glucose 6-phosphate. During the enhanced dark fixation period, there was a slow formation of sugar phosphates which subsequently continued at 5 times the initial rate long after the cessation of ¹⁴CO₂ uptake. To explain the kinetics of changes in the labelling patterns and in the limited formation of the sugar phosphates during enhanced dark CO₂ fixation, the suggestion is made that most of the reductant mediating these effects did not have its origin in the preillumination phase.

It is concluded that a complete photosynthetic carbon reduction cycle operates to a limited extent, if at all, in the dark period subsequent to preillumination.

     

Ecosystem carbon exchange in two terrestrial ecosystem mesocosms under changing atmospheric CO2 concentrations

The ecosystem-level carbon uptake and respiration were measured under different CO₂ concentrations in the tropical rainforest and the coastal desert of Biosphere 2, a large enclosed facility. When the mesocosms were sealed and subjected to step-wise changes in atmospheric CO₂ between daily means of 450 and 900 μmol mol⁻¹, net ecosystem exchange (NEE) of CO₂ was derived using the diurnal changes in atmospheric CO₂ concentrations. The step-wise CO₂ treatment was effectively replicated as indicated by the high repeatability of NEE measurements under similar CO₂ concentrations over a 12-week period. In the rainforest mesocosm, daily NEE was increased significantly by the high CO₂ treatments because of much higher enhancement of canopy CO₂ assimilation relative to the increase in the nighttime ecosystem respiration under high CO₂. Furthermore, the response of daytime NEE to increasing atmospheric CO₂ in this mesocosm was not linear, with a saturation concentration of 750 μmol mol⁻¹. In the desert mesocosm, a combination of a reduction in ecosystem respiration and a small increase in canopy CO₂ assimilation in the high CO₂ treatments also enhanced daily NEE. Although soil respiration was not affected by the short-term change in atmospheric CO₂ in either mesocosm, plant dark respiration was increased significantly by the high CO₂ treatments in the rainforest mesocosm while the opposite was found in the desert mesocosm. The high CO₂ treatments increased the ecosystem light compensation points in both mesocosms. High CO₂ significantly increased ecosystem radiation use efficiency in the rainforest mesocosm, but had a much smaller effect in the desert mesocosm. The desert mesocosm showed much lower absolute response in NEE to atmospheric CO₂ than the rainforest mesocosm, probably because of the presence of C₄ plants. This study illustrates the importance of large-scale experimental research in the study of complex global change issues. Received: 30 October 1998 / Accepted: 2 December 1998