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1.
Red light exposures given to dark-grown wheat seedlings (Triticum aestivum L.) prior to etioplast isolation reduced the ability of these organelles to consume O2. The same preharvest red light exposures also decreased protochlorophyll(ide) content of etioplasts. In addition, regeneration of both O2 uptake rates as well as protochlorophyll(ide) levels followed a parallel time course. These similarities suggested that photoconversion of protochlorophyll(ide)-650 to chlorophyll(ide) may mediate some process with O2 as the electron acceptor. This process appears to involve photooxidation of nonphotoconvertible protochlorophyll(ide) as well as of newly formed chlorophyll(ide). This hypothesis is further supported by the observations that: (a) the in vitro light induced O2 uptake phenomenon was observed in solubilized protochlorophyll(ide) holochrome preparations; and (b) photoinduced O2 uptake was reduced to zero rate by light exposure time equivalent to that required for chlorophyll(ide) and nonphotoconvertible protochlorophyll(ide) destruction.  相似文献   

2.
Oxygen exchange in leaves in the light   总被引:30,自引:20,他引:10       下载免费PDF全文
Photosynthetic O2 production and photorespiratory O2 uptake were measured using isotopic techniques, in the C3 species Hirschfeldia incana Lowe., Helianthus annuus L., and Phaseolus vulgaris L. At high CO2 and normal O2, O2 production increased linearly with light intensity. At low O2 or low CO2, O2 production was suppressed, indicating that increased concentrations of both O2 and CO2 can stimulate O2 production. At the CO2 compensation point, O2 uptake equaled O2 production over a wide range of O2 concentrations. O2 uptake increased with light intensity and O2 concentration. At low light intensities, O2 uptake was suppressed by increased CO2 concentrations so that O2 uptake at 1,000 microliters per liter CO2 was 28 to 35% of the uptake at the CO2 compensation point. At high light intensities, O2 uptake was stimulated by low concentrations of CO2 and suppressed by higher concentrations of CO2. O2 uptake at high light intensity and 1000 microliters per liter CO2 was 75% or more of the rate of O2 uptake at the compensation point. The response of O2 uptake to light intensity extrapolated to zero in darkness, suggesting that O2 uptake via dark respiration may be suppressed in the light. The response of O2 uptake to O2 concentration saturated at about 30% O2 in high light and at a lower O2 concentration in low light. O2 uptake was also observed with the C4 plant Amaranthus edulis; the rate of uptake at the CO2 compensation point was 20% of that observed at the same light intensity with the C3 species, and this rate was not influenced by the CO2 concentration. The results are discussed and interpreted in terms of the ribulose-1,5-bisphosphate oxygenase reaction, the associated metabolism of the photorespiratory pathway, and direct photosynthetic reduction of O2.  相似文献   

3.
The role of the oxyhydrogen reaction in the nitrogen metabolism of Anabaena cylin-drica, particularly under conditions of dinitrogen starvation, was investigated. It was shown that although this reaction supports nitrogenase activity in the dark, when the cells are deprived of nitrogen the rate of hydrogen uptake is little changed. Measurements of ammonia excretion into the medium in the presence of methionine sulfoximine under such conditions indicated that hydrogen uptake supported the turnover of cell protein as an alternative source of nitrogen. In the absence of H2 and O2 in the dark, nitrogenase activity was negligible but protein turnover continued. In their presence nitrogenase activity was greatly stimulated; turnover was also stimulated but to a greater extent in the absence of nitrogenase substrates. The oxyhydrogen reaction also stimulated uptake of ammonium ions by intact filaments in argon in the dark. Only at very low hydrogen tensions can net hydrogen formation be obtained in argon/CO2 in the light, casting considerable doubt on the suitability of hydrogenase-containing organisms for biophotolytic hydrogen formation. Addition of exogenous ammonia to the cultures incubated in argon resulted in a pronounced stimulation of H2 uptake; nitrate and its derivatives had no such effect, nor did various amino acid derivatives of ammonia.  相似文献   

4.
The Kok Effect in Chlamydomonas reinhardi   总被引:2,自引:1,他引:1       下载免费PDF全文
A Haxo-Blinks rate-measuring oxygen electrode together with a modulated light source gave an average current signal (change in net O2 exchange) and a modulated current signal (photosynthetic O2 evolution). Using this apparatus, net O2 exchange and photosynthetic O2 evolution at low intensities have been studied in the green alga, Chlamydomonas reinhardi. At both 645 nm and 695 nm, the curves of net O2 exchange as a function of light intensity were steeper at lowest intensities than about compensation, indicative of the Kok effect. The effect was greater at 695 nm than at 645 nm. The corresponding curves of photosynthetic O2 evolution, on the other hand, showed no Kok effect; here, the slope was lowest at lowest intensity. The absence of the Kok effect in O2 evolution, together with its sensitivity to monofluoroacetic acid, show that it is due to an interaction of photosynthesis and respiration. The effect was exaggerated by 3-(3,4-dichlorophenyl)-1,1-dimethylurea. In the presence of concentrations of this inhibitor sufficient to inhibit O2 evolution completely, a light-induced change in net O2 exchange remained. This was interpreted as a system I dependent depression of respiratory O2 uptake. The Kok effect remained undiminished in concentrations of carbonyl cyanide m-chlorophenylhydrazone and 2,4-dinitrophenol which partially uncoupled either oxidative phosphorylation alone or both oxidative and photosynthetic phosphorylations. The above results can be explained within a model of the Kok effect in which O2 uptake is depressed by diversion of reductant away from respiratory electron transport and into photosystem I. The same photodepression of O2 uptake also appears to account for a transient in net O2 exchange seen in several algae upon turning off the light.  相似文献   

5.
The response of net O2 exchange to light intensity by intact Anacystis nidulans cells in the presence of saturating NaHCO3 concentrations followed a curve with an inflection near the light-compensation point. Addition of either KNO3 or NH4Cl stimulated O2 uptake in the dark and at light intensities below the light-compensation point. This resulted in steeper slopes of the curve calculated below and above the light-compensation point. At O2 concentrations limiting dark respiration, addition of inorganic nitrogen had no effect on either dark respiration or O2 exchange in the light. The apparent changes in photosynthetic yield observed under normal O2 concentration disappeared when respiration was limited by O2 availability, indicating that the effects of inorganic nitrogen on O2 exchange at low light intensities are due to stimulation of respiration rather than to increases in photosynthetic yield.  相似文献   

6.
Technical features and examples of application of a special emitter–detector module for highly sensitive measurements of the electrochromic pigment absorbance shift (ECS) via dual-wavelength (550–520 nm) transmittance changes (P515) are described. This device, which has been introduced as an accessory of the standard, commercially available Dual-PAM-100 measuring system, not only allows steady-state assessment of the proton motive force (pmf) and its partitioning into ΔpH and ΔΨ components, but also continuous recording of the overall charge flux driven by photosynthetic light reactions. The new approach employs a double-modulation technique to derive a continuous signal from the light/dark modulation amplitude of the P515 signal. This new, continuously measured signal primarily reflects the rate of proton efflux via the ATP synthase, which under quasi-stationary conditions corresponds to the overall rate of proton influx driven by coupled electron transport. Simultaneous measurements of charge flux and CO2 uptake as a function of light intensity indicated a close to linear relationship in the light-limited range. A linear relationship between these two signals was also found for different internal CO2 concentrations, except for very low CO2, where the rate of charge flux distinctly exceeded the rate of CO2 uptake. Parallel oscillations in CO2 uptake and charge flux were induced by high CO2 and O2. The new device may contribute to the elucidation of complex regulatory mechanisms in intact leaves.  相似文献   

7.
A mass spectrometric method combining 16O/18O and 12C/13C isotopes was used to quantify the unidirectional fluxes of O2 and CO2 during a dark to light transition for guard cell protoplasts and mesophyll cell protoplasts of Commelina communis L. In darkness, O2 uptake and CO2 evolution were similar on a protein basis. Under light, guard cell protoplasts evolved O2 (61 micromoles of O2 per milligram of chlorophyll per hour) almost at the same rate as mesophyll cell protoplasts (73 micromoles of O2 per milligram of chlorophyll per hour). However, carbon assimilation was totally different. In contrast with mesophyll cell protoplasts, guard cell protoplasts were able to fix CO2 in darkness at a rate of 27 micromoles of CO2 per milligram of chlorophyll per hour, which was increased by 50% in light. At the onset of light, a delay observed for guard cell protoplasts between O2 evolution and CO2 fixation and a time lag before the rate of saturation suggested a carbon metabolism based on phosphoenolpyruvate carboxylase activity. Under light, CO2 evolution by guard cell protoplasts was sharply decreased (37%), while O2 uptake was slowly inhibited (14%). A control of mitochondrial activity by guard cell chloroplasts under light via redox equivalents and ATP transfer in the cytosol is discussed. From this study on protoplasts, we conclude that the energy produced at the chloroplast level under light is not totally used for CO2 assimilation and may be dissipated for other purposes such as ion uptake.  相似文献   

8.
The nature of the different processes of O2 uptake involved in the light in the red macroalga Chondrus crispus Stackhouse (Rhodophyta, Gigartinales) was investigated. At limiting CO2, INH (2.5 mM) did not alter the O2 uptake rate. Glycolate was not excreted and did not accumulate within the cells. KCN reduced the rate of O2 uptake in the light by 76% at limiting CO2 and by 43% at saturating CO2, but caused > 95% inhibition of O2 evolution. DCMU (5 μM) totally blocked the photosynthetic electron transport chain, but allowed a residual O2 uptake of 3.0±0.6 μmol O2 .h?1.g?1 FW, irrespective of the CO2 concentration. In saturating CO2, a high light intensity pretreatment significantly stimulated the rate of O2 uptake compared to net O2 evolution, suggesting the persistence, in the light, of mitochondrial respiration. Irrespective of the CO2 concentration, the optimum temperature for O2 evolution was 17°C whereas dark O2 uptake increased linearly with temperature. In contrast, O2 uptake in the light showed an optimum at 17°C in limiting CO2, and 21–25° C in saturating CO2; its Q10 was 2.4 at limiting CO2, a value close to that of RuBP oxygenase, and 3.1 at saturating CO2, a value close to that of dark respiration. It is concluded that: 1) mitochondrial respiration and Mehler reaction are both involved at all CO2 concentrations, 2) RuBP oxygenase activity cannot account for more than 45%, and Mehler reaction for less than 20%, of the total O2 uptake observed in the light at limiting CO2.  相似文献   

9.
The mass transfer rate of 14C-sucrose translocation from sugar beet (Beta vulgaris, L.) leaves was measured over a range of net photosynthesis rates from 0 to 60 milligrams of CO2 decimeters−2 hour−1 under varying conditions of light intensity, CO2 concentration, and O2 concentration. The resulting rate of translocation of labeled photosynthate into total sink tissue was a linear function (slope = 0.18) of the net photosynthesis rate of the source leaf regardless of light intensity (2000, 3700, or 7200 foot-candles), O2 concentration (21% or 1% O2), or CO2 concentration (900 microliters/liter of CO2 to compensation concentration). These data support the theory that the mass transfer rate of translocation under conditions of sufficient sink demand is limited by the net photosynthesis rate or more specifically by sucrose synthesis and this limitation is independent of light intensity per se. The rate of translocation was not saturated even at net photosynthesis rates four times greater than the rate occurring at 300 microliters/liter of CO2, 21% O2, and saturating light intensity.  相似文献   

10.
Woo KC 《Plant physiology》1983,72(2):313-320
This study examines the effect of antimycin A and nitrite on 14CO2 fixation in intact chloroplasts isolated from spinach (Spinacia oleracea L.) leaves. Antimycin A (2 micromolar) strongly inhibited CO2 fixation but did not appear to inhibit or uncouple linear electron transport in intact chloroplasts. The addition of small quantities (40-100 micromolar) of nitrite or oxaloacetate, but not NH4Cl, in the presence of antimycin A restored photosynthesis. Antimycin A inhibition, and the subsequent restoration of photosynthetic activities by nitrite or oxaloacetate, was observed over a wide range of CO2 concentration, light intensity, and temperature. High O2 concentration (up to 240 micromolar) did not appear to influence the extent of the inhibition by antimycin A, nor the subsequent restoration of photosynthetic activity by nitrite or oxaloacetate. Studies of O2 exchanges during photosynthesis in cells and chloroplasts indicated that 2 micromolar antimycin A stimulated O2 uptake by about 25% while net O2 evolution was inhibited by 76%. O2 uptake in chloroplasts in the presence of 2 micromolar antimycin A was 67% of total O2 evolution. These results suggest that only a small proportion of the O2 uptake measured was directly linked to ATP generation. The above evidence indicates that cyclic photophosphorylation is the predominant energy-balancing reaction during photosynthesis in intact chloroplasts. On the other hand, pseudocyclic O2 uptake appears to play only a minimal role.  相似文献   

11.
Pisum sativum L. cv Alsweet (garden pea) and Lycopersicon esculentum flacca Mill. (tomato) were used to evaluate the phytotoxicity of SO2 and O3 in the light and dark. Plants were grown in controlled environment chambers and exposed to SO2 or O3 in the light or dark at the same environmental conditions at which they were grown. The pea plants were treated with fusicoccin to ensure open stomata in the dark; the stomata of the tomato mutant remained open in the dark. Both species exhibited 64% to 80% less foliar necrosis following exposure to SO2 (0.5 to 1.0 microliter per liter for 2 hours) in the light than in the dark. The decrease in SO2 injury for light versus dark exposed plants was greater in fully expanded than expanding leaves. Both species exhibited 30% greater foliar necrosis following exposure to O3 (0.2 microliter per liter for 2 hours) in the light than dark. The increase in O3 injury in the light versus dark was similar for leaves at all stages of expansion. Leaf conductance to water vapor was 7% to 11% and 23% higher in the light than dark for fusicoccin-treated peas and tomato plants, respectively, indicating greater foliar uptake of both pollutants in the light than dark. Thus, the decreased SO2 toxicity in the light was not associated with pollutant uptake, but rather the metabolism of SO2. In contrast, the increased toxicity of O3 in the light was at least in part associated with increased uptake or could not be separated from it.  相似文献   

12.
The substrate-dependent O2 uptake by sycamore (Acer pseudoplatanus L.) cell mitochondria in the presence of ADP and limiting Pi concentrations has been measured. The Pi concentration for half-maximum O2 uptake rate was found to be in the range 20 to 50 micromolar for all the substrates tested. 31P NMR of intact sycamore cells indicated that the Pi concentration in the cytoplasm was in the range 5 to 6 millimolar, approximately 100-fold higher than the Pi concentration required for maximum O2 uptake rates by isolated mitochondria. When sycamore cells were transferred to a culture medium devoid of Pi, the cytoplasmic Pi concentration decreased from 6 to less than 3 millimolar, but the intact cell respiration remained practically constant for at least 4 days. These results strongly suggest that, in vivo, the respiration rate of sycamore cells is not limited by the quantity of Pi supplied to the mitochondria.  相似文献   

13.
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 Km (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.  相似文献   

14.
Photosynthetic efficiency, primary productivity, and N2 reduction were determined in peas (Pisum sativum L. var. Alaska) grown at light intensities ranging from severely limiting to saturating. Plants grown under higher light intensities showed greater carboxylation and light capture potential and higher rates of net C exchange. Uptake of N2, computed from measured C2H2 reduction and H2 evolution rates, also increased with growth light intensity, while the previously proposed relative efficiency of N2 fixation, based on these same parameters, declined. The plot of N/C ratios (total nitrogen content/plant dry weight) increased hyperbolically with light intensity, and the plot of N2/CO2 uptake ratios (N2 uptake rate/net CO2 uptake rate) increased linearly. Both plots extrapolated to the light compensation point. The data indicate that the relative efficiency of N2 fixation is not necessarily correlated with maximum plant productivity and that evaluation of a plant's capacity to reduce N2 is related directly to concurrent CO2 reduction. A measure of whole plant N2 fixation efficiency based on the N2/CO2 uptake ratio is proposed.  相似文献   

15.
The nature of the process responsible for the stationary O2 uptake occurring in the light under saturating CO2 concentration in Chlamydomonas reinhardii has been investigated. For this purpose, a mass spectrometer with a membrane inlet system was used to measure O2 uptake and evolution in the algal suspension. First, we observed that the O2 uptake rate was constant (about 0.5 micromoles of O2 per milligram chlorophyll per minute) during a light to dark transition and was not affected by 3-(3,4-dichlorophenyl)-1,1-dimethylurea. Salicylhydroxamic acid had no effect on O2 uptake in the dark or in the light, but was found to have the same inhibitory effect either in the dark or in the light when added to cyanide-treated algae. The stimulation of the O2 uptake rate due to the uncoupling effect of carbonyl cyanide m-chlorophenylhydrazone was about the same in the dark or in the light. From these results, we conclude that mitochondrial respiration is maintained during illumination and therefore is not inhibited by high ATP levels. Another conclusion is that in conditions where photorespiration is absent, no other light-dependent O2 uptake process occurs. If Mehler reactions are involved, in Chlamydomonas, under conditions where both photosynthetic carbon oxidation and reduction cycles cannot operate (as in cyanide-treated algae), their occurrence in photosynthesizing algae either under saturating CO2 concentration or at the CO2 compensation point appears very unlikely. The comparison with the situation previously reported in Scenedesmus (R. J. Radmer and B. Kok 1976 Plant Physiol 58: 336-340) suggests that different O2 uptake processes might be present in these two algal species.  相似文献   

16.
The reversible hydrogenase from Anabaena 7120 appeared when O2 was continuously removed from a growing culture. Activity increased further when cells were incubated under argon in the dark or in the light plus 3-(3,4-dichlorophenyl)-1,1-dimethylurea. Hydrogenase existed in an inactive state during periods of O2 evolution. It could be reductively activated by exposure to reduced methyl viologen or by dark, anaerobic incubation. Hydrogenase-containing cells evolved H2 slowly during dark anaerobic incubations, and the rate of H2 evolution was increased by illumination with low intensity light. Light enhancement of H2 evolution was of short duration and was eliminated by the ferredoxin antagonist disalicylidene diaminopropane. Physiological acceptors that supported H2 uptake included NO3, NO2, and HSO3, and light had a slight influence on the rate of H2 uptake with these acceptors. Low levels of O2 supported H2 uptake, but higher concentrations of O2 inactivated the hydrogenase. Hydrogen uptake with HCO3 as acceptor was the most rapid reaction measured, and it was strictly light-dependent. It occurred only at low light intensities, and higher light intensities restored normal O2-evolving photosynthesis. It is suggested that hydrogenase is present to capture exogenous H2 as a source of reducing equivalents during growth in anaerobic environments.  相似文献   

17.
Photosynthesis and photorespiration in whole plants of wheat   总被引:12,自引:11,他引:1       下载免费PDF全文
Wheat was cultivated in a small phytotronic chamber. 18O2 was used to measure the O2 uptake by the plant, which was recorded simultaneously with the O2 evolution, net CO2 uptake, and transpiration. At normal atmospheric CO2 concentration, photorespiration, measured as O2 uptake, was as important as the net photosynthesis. The level of true O2 evolution was independent of CO2 concentration and stayed nearly equal to the sum of net CO2 photosynthesis and O2 uptake. We conclude that at a given light intensity, O2 and CO2 compete for the reducing power produced at constant rate by the light reactions of photosynthesis.  相似文献   

18.
A closed system consisting of an assimilation chamber furnished with a membrane inlet from the liquid phase connected to a mass spectrometer was used to measure O2 evolution and uptake by Chlamydomonas reinhardtii cells grown in ambient (0.034% CO2) or CO2-enriched (5% CO2) air. At pH = 6.9, 28°C and concentrations of dissolved inorganic carbon (DIC) saturating for photosynthesis, O2 uptake in the light (Uo) equaled O2 production (Eo) at the light compensation point (15 micromoles photons per square meter per second). Eo and Uo increased with increasing photon fluence rate (PFR) but were not rate saturated at 600 micromoles photons per square meter per second, while net O2 exchange reached a saturation level near 500 micromoles photons per square meter per second which was nearly the same for both, CO2-grown and air-grown cells. Comparison of the Uo/Eo ratios between air-grown and CO2-grown C. reinhardtii showed higher values for air-grown cells at light intensities higher than light compensation. For both, air-grown and CO2-grown algae the rates of mitochondrial O2 uptake in the dark measured immediately before and 5 minutes after illumination were much lower than Uo at PFR saturating for net photosynthesis. We conclude that noncyclic electron flow from water to NADP+ and pseudocyclic electron flow via photosystem I to O2 both significantly contribute to O2 exchange in the light. In contrast, mitochondrial respiration and photosynthetic carbon oxidation cycle are regarded as minor O2 consuming reactions in the light in both, air-grown and CO2-grown cells. It is suggested that the “extra” O2 uptake by air-grown algae provides ATP required for the energy dependent CO2/HCO3 concentrating mechanism known to be present in these cells.  相似文献   

19.
A mass spectrometric 16O2/18O2-isotope technique was used to analyse the rates of gross O2 evolution, net O2 evolution and gross O2 uptake in relation to photon fluence rate by Dunaliella tertiolecta adapted to 0.5, 1.0, 1.5, 2.0 and 2.5 M NaCl at 25°C and pH 7.0.At concentrations of dissolved inorganic carbon saturating for photosynthesis (200 M) gross O2 evolution and net O2 evolution increased with increasing salinity as well as with photon fluence rate. Light compensation was also enhanced with increased salinities. Light saturation of net O2 evolution was reached at about 1000 mol m-2s-1 for all salt concentrations tested. Gross O2 uptake in the light was increased in relation to the NaCl concentration but it was decreased with increasing photon fluence rate for almost all salinities, although an enhanced flow of light generated electrons was simultaneously observed. In addition, a comparison between gross O2 uptake at 1000 mol photons m-2s-1, dark respiration before illumination and immediately after darkening of each experiment showed that gross O2 uptake in the light paralleled but was lower than mitochondrial O2 consumption in the dark.From these results it is suggested that O2 uptake by Dunaliella tertiolecta in the light is mainly influenced by mitochondrial O2 uptake. Therefore, it appears that the light dependent inhibition of gross O2 uptake is caused by a reduction in mitochondrial O2 consumption by light.Abbreviations DCMU 3-(3, 4-dichlorophenyl)-1, 1-dimethylurea - DHAP dihydroxy-acetonephosphate - DIC dissolved inorganic carbon - DRa rate of dark respiration immediately after illumination - DRb rate of dark respiration before illumination - E0 rate of gross oxygen evolution in the light - NET rate of net oxygen evolution in the light - PFR photon fluence rate - RubP rubulose-1,5-bisphosphate - SHAM salicyl hydroxamic acid - U0 rate of gross oxygen uptake in the light  相似文献   

20.
Lara C  Romero JM 《Plant physiology》1986,81(2):686-688
The effect of light intensity on the rates of ammonium and nitrate uptake and of CO2 fixation has been determined in intact Anacystis nidulans cells. Ammonium uptake became saturated at photon flux values of about 60 microeinsteins per square meter per second, whereas both nitrate uptake and CO2 fixation reached saturation at about 250 microeinsteins per square meter per second, the rates of the two latter processes being tightly correlated at any light intensity assayed. Inhibition of ammonium assimilation resulted in the loss of correlation between CO2 fixation and nitrate uptake, the latter process exhibiting then a reduced light requirement. The results establish a clear distinction between ammonium utilization and nitrate utilization with regard to their light requirement and to the nature of their dependence upon CO2 fixation.  相似文献   

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