Nutrient-enhanced survival of and phenanthrene mineralization by alginate-encapsulated and freePseudomonas sp. UG14Lr cells in creosote-contaminated soil slurries

The effects of nutrient amendment and alginate encapsulation on survival of and phenanthrene mineralization by the bioluminescentPseudomonas sp. UG14Lr in creosote-contaminated soil slurries were examined. UG14Lr was inoculated into creosote-contaminated soil slurries either as a free cell suspension or encapsulated in alginate beads prepared with montmorillonite clay and skim milk. Additional treatments were free-cell-inoculated slurries amended with sterile alginate beads, free-cell-inoculated and uninoculated slurries amended with skim milk only, and uninoculated, unamended slurries. Mineralization was determined by measuring¹⁴CO₂ released from radiolabelled phenanthrene. Survival was measured by selective plating and bioluminescence. Inclusion of skim milk was found to enhance both survival of and phenanthrene mineralization by free and encapsulated UG14Lr cells.     

Photosynthetic characteristics of coccoid marine cyanobacteria

Two strains of marine Synechococcus possessed a much greater potential for photorespiration than other marine algae we have studied. This conclusion was based on the following physiological and biochemical characteristics: a) a light-dependent O₂ inhibition of photosynthetic CO₂ assimilation at atmospheric O₂ concentrations. The degree of inhibition was dependent on the relative concentrations of dissolved O₂ and CO₂, being greatest at 100% O₂ with no extra bicarbonate added to the medium; b) actively photosynthesizing cells had high levels of ribulose-1,5-bisphosphate carboxylase compared with phosphoenolpyruvate carboxylase; ribulose-1,5-bisphosphate oxygenase activities were three times greater than ribulose-1,5-bisphosphate carboxylase activities; c) cells photosynthesizing in 21% O₂, showed significant ¹⁴C-labelling of phosphoglycolate and glycolate and the percentage of total carbon-14 incorporated into these two compounds increased when the O₂ concentration was 100%; d) at 100% O₂, there was a post-illumination enhanced rate of O₂ consumption, which was three times greater than dark respiration, and the rate declined with increasing bicarbonate concentrations. The inhibitory effect of O₂ on photosynthesis did not appear to be solely due to photorespiration, since O₂ inhibition of photosynthetic O₂ evolution was much greater than that of photosynthetic CO₂ assimilation. Also, O₂ inhibition of photosynthetic O₂ evolution declined only slightly with decreasing light intensities, while the inhibition of CO₂ assimilation declined rapidly with decreasing light intensity.     

Microbial Dynamics Associated with Multiphasic Decomposition of 14C-Labeled Cellulose in Soil

Abstract Recent emphasis on residue management in sustainable agriculture highlights the importance of elucidating the mechanisms of microbial degradation of cellulose. Cellulose decomposition and its associated microbial dynamics in soil were investigated in incubation experiments. Population dynamics of actinomycetes, bacteria, and fungi were monitored by direct counts. Populations of oligotrophic bacteria in cellulose-amended soil were determined by plate count using a low C medium containing 4 mg C liter⁻¹ agar, and copiotrophs using a high C medium. Cumulative ¹⁴CO₂ evolution from ¹⁴C-labeled cellulose was best described by a multiphasic curve in a 28-day incubation experiment. The initial phase of decomposition was attributed mainly to the activity of bacterial populations with a low oligotroph-to-copiotroph ratio, and the second phase mainly to fungal populations. An increase in oligotroph-to-copiotroph ratio coincided with the emergence of a rapid ¹⁴CO₂ evolution stage. Streptomycin reduced ¹⁴CO₂ evolution during the first phase and prompted earlier emergence of the second phase, compared to the control. Cycloheximide initially promoted ¹⁴CO₂ evolution but subsequently had a lasting negative effect on ¹⁴CO₂ evolution. Cycloheximide addition significantly increased bacterial biomass and resulted in substantially stronger oscillation of active bacterial populations, whereas it initially reduced, and then stimulated, active fungal biomass. The observed changes in ¹⁴CO₂ evolution could not be explained by observed shifts in fungal and bacterial biomass, probably because functional groups of fungi and bacteria could not be distinguished. However, it was suggested that oligotrophic bacteria prompted activation of cellulolytic enzumes in fungi and played an important role in leading to fungal dominance of cellulose decomposition. Received: 2 October 1995; Accepted: 10 February 1996     

Photosynthesis in enzymatically isolated leaf cells from the CAM plant Sedum telephium L.

A technique has been developed for the enzymatic isolation of leaf cells from the Crassulacean acid-metabolism plant Sedum telephium. The cells exhibited high activity in both ¹⁴CO₂ incorporation (30–70 mol CO₂ mg^-1 chlorophyll h^-1) and O₂ evolution in the presence of bicarbonate (60–110 mol O₂ mg^-1 chlorophyll h^-1). Half-maximum saturation of ¹⁴CO₂ incorporation occurred at a bicarbonate concentration of ca. 2 mM (20 M CO₂) at pH 8.4 and 30°C. Two types of light-dependent O₂ evolution are reported: O₂ evolution in the absence of exogenously supplied bicarbonate (endogenous O₂ evolution), and bicarbonate-stimulated O₂ evolution. Oxygen evolution in the presence of approximately ambient concentrations of CO₂ appeared to be a combination of the endogenous O₂ evolution and O₂ evolution from fixation of the exogenously supplied CO₂.Abbreviations CAM Crassulacean acid metabolism - cirlo chlorophyll - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - PEP phosphoenolpyruvate - RuDP ribulose-1,5-diphosphate     

Metabolic Activity and Population Dynamics of Rhizobia Introduced into Unamended and Bentonite-Amended Loamy Sand

Respiration measurements showed that the cumulative amount of CO₂ respired by rhizobia introduced into sterile bentonite-amended loamy sand was significantly higher than it was in unamended loamy sand. The maintenance respiration of rhizobial cells was not influenced by the presence of bentonite clay. Carbon was used more efficiently during growth in bentonite-amended than in unamended loamy sand. The presence of bentonite clay increased the growth rate of rhizobia introduced into sterile soil. Survival studies performed in nonsterile bentonite-amended loamy sand showed that the use of high (10¹⁰ cells per g of dry soil) rather than lower (10⁴ to 10⁷ cells per g of dry soil) inoculum densities increased the final survival levels of introduced rhizobia. In unamended loamy sand, the application of 10¹⁰ or 10⁷ cells per g of dry soil resulted in similar final survival levels. Pore shape and the continuity of the water-filled pore system were suggested to largely determine the colonization rate of protective microhabitats.     

Photorespiration in Air and High CO(2)-Grown Chlorella pyrenoidosa

Oxygen inhibition of photosynthesis and CO₂ evolution during photorespiration were compared in high CO₂-grown and air-grown Chlorella pyrenoidosa, using the artificial leaf technique at pH 5.0. High CO₂ cells, in contrast to air-grown cells, exhibited a marked inhibition of photosynthesis by O₂, which appeared to be competitive and similar in magnitude to that in higher C₃ plants. With increasing time after transfer to air, the photosynthetic rate in high CO₂ cells increased while the O₂ effect declined. Photorespiration, measured as the difference between ¹⁴CO₂ and ¹²CO₂ uptake, was much greater and sensitive to O₂ in high CO₂ cells. Some CO₂ evolution was also present in air-grown algae; however, it did not appear to be sensitive to O₂. True photosynthesis was not affected by O₂ in either case. The data indicate that the difference between high CO₂ and air-grown algae could be attributed to the magnitude of CO₂ evolution. This conclusion is discussed with reference to the oxygenase reaction and the control of photorespiration in algae.     

Microbial Activity and 13C/12C Ratio as Evidence of N-Hexadecane and N-Hexadecanoic Acid Biodegradation in Agricultural and Forest Soils

The dynamics of microbial degradation of exogenous contaminants, n-hexadecane and its primary microbial oxidized metabolite, n-hexadecanoic (palmitic) acid, was studied for topsoils, under agricultural management and beech forest on the basis the changes in O₂ uptake, CO₂ evolution and its associated carbon isotopic signature, the respiratory quotient (RQ) and the priming effect (PE) of substrates. Soil microbial communities in agricultural soil responded to the n-hexadecane addition more rapidly compared to those of forest soil, with lag-periods of about 23 ± 10 and 68 ± 13 hours, respectively. Insignificant difference in the lag-period duration was detected for agricultural (t_lag = 30 ± 13 h) and forest (t_lag = 30 ± 14 h) soils treated with n-hexadecanoic (palmitic) acid. These results demonstrate that the soil microbiota has different metabolic activities for using n-hexadecane as a reductive hydrocarbon and n-hexadecanoic acid as a partly oxidized hydrocarbon. The corresponding δ¹³C of respired CO₂ after the addition of the hydrocarbon contaminants to soils indicates a shift in microbial activity towards the consumption of exogenous substrates with a more complete degradation of n-hexadecane in the agricultural soil, for which some initial contents of hydrocarbons are inherent. It is supposed that the observed deviation of RQ from theoretically calculated value under microbial substrate mineralization is determined by difference in the time (Δt_i) of registration of CO₂ production and O₂ consumption. Positive priming effect (PE) of n-hexadecane and negative PE of n-hexadecanoic (palmitic) acid were detected in agricultural and forest soils. It is suggested that positive PE of n-hexadecane is conditioned by the induction of microbial enzymes that perform hydroxylation/oxygenation of stable SOM compounds mineralized by soil microbiota to CO₂. The microbial metabolism coupled with oxidative decarboxylation of n-hexadecanoic acid is considered as one of the most probable causes of the revealed negative PE value.     

Effects of Various Concentrations of Carbon Dioxide on Respiration and Potassium Uptake in Barley Roots

Barley roots contain a CO₂ sensitive respiratory fraction which is inhibited in 50 per cent CO₂ and is partially restored upon subsequent exposure to air. The residual O₂ consumption occurring at CO₂ concentrations between 50 per cent and 95 per cent amounts to about 40 per cent of the O₂ uptake in air and can support K⁺ uptake for a limited time at a rate equal to or higher than occurs in air. Above 95 per cent CO₂ both O₂ and K⁺ uptakes decrease rapidly. 2,4-dinitrophenol (DNP), in the range of 10^?6 to 10^?5M, stimulates O₂ uptake by the roots in air. The stimulation is absent when roots are treated with DNP in 80 per cent CO₂, presumably because of the reduced demand for inorganic phosphate and phosphate acceptor at the lower respiratory level in high CO₂. In either air or CO₂, K⁺ uptake is strongly inhibited by DNP. A comparison of the respiratory and K⁺ uptake data indicates that O₂ consumption is a necessary requirement for the uptake process in high CO₂. Protoplasmic streaming in the root cells is rapidly stopped by high CO₂ although K⁺ uptake and O₂ consumption continue. The cation uptake mechanism in high CO₂ appears to be limited to the stationary cytoplasm. It is also possible that a similar mechanism may be involved in cation uptake in air.     

Trophic interactions in soils as they affect energy and nutrient dynamics. IV. Flows of metabolic and biomass carbon

Flows of biomass and respiratory carbon were studied in a series of propylene-oxide sterilized soil microcosms. One-half of the microcosms received three pulsed additions of 200 ppm glucose-carbon to mimic rhizosphere carbon inputs. Biotic variables were: bacteria (Pseudomonas) alone, or amoebae (Acanthamoeba) and nematodes (Mesodiplogaster) singly, or both combined in the presence of bacteria.Over the 24-day experiment, respiration was significantly higher in the microcosms containing the bacterial grazers. Biomass accumulation by amoebae was significantly higher than that by nematodes. The nematodes respired up to 30-fold more CO₂ per unit biomass than did amoebae. Similar amounts of carbon flowed into both respiratory and biomass carbon in microcosms with fauna, compared with the bacteria-alone microcosms. However, partitioning of available carbon by the microfauna varied considerably, with little biomass production and relatively more CO₂-C produced in the nematode-containing microcosms. The amoebae, in contrast, allocated more carbon to tissue production (about 40% assimilation efficiency) and correspondingly less to CO₂.     

Ferric reduction by iron-limited Chlamydomonas cells interacts with both photosynthesis and respiration

Iron limitation led to a large increase in extracellular ferricyanide (Fe[III]) reductase activity in cells of the green alga Chlamydomonas reinhardtii Dangeard. Mass-spectrometric measurement of gas exchange indicated that ferricyanide reduction in the dark resulted in a stimulation of respiratory CO₂ production without affecting the rate of respiratory O₂ consumption, consistent with the previously postulated activation of the oxidative pentose phosphate pathway in support of Fe(III) reduction by iron-limited Chlamydomonas cells (X. Xue et al., 1998, J. Phycol. 34: 939–944). At saturating irradiance, the rate of ferricyanide reduction was stimulated almost 3-fold, and this stimulation was inhibited by 3-(3′,4′-dichlorophenyl)-1,1-dimethylurea. Ferricyanide reduction during photosynthesis resulted in approximately a 50% inhibition of photosynthetic CO₂ fixation at saturating irradiance, and almost 100% inhibition of CO₂ fixation at sub-saturating irradiance. Photosynthesis by iron-sufficient cells was not affected by ferricyanide addition. Addition of 250 μM ferricyanide to iron-limited cells in which photosynthesis was inhibited (either by the presence of glycolaldehyde, or by maintaining the cells at the CO₂ compensation point) resulted in a stimulation in the rate of gross photosynthetic O₂ evolution. Chlorophyll a fluorescence measurements indicated a large increase in non-photochemical quenching during ferricyanide reduction in the light; the increase in nonphotochemical quenching was abolished by the addition of nigericin. These results suggest that reduction of extracellular ferricyanide (mediated at the plasma membrane) interacts with both photosynthesis and respiration, and that both of these processes contribute NADPH in the light. Received: 15 September 1999 / Accepted: 14 October 1999     

Carbon fluxes in the rhizosphere of sweet chestnut seedlings (Castanea sativa) grown under two atmospheric CO2 concentrations: 14C partitioning after pulse labelling

Partitioning of ¹⁴C was assessed in sweet chestnut seedlings (Castanea sativa Mill.) grown in ambient and elevated atmospheric [CO₂] environments during two vegetative cycles. The seedlings were exposed to ¹⁴CO₂ atmosphere in both high and low [CO₂] environments for a 6-day pulse period under controlled laboratory conditions. Six days after exposure to ¹⁴CO₂, the plants were harvested, their dry mass and the radioactivity were evaluated. ¹⁴C concentration in plant tissues, root-soil system respiratory outputs and soil residues (rhizodeposition) were measured. Root production and rhizodeposition were increased in plants growing in elevated atmospheric [CO₂]. When measuring total respiration, i.e. CO₂ released from the root/soil system, it is difficult to separate CO₂ originating from roots and that coming from the rhizospheric microflora. For this reason a model accounting for kinetics of exudate mineralization was used to estimate respiration of rhizospheric microflora and roots separately. Root activity (respiration and exudation) was increased at the higher atmospheric CO₂ concentration. The proportion attributed to root respiration accounted for 70 to 90% of the total respiration. Microbial respiration was related to the amount of organic carbon available in the rhizosphere and showed a seasonal variation dependent upon the balance of root exudation and respiration. The increased carbon assimilated by plants grown under elevated atmospheric [CO₂] stayed equally distributed between these increased root activities. ei]H Lambers     

Photosynthetic studies with mesophyll protoplasts from Notonia grandiflora, a crassulacean acid metabolism plant

A method is described for rapid enzymatic isolation of mesophyll protoplasts and cells from the crassulacean acid metabolism (CAM) plant Notonia grandiflora DC. The mesophyll protoplasts exhibited high rates of ¹⁴CO₂ fixation both in the light (45 μmol of CO₂ fixed mg^?1 Chl h^?1) and in the dark (20 μmol of CO₂ fixed mg^?1 Chl h^?1). The protoplasts also showed O₂ evolution (40 μmol of O₂ evolved mg^?1 Chl h^?1) without added bicarbonate. Exogenously added bicarbonate had no stimulating effect on the O₂ evolution. Analyses of early photosynthetic products in the light showed the formation of both C₃ and C₄ acids. Aspartate was found to be a predominant photosynthate.     

Root and microbial involvement in the kinetics of 14C-partitioning to rhizosphere respiration after a pulse labelling of maize assimilates

In a previous study, we examined the kinetics of radioactivity evolution from rhizosphere respiration after the pulse labelling of maize shoots with ¹⁴CO₂ (Nguyen et al., 1999). The specific activity of rhizosphere respiration demonstrated two peaks of ¹⁴CO₂ production. The first one occurred a few hours after the pulse of ¹⁴CO₂ and was followed by a second peak, which took place during the night following the labelling. In the present work, we demonstrate that the second phase of activity occurred in both sterile and non sterile plant–soil systems. This was inconsistent with the results obtained for wheat by Warembourg and Billès (1979) who observed the second peak solely in the case of non-sterile cultures. These authors suggested that this second phase of ¹⁴CO₂ production was related to microbial mineralisation of labelled complex compounds. Their synthesis and breakdown into smaller molecules delayed their utilisation by micro-organisms. However, in the present work, we also demonstrate that the second phase of activity was closely related to photoperiod. When plants were transferred from a 16 h to 20 h photoperiod, the second mineralisation of labelled rhizosphere compounds occurred sooner after the initiation of the dark period and it was strongly attenuated. Therefore, we suggest that the second phase of activity resulted from the utilisation by roots and by micro-organisms of stored ¹⁴C-compounds, which accumulated during the previous light period.     

Seasonal fluctuations of aerobic cellulolytic bacteria,and cellulase and respiratory activities in a soil profile under a forest

The amount of cellulase activity and¹⁴CO₂ evolution declined with profile depth. These properties varied seasonally, being highest in autumn and lowest in winter. Cytophaga hutchinsonii andCytophaga rubra were the most common species of cellulolytic bacteria found by the dilution-plate method;Bacillus circulans andCellulomonas fimi were also isolated.Cellulolytic bacterial numbers-¹⁴CO₂ evolution, cellulolytic bacterial numbers-cellulase activity and¹⁴CO₂ evolution-cellulase activity were correlated positive-linear and significantly.     

On-line biomass estimation using a modified oxygen utilization rate

A procedure for estimating biomass during batch fermentation from on-line gas analysis is presented. First, the respiratory quotient was used to determine the fraction of the total oxygen utilization rate required for cell maintenance and growth versus product synthesis. The modified oxygen utilization rate was then used to estimate biomass on-line by integrating the oxygen balance for cell synthesis-maintenance. The method is illustrated for the case of L-lysine synthesis by Corynebacterium glutamicum.List of Symbols CER mmol CO₂/l · h carbon dioxide evolution rate - M _{O 2}/x mmol O₂/h · g cells maintenance coefficient - OUR mmol O₂/l · h oxygen utilization rate - OUR _X mmol O₂/l · h OUR fraction for cell maintenance and growth - RQ mmol CO₂/mmol O₂ respiratory quotient(CER/OUR) - X g cells/l biomass concentration - Y _X/O₂ yield coefficients     

Soil microorganisms at different CO2 and O2 tensions

Microbial biomass and activity were determined in cambisol incubated under ambient and increased (up to 2.23 mmol/L) CO₂ concentrations. An immediate negative response of the soil microbial community to [CO]₂ increase was observed during the first day with respect to microbial biomass, soil respiration and specific respiration activity (both expressed as CO₂ evolution). In contrast, O₂ consumption was not affected but anabolic utilization of available substrate increased. These phenomena were observed under conditions of increased CO₂ tension but without any change in O₂ concentration.     

An improved method for the simultaneous determination of photosynthetic O2 evolution and CO2 consumption in Rhizophora mucronata leaves

The photosynthetic gas-exchange has been assessed traditionally either as O₂ evolution or CO₂ consumption. In this study, we used a liquid-phase O₂ electrode combined with CO₂ optodes to examine simultaneously photosynthesis in intact leaves of mangrove Rhizophora mucronata. We verified suitable conditions for leaf photosynthetic rates by assessing pH levels and NaHCO₃ concentrations and compared these to the gas-exchange method at various PAR levels. The photosynthetic rate in response to pH exhibited a similar pattern both for O₂ evolution and CO₂ consumption, and higher rates were associated with intermediate pH compared with low and high pH values. The net photosynthetic quotient (PQ) of R. mucronata leaves ranged from 1.04–1.28. The PQ values, which were never lesser than 1, suggested that photorespiration did not occur in R. mucronata leaves under aqueous conditions. The similar maximum photosynthetic rates suggested that all measurements had a high capacity to adjust the photosynthetic apparatus under a light saturation condition. The simultaneous measurements of O₂ evolution and CO₂ consumption using the Clark oxygen electrode polarographic sensor with the CO₂ optode sensor provided a simple, stable, and precise measurement of PQ under aqueous and saturated light conditions.     

Bacterial conjugation betweenEscherichia coli andPseudomonas spp. donor and recipient cells in soil

Summary Experiments conducted in microcosms containing loam soil samples inoculated with eitherE. coli orPseudomonas spp. donor and recipient cells showed that bacterial cells survived and conjugated over a 24-h incubation period.E. coli transconjugants were detected 6 h after donor and recipient strains were introduced into sterile soil samples. In non-sterile soil samples, transconjugants were detected between 8 and 24 h incubation.Pseudomonas transconjugants were recovered from sterile soil samples between 6 and 12 h after their introduction and as early as 2 h in non-sterile soil. The results show that genetic interactions occur in non-sterile soil in relatively short periods of time at relatively high transfer frequencies (10^–3 to 10^–4). Studies on genetic interactions in soil are becoming necessary in risk assessment/environmental impact studies prior to the release of genetically engineered or modified organisms into uncontained environments.     

Acclimation of leaf respiratory properties in Alocasia odora following reciprocal transfers of plants between high- and low-light environments

Acclimation of respiration to the light environments is important for a plant’s carbon balance. Respiratory rates of mature leaves of Alocasia odora, a typical shade‐tolerant species, were measured during the night for 14 d after reciprocal transfers between high‐ (330 µ mol m^?2 s^?1) and low‐light (20 µ mol m^?2 s^?1) environments. Following the transfer, both the rate of CO₂ efflux and that of O₂ uptake of A. odora leaves adjusted to the new light environments. The O₂‐uptake rates changed more slowly than the CO₂‐efflux rates under the new environments. Leaf mass per area also changed after the transfer. We analysed whether substrate availability or ATP‐consumption rates influence the respiratory acclimation. Since the addition of sucrose to leaf segments did not influence the O₂‐uptake rates, the change of respiratory substrate availability was not responsible for the respiratory acclimation. The addition of an uncoupler induced increases in the O₂‐uptake rates, and the degree of enhancement significantly decreased after the transfer from low to high irradiance. Thus, the change in ATP‐consumption rates was responsible for the changes in respiratory rates in the plants transferred from low to high light. Potential rates of O₂ uptake, as measured in the presence of both the substrate and the uncoupler, changed after the transfer, and strongly correlated with the O₂‐uptake rates, irrespective of the directions of transfer (r = 0·961). There was a strong correlation between maximal activities of NAD‐isocitrate dehydrogenase and the potential rates of O₂ uptake (r = 0·933), but a weaker correlation between those of cytochrome c oxidase and the potential rates (r = 0·689). These data indicate that the changes of light environments altered the respiratory rates via the change of the respiratory ATP demand, and that the altered rates of respiration will induce the changes of the respiratory capacities.     

Effect of O2 and CO2 on net CO2 exchange in a high-CO2-requiring mutant of Chlamydomonas reinhardtii during dark-light-dark transitions

Net CO₂ exchange was monitored through a dark-light-dark transition, under 2% and 21% O₂ in the presence and absence of CO₂, in Chlamydomonas reinhardtii wild type and the high-CO₂-requiring mutant ca-1-12-1C. Upon illumination at 350 l/l CO₂, ca-1-12-1C cell exhibited a large decrease in net CO₂ uptake following an initial surge of CO₂ uptake. Net CO₂ uptake subsequently attained a steady-state rate substantially lower than the maximum. A large, O₂-enchanced post-illumination burst of CO₂ efflux was observed after a 10-min illumination period, corresponding to a minimum in the net CO₂ uptake rate. A smaller, but O₂-insensitive post-illumination burst was observed following a 30-min illumination period, when net CO₂ uptake was at a steady-state rate. These post-illumination bursts appeared to reflect the release of an intracellular pool of inorganic carbon, which was much larger following the initial surge of net CO₂ uptake than during the subsequent steady-state CO₂ uptake period.With the mutant in CO₂-free gas, O₂-stimulated, net CO₂ efflux was observed in the light, and a small, O₂-dependent post-illumination burst was observed. With wild-type cells no CO₂ efflux was observed in the light in CO₂-free gas under either 2% or 21% O₂, but a small, O₂-dependent post-illumination burst was observed. These results were interpreted as indicating that photorespiratory rates were similar in the mutant and wild-type cells in the absence of CO₂, but that the wild-type cells were better able to scavenge the photorespiratory CO₂.