Floating Aquatic Macrophytes Can Substantially Offset Open Water CO<Subscript>2</Subscript> Emissions from Tropical Floodplain Lake Ecosystems

Tropical floodplain lake ecosystems are recognized as important sources of carbon (C) from the water to the atmosphere. They receive large amounts of organic matter and nutrients from the watershed, leading to intense net heterotrophy and carbon dioxide (CO₂) emission from open waters. However, the role of extensive stands of floating macrophytes colonizing floodplains areas is still neglected in assessments of net ecosystem exchange of CO₂ (NEE). We assessed rates of air-lake CO₂ flux using static chambers in both open waters and waters covered by the widespread floating aquatic macrophyte (water hyacinth; Eichornia sp.) in two tropical floodplain lakes in Pantanal, Brazil during different hydrological seasons. In both lakes, areas colonized by floating macrophytes were a net CO₂ sink during all seasons. In contrast, open waters emitted CO₂, with higher emissions during the rising and high water periods. Our results indicate that the lake NEE can be substantially overestimated (fivefold or more in the studied lakes) if the carbon fixation by macrophytes is not considered. The contribution of these plants can lead to neutral or negative NEE (that is, net uptake of CO₂) on a yearly basis. This highlights the importance of floating aquatic macrophytes for the C balance in shallow lakes and extensive floodplain areas.     

Changes in benthic macroinvertebrate abundance and lake isotope (C,N) signals following biomanipulation: an 18-year study in shallow Lake Vaeng,Denmark

Change in the abundance of benthic macroinvertebrates and the stable isotope composition (C, N) of benthic invertebrates and zooplankton in Lake Vaeng, Denmark, was investigated over an 18-year period following biomanipulation (removal of cyprinids). During the first nine years after biomanipulation, the lake was clear and submerged macrophytes were abundant; after this period, a shift occurred to low plant abundance and high turbidity. Two years after the biomanipulation, total density of benthic macroinvertebrates reached a maximum of 17042 (±2335 SE) individuals m⁻² and the density was overall higher when the lake was in a clear state. Redundancy analysis (RDA) suggested macrophyte abundance and total nitrogen (TN) concentration were the dominant structuring forces on the benthic macroinvertebrate assemblage. Stable isotope analysis revealed that δ¹³C of macroinvertebrates and zooplankton was markedly higher in years with high submerged macrophyte abundance than in years without macrophytes, most likely reflecting elevated δ¹³C of phytoplankton and periphyton mediated by a macrophyte-induced lowering of lake water CO₂ concentrations. We conclude that the strong relationship between macrophyte coverage and δ¹³C of macroinvertebrates and cladocerans may be useful in paleoecological studies of past changes in the dynamics of shallow lakes, as change in macrophyte abundance may be tracked by the δ¹³C of invertebrate remains in the sediment.     

Regional Variability and Drivers of Below Ice CO<Subscript>2</Subscript> in Boreal and Subarctic Lakes

Northern lakes are ice-covered for considerable portions of the year, where carbon dioxide (CO₂) can accumulate below ice, subsequently leading to high CO₂ emissions at ice-melt. Current knowledge on the regional control and variability of below ice partial pressure of carbon dioxide (pCO₂) is lacking, creating a gap in our understanding of how ice cover dynamics affect the CO₂ accumulation below ice and therefore CO₂ emissions from inland waters during the ice-melt period. To narrow this gap, we identified the drivers of below ice pCO₂ variation across 506 Swedish and Finnish lakes using water chemistry, lake morphometry, catchment characteristics, lake position, and climate variables. We found that lake depth and trophic status were the most important variables explaining variations in below ice pCO₂ across the 506 lakes_. Together, lake morphometry and water chemistry explained 53% of the site-to-site variation in below ice pCO₂. Regional climate (including ice cover duration) and latitude only explained 7% of the variation in below ice pCO₂. Thus, our results suggest that on a regional scale a shortening of the ice cover period on lakes may not directly affect the accumulation of CO₂ below ice but rather indirectly through increased mobility of nutrients and carbon loading to lakes. Thus, given that climate-induced changes are most evident in northern ecosystems, adequately predicting the consequences of a changing climate on future CO₂ emission estimates from northern lakes involves monitoring changes not only to ice cover but also to changes in the trophic status of lakes.     

Primary production of aquatic macrophytes and their epiphytes in two shallow lakes (Peipsi and Võrtsjärv) in Estonia

In shallow lakes with large littoral zones, epiphytes and submerged macrophytes can make an important contribution to the total annual primary production. We investigated the primary production (PP) of phytoplankton, submerged macrophytes, and their epiphytes, from June to August 2005, in two large shallow lakes. The production of pelagic and littoral phytoplankton and of the dominant submerged macrophytes in the littoral zone (Potamogeton perfoliatus in Lake Peipsi and P. perfoliatus and Myriopyllum spicatum in Lake Võrtsjärv) and of their epiphytes was measured using a modified ¹⁴C method. The total PP of the submerged macrophyte area was similar in both lakes: 12.4 g C m^?2 day^?1 in Peipsi and 12.0 g C m^?2 day^?1 in Võrtsjärv. In Peipsi, 84.2% of this production was accounted for by macrophytes, while the shares of phytoplankton and epiphytes were low (15.6 and 0.16%, respectively). In Võrtsjärv, macrophytes contributed 58%, phytoplankton 41.9% and epiphytes 0.1% of the PP in the submerged macrophyte area. Epiphyte production in both lakes was very low in comparison with that of phytoplankton and macrophytes: 0.01, 5.04, and 6.97 g C m^?2 day^?1, respectively, in Võrtsjärv, and 0.02, 1.93, and 10.5 g C m^?2 day^?1, respectively, in Peipsi. The PP of the littoral area contributed 10% of the total summer PP of Lake Peipsi sensu stricto and 35.5% of the total summer PP of Lake Võrtsjärv.     

Can artificial plant beds be used to enhance macroinvertebrate food resources for perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) during the initial phase of lake restoration by cyprinid removal?

Restoration of shallow lakes to a clear-water state, often characterized by high submerged macrophyte cover and a high proportion of piscivores such as perch, Perca fluviatilis L., frequently involves removal of a large proportion of the zoobenthivorous fish, such as bream, Abramis brama L., and roach, Rutilus rutilus L. (i.e. biomanipulation). However, establishment of submerged macrophytes is often delayed following fish removal. This is unfortunate because plant beds typically host high densities of the macroinvertebrates constituting the diet of small perch and thus help perch to go through the bottleneck from feeding on macroinvertebrates to feeding on fish. Establishment of artificial plant beds may be a useful tool to enhance macroinvertebrate population growth and thus food resources for small perch until the natural plants have established. To investigate this restoration option, we studied during two growing seasons (June–October) the composition and abundance of the macroinvertebrate community in artificial plant beds installed in shallow Lake Væng (Denmark) comprising the initial phase of a biomanipulation effort by fish removal. Lake areas with artificial plant beds exhibited substantially higher macroinvertebrate densities than the lake bottom. This suggests that artificial plant beds may be used as feeding grounds for small perch, similarly to the well-known refuge effect for zooplankton against fish predation. In this way, artificial plant beds could help maintain a clear-water state during the transient period when natural submerged vegetation is not yet established in the lake.     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Impacts of sediment type on the performance and composition of submerged macrophyte communities

To restore deteriorated lake ecosystems, it is important to identify environmental factors that influence submerged macrophyte communities. While sediment is a critical environmental factor for submerged macrophytes and many studies have examined effects of sediment type on the growth of individual submerged macrophytes, very few have tested how sediment type affects the growth and species composition of submerged macrophyte communities. We constructed submerged macrophyte communities containing four co-occurring submerged macrophytes (Hydrilla verticillata, Myriophyllum spicatum, Ceratophyllum demersum and Chara fragilis) and subjected them to three sediment treatments, i.e., clay, a mixture of clay and quartz sand at a volume ratio of 1:1 and a mixture at a volume ratio of 1:4. Compared to the clay, the 1:1 mixture treatment greatly increased overall biomass, number of shoot nodes and shoot length of the community, but decreased its diversity. This was because it substantially promoted the growth of H. verticillata within the community, making it the most abundant species in the mixture sediment, but decreased that of M. spicatum and C. demersum. The sediment type had no significant effects on the growth of C. fragilis. As a primary nutrient source for plant growth, sediment type can have differential effects on various submerged macrophyte species and 1:1 mixture treatment could enhance the performance of the communities, increasing the overall biomass, number of shoot nodes and shoot length by 39.03%, 150.13% and 9.94%, respectively, compared to the clay treatment. Thus, measures should be taken to mediate the sediment condition to restore submerged macrophyte communities with different dominant species.     

Seasonality of chlorophyll and nutrients in Lake Erken – effects of weather conditions

The effect of submerged macrophytes on interactions among epilimnetic phosphorus, phytoplankton, and heterotrophic bacterioplankton has been acknowledged, but remains poorly understood. Here, we test the hypotheses that the mean summer phytoplankton biomass (chlorophyll a): phosphorus ratios decrease with increased macrophyte cover in a series of nine lakes. Further, we test that both planktonic respiration and bacterioplankton production increase with respect to phytoplankton biomass along the same gradient of increasing macrophyte cover. Increased macrophyte cover was associated with a lower fraction of particulate phosphorus in epilimnia, with total particulate phosphorus declining from over 80% of total phosphorus in a macrophyte free lake to less than 50% in a macrophyte rich lake. Phytoplankton biomass (chlorophyll a) too was lower in macrophyte dominated lakes, despite relatively high levels of total dissolved phosphorus. Planktonic respiration and bacterioplankton production were higher in macrophyte rich lakes than would be expected from phytoplankton biomass alone, pointing to a subsidy of bacterioplankton metabolism by macrophyte beds at the whole lake scale. The results suggest that the classical view of pelagic interactions, which proposes phosphorus determines phytoplankton abundance, which in turn determines bacterial abundance through the production of organic carbon, becomes less relevant as macrophyte cover increases.     

Diversity and composition of bacterial community in the rhizosphere sediments of submerged macrophytes revealed by 454 pyrosequencing

Freshwater lake sediments support a variety of submerged macrophytes that may host groups of bacteria exerting important ecological functions. We collected three kinds of commonly found submerged macrophyte species (Ceratophyllum demersum, Vallisneria spiralis and Elodea nuttallii) to investigate the bacterial community associated with their rhizosphere sediments. High-throughput 454 pyrosequencing and bioinformatics analyses were performed to examine the diversity and composition of the bacterial community. The results obtained indicated that the diversity of the bacterial community associated with the rhizosphere sediments of submerged macrophytes was significantly lower than that of the bulk sediment. Remarkable differences in the bacterial community composition between the rhizosphere and bulk sediments were also observed.     

Macrophytes as dispersal vectors of zooplankton resting stages in a subtropical riverine floodplain

The resting stages of freshwater zooplankton constitute a special mechanism for passive dispersal, often displaying a variety of adaptations so as to ease transport. In floodplain systems, macrophytes are one of the most representative biotic groups showing interactions with the zooplankton community. The annual fluctuations in the hydrometric level of the Paraná River favour the displacement of this aquatic vegetation in floodplain environments. This paper hypothesizes that the roots and submerged portions of different macrophytes contain zooplankton resting stages which are able to hatch when environmental conditions are favourable. In turn, this contributes to the dispersal of zooplankton by plants when they are displaced by the flood pulse. Six macrophyte species were sampled (Eichhornia crassipes, Azolla filiculoides, Limnobium spongia, Pistia stratiotes, Eichhornia azurea and Nymphoides indica) from lakes within the Paraná River floodplain. Roots and submerged portions of vegetation were stored (90 days) at 4 °C then incubated at 25 °C for 90 days. Hatchling emergence was recorded at 2-day intervals during this period. In total, 70 zooplankton taxa were recorded in all macrophyte samples; rotifers were the most representative group (69%) followed by cladocerans (28%) and copepods (3%). The roots and submerged parts of aquatic vegetation house viable zooplankton resting stages. This phenomenon allows the dispersal of resting stages and therefore colonization of new habitats during the displacement of macrophyte species.     

Above- and belowground competition between two submersed macrophytes

Several studies have shown that submerged macrophytes provide a refuge for zooplankton against fish predation, whereas the role of emergent and floating-leaved species, which are often dominant in eutrophic turbid lakes, is far less investigated. Zooplankton density in open water and amongst emergent and floating-leaved vegetation was monitored in a small, eutrophic lake (Frederiksborg Slotssø) in Denmark during July–October 2006. Emergent and floating-leaved macrophytes harboured significantly higher densities of pelagic as well as plant-associated zooplankton species, compared to the open water, even during periods where the predation pressure was presumably high (during the recruitment of 0+ fish fry). Zooplankton abundance in open water and among vegetation exhibited low values in July and peaked in August. Bosmina and Ceriodaphnia dominated the zooplankton community in the littoral vegetated areas (up to 4,400 ind l^?1 among Phragmites australis and 11,000 ind l^?1 between Polygonum amphibium stands), whereas the dominant species in the pelagic were Daphnia (up to 67 ind l^?1) and Cyclops (41 ind l^?1). The zooplankton density pattern observed was probably a consequence of concomitant modifications in the predation pressure, refuge availability and concentration of cyanobacteria in the lake. It is suggested that emergent and floating-leaved macrophytes may play an important role in enhancing water clarity due to increased grazing pressure by zooplankton migrating into the plant stands. As a consequence, especially in turbid lakes, the ecological role of these functional types of vegetation, and not merely that of submerged macrophyte species, should be taken into consideration.     

Assessment of anthropogenic pressures on South European Atlantic bogs (NW Spain) based on hydrochemical data

Cyanobacterial exudates are known to allelopathically inhibit submerged macrophytes, but the influence of the cyanobacteria growth phase on this effect is yet unknown. We compared the effect of exudates of the exponential growth phase of Microcystis aeruginosa Kütz. Elenk with exudates during the decline phase on seedlings of the macrophyte species Potamogeton crispus L. Biomass, chlorophyll content, the ratio of variable–maximum fluorescence (F _v/F _m), and light response capacity of P. crispus seedings were significantly inhibited when affected by M. aeruginosa exudates of the exponential growth phase but promoted by exudates of the decline phase. Tiller numbers of P. crispus increased by 350% under the influence of exponential phase exudates, but decreased by 60% when decline phase exudates were applied. Both exudates increased the malondialdehyde contents and decreased the activity of superoxide dismutase and peroxidase in P. crispus seedlings. We conclude that the exponential growth phase of cyanobacteria rather than the decline phase is important for disrupting photosynthesis and for inducing oxidative stress in submerged macrophytes. Planting P. crispus should thus not be applied in summer but during the cyanobacteria decline phase.     

Effects of phytoplankton on the distribution of submerged macrophytes in a small canal

Submerged macrophytes have been disappearing from the Kanto Plain, Japan since the 1960s. This disappearance is usually attributable to the interaction between macrophytes and phytoplankton. Phytoplankton contributes to shading of the available light and changes the availability of inorganic carbon from free CO₂ to HCO ₃ ^? for use in photosynthesis. However, limited information is available about the interaction between carbon fraction and submerged macrophytes through phytoplankton abundance. In this short note, we observe the distribution of submerged macrophytes and phytoplankton in a small canal. We found that, despite high photosynthetically active radiation (PAR) in the downstream region, low free CO₂ concentration through phytoplankton abundance can deplete free CO₂ for submerged macrophytes. In contrast, the upstream region exhibited macrophytes in an environment with high free CO₂ concentration. The stable carbon isotope ratio of submerged macrophytes follows this pattern, with more positive values occurring in the downstream region and more negative values in the upstream region. It has been reported that phytoplankton limits light availability for submerged macrophytes, but carbon availability could also be a factor in the distribution of submerged macrophytes. Because the source of water for submerged macrophytes is groundwater, its preservation possibly plays a key role for the restoration of submerged macrophytes.     

Ontogenetic changes in the predator–prey interactions between threadsail filefish and moon jellyfish

The global climate change may lead to more extreme climate events such as severe flooding creating excessive pulse-loading of nutrients, including nitrogen (N), to freshwaters. We conducted a 3-month mesocosm study to investigate the responses of phytoplankton, zooplankton and Vallisneria spinulosa to different N loading patterns using weekly and monthly additions of in total 14 g N m^?2 month^?1 during the first 2 months. The monthly additions led to higher phytoplankton chlorophyll a and total phytoplankton biomass than at ambient conditions as well as lower leaf biomass and a smaller ramet number of V. spinulosa. Moreover, the biomass of cyanobacteria was higher during summer (August) in the monthly treatments than those with weekly or no additions. However, the biomass of plankton and macrophytes did not differ among the N treatments at the end of the experiment, 1 month after the termination of N addition. We conclude that by stimulating the growth of phytoplankton (cyanobacteria) and reducing the growth of submerged macrophytes, short-term extreme N loading may have significant effects on shallow nutrient-rich lakes and that the lakes may show fast recovery if they are not close to the threshold of a regime shift from a clear to a turbid state.     

Performance of a new phytoplankton composition metric along a eutrophication gradient in Nordic lakes

A new phytoplankton metric is presented, which is developed from a large dataset of Norwegian lakes (>2,000 samples from >400 lakes). In contrast to previous metrics, this index is not built on selected ‘indicative’ taxa, but uses all available taxonomic information at genus and species level. Taxa optima with respect to lake trophic status (derived from total phosphorus concentrations) are used to calculate a phytoplankton trophic index (TI) for each sample. Analysis of the TI shows that phytoplankton communities exhibit highly non-linear responses to eutrophication in Norwegian lakes. Reference lakes are characterized by very similar TIs despite having considerable variation in total phosphorus and chlorophyll a concentrations. TI exhibits a non-linear distribution along the eutrophication gradient which separates unimpacted from impacted sites in the study area. We further show that TI exhibits smaller seasonal variations than chlorophyll a, making it a more reliable indicator for lake monitoring. Implications for its applicability within the WFD are discussed.     

Effects of macrophytes on lake‐water quality across latitudes: a meta‐analysis

Macrophytes are widely recognized for improving water quality and stabilizing the desirable clear‐water state in lakes. The positive effects of macrophytes on water quality have been noted to be weaker in the (sub)tropics compared to those of temperate regions. We conducted a global meta‐analysis using 47 studies that met our set criteria to assess the overall effects of macrophytes on water quality (measured by phytoplankton chlorophyll a concentration, total nitrogen concentration, total phosphorus concentration, Secchi depth and the trophic state index) and to investigate how these effects correlate with latitude using meta‐regressions. We also examined if the effects of macrophytes on lake‐water quality differ with growth form and study design in (sub)tropical and temperate areas by grouping the data and then comparing the effect sizes. We found that macrophytes significantly reduced phytoplankton chlorophyll a concentration, total nitrogen concentration, total phosphorus concentration, as well as the trophic state index, but they did not have a significant overall effect on Secchi depth. The effects of macrophytes on reducing phytoplankton chlorophyll a concentration, total nitrogen concentration and the trophic state index did not differ with latitude. However, the reduction of total phosphorus concentration was greater at lower latitudes. We showed that at lower latitudes, the positive effects of macrophytes on water quality are similar to or greater than those at higher latitudes, thus challenging the prevailing paradigm of macrophytes being less effective at enhancing lake‐water quality in the (sub)tropics. Furthermore, our data showed that the macrophyte effects vary by growth forms, and the growth forms that positively affect water quality differ between the (sub)tropical and temperate areas. We showed a lack of significant macrophyte effects in surveys within and outside macrophyte stands, suggesting difference in the sensitivities of study designs or possibly weaker effects of macrophytes in lakes compared to experimental settings.     

Effect of red and blue light on acclimation of <Emphasis Type="Italic">Chlamydomonas reinhardtii</Emphasis> to CO<Subscript>2</Subscript>-limiting conditions

Effects of red (RL) and blue (BL) light on acclimation of the unicellular green alga Chlamydomonas reinhardtii to the low level of ambient CO₂ were studied. C. reinhardtii cells grown at 5% CO₂ and under white light (170 μmol/(m²s)) had a relatively low activity of extracellular carbonic anhydrase (CA), a low affinity for dissolved inorganic carbon, and a low rate of photosynthesis under CO₂-limiting conditions. These cells readily started acclimation to the low CO₂ concentration when they were exposed to atmospheric air (～ 0.03% CO₂) under RL or BL (150 μmol/(m² s) each). The acclimation was manifested in a significant increase in the CO₂-limited rate of photosynthesis, the affinity for dissolved inorganic carbon, and the extracellular CA activity with no difference between RL-and BL-cells. Independently of light quality, the acclimation was completed for 5–7 h after cell exposure to air. As is evident from RL-and BL-dependent changes in the sum of chlorophylls and chlorophyll a/b ratio, transfer of C. reinhardtii cells to air and RL or BL triggered also the process of algal photosynthetic adaptation to light quality. However, this process did not interfere with acclimation to low CO₂ because started 4 h later. On the basis of similarity in the low CO₂-induced changes under RL and BL, it is concluded that acclimation of C. reinhardtii to CO₂-limiting conditions does not depend on light quality.     

Simple generalisation of a mesophyll resistance model for various intracellular arrangements of chloroplasts and mitochondria in C<Subscript>3</Subscript> leaves

The classical definition of mesophyll conductance (g _m) represents an apparent parameter (g _m,app) as it places (photo)respired CO₂ at the same compartment where the carboxylation by Rubisco takes place. Recently, Tholen and co-workers developed a framework, in which g _m better describes a physical diffusional parameter (g _m,dif). They partitioned mesophyll resistance (r _m,dif = 1/g _m,dif) into two components, cell wall and plasmalemma resistance (r _wp) and chloroplast resistance (r _ch), and showed that g _m,app is sensitive to the ratio of photorespiratory (F) and respiratory (R _d) CO₂ release to net CO₂ uptake (A): g _m,app = g _m,dif/[1?+?ω(F?+?R _d)/A], where ω is the fraction of r _ch in r _m,dif. We herein extend the framework further by considering various scenarios for the intracellular arrangement of chloroplasts and mitochondria. We show that the formula of Tholen et al. implies either that mitochondria, where (photo)respired CO₂ is released, locate between the plasmalemma and the chloroplast continuum or that CO₂ in the cytosol is completely mixed. However, the model of Tholen et al. is still valid if ω is replaced by ω(1?σ), where σ is the fraction of (photo)respired CO₂ that experiences r _ch (in addition to r _wp and stomatal resistance) if this CO₂ is to escape from being refixed. Therefore, responses of g _m,app to (F?+?R _d)/A lie somewhere between no sensitivity in the classical method (σ =1) and high sensitivity in the model of Tholen et al. (σ =0).     

Stomatal conductance in mature deciduous forest trees exposed to elevated CO<Subscript>2</Subscript>

Stomatal conductance (g _s) of mature trees exposed to elevated CO₂ concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g _s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO₂ compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO₂ treatment. During the 2003 drought in the third treatment year, the CO₂ effect became reversed in Carpinus, resulting in higher g _s in trees exposed to elevated CO₂ compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO₂ enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO₂-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO₂.     

Photosynthetic CO<Subscript>2</Subscript> uptake in seedlings of two tropical tree species exposed to oscillating elevated concentrations of CO<Subscript>2</Subscript>

Do short-term fluctuations in CO₂ concentrations at elevated CO₂ levels affect net CO₂ uptake rates of plants? When exposed to 600 μl CO₂ l^?1, net CO₂ uptake rates in shoots or leaves of seedlings of two tropical C₃ tree species, teak (Tectona grandis L. f.) and barrigon [Pseudobombax septenatum (Jacq.) Dug.], increased by 28 and 52% respectively. In the presence of oscillations with half-cycles of 20 s, amplitude of ca. 170 μl CO₂ l^?1 and mean of 600 μl CO₂ l^?1, the stimulation in net CO₂ uptake by the two species was reduced to 19 and 36%, respectively, i.e. the CO₂ stimulation in photosynthesis associated with a change in exposure from 370 to 600 μl CO₂ l^?1 was reduced by a third in both species. Similar reductions in CO₂-stimulated net CO₂ uptake were observed in T. grandis exposed to 40-s oscillations. Rates of CO₂ efflux in the dark by whole shoots of T. grandis decreased by 4.8% upon exposure of plants grown at 370 μl CO₂ l^?1 to 600 μl CO₂ l^?1. The potential implications of the observations on CO₂ oscillations and dark respiration are discussed in the context of free-air CO₂ enrichment (FACE) systems in which short-term fluctuations of CO₂ concentration are a common feature.