Photosynthesis of amphibious and obligately submerged plants in CO2-rich lowland streams

Small unshaded streams in lowland regions receive drainage water with high concentrations of free␣CO₂, and they support an abundant growth of amphibious and obligately submerged plants. Our first objective was to measure the CO₂ regime during summer in a wide range of small alkaline Danish streams subject to wide variation in temperature, O₂ and CO₂ during the day. The second objective was to determine the effect of these variations on daily changes in light-saturated photosynthesis in water of a homophyllous and a heterophyllous amphibious species that only used CO₂, and an obligately submerged species capable of using both HCO⁻ ₃ and CO₂. We found that the median CO₂ concentrations of the streams were 11 and 6 times above air saturation in the morning and the afternoon, respectively, but stream sites with dense plant growth had CO₂ concentrations approaching air saturation in the afternoon. In contrast, outlets from lakes had low CO₂ concentrations close to, or below, air saturation. The amphibious species showed a reduction of photosynthesis in water from morning to afternoon along with the decline in CO₂ concentrations, while increasing temperature and O₂ had little effect on photosynthesis. Photosynthesis of the obligately submerged species varied little with the change of CO₂ because of HCO₃ ⁻- use, and variations were mostly due to changes in O₂ concentration. Independent measurements showed that changes in temperature, O₂ and CO₂ could account for the daily variability of photosynthesis of all three species in water. The results imply that CO₂ supersaturation in small lowland streams is important for the rich representation of amphibious species and their contribution to system photosynthesis. Received: 2 March 1998 / Accepted: 24 July 1998     

Regulation of photosynthetic rates of submerged rooted macrophytes

Summary Fourteen temperate, submerged macrophytes were cultivated in the laboratory at high DIC levels (3.3–3.8 mM), 10.4–14.4 mol photons (PAR) m^-2 d^-1 and 15°C. Photosynthesis at photosaturation ranged between 0.59 and 17.98 mg O₂ g^-1 DW h^-1 at ambient pH (8.3) and were markedly higher between 1.76 and 47.11 mg O₂ g^-1 DW h^-1 at pH 6.5 under elevated CO₂ concentrations. Photosynthetic rates were significantly related to both the relative surface area and the chlorophyll content of the leaves. Consequently, the photosynthetic rate was much less variable among the species when expressed per surface area and chlorophyll content instead of dry mass. The chlorophyll content was probably a main predictor of photosynthesis of submerged leaves because of the direct relationship of chlorophyll to the light harvesting capacity and/or a coupling to the capacity for photosynthetic electron transport and carboxylation. A comparison with terrestrial leaves characterized the submerged leaves by their low chlorophyll concentrations and low photosynthetic rates per surface area due to the thin leaves. Photosynthetic rates per chlorophyll content in submerged leaves at CO₂ saturation, however, were at the same level as photosynthesis in terrestrial leaves measured at ambient CO₂ when appropriate corrections were made for differences in incubation temperature.     

Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water

The transition zone between terrestrial and freshwater habitats is highly dynamic, with large variability in environmental characteristics. Here, we investigate how these characteristics influence the nutritional status and performance of plant life forms inhabiting this zone. Specifically, we hypothesised that: (i) tissue nutrient content differs among submerged, amphibious and terrestrial species, with higher content in submerged species; and (ii) PNUE gradually increases from submerged over amphibious to terrestrial species, reflecting differences in the availability of N and P relative to inorganic C across the land–water ecotone. We found that tissue nutrient content was generally higher in submerged species and C:N and C:P ratios indicated that content was limiting for growth for ca. 20% of plant individuals, particularly those belonging to amphibious and terrestrial species groups. As predicted, the PNUE increased from submerged over amphibious to terrestrial species. We suggest that this pattern reflects that amphibious and terrestrial species allocate proportionally more nutrients into processes of importance for photosynthesis at saturating CO₂ availability, i.e. enzymes involved in substrate regeneration, compared to submerged species that are acclimated to lower availability of CO₂ in the aquatic environment. Our results indicate that enhanced nutrient loading may affect relative abundance of the three species groups in the land–water ecotone of stream ecosystems. Thus, species of amphibious and terrestrial species groups are likely to benefit more from enhanced nutrient availability in terms of faster growth compared to aquatic species, and that this can be detrimental to aquatic species growing in the land–water ecotone, e.g. Ranunculus and Callitriche.     

Plant distribution and abundance in relation to physical conditions and location within Danish stream systems

The distribution of obligate submerged plants, amphibious plants and terrestrial plants in streams was examined in relation to water depth, substrate and distance to the banks using univariate and mulitivariate analyses. The analyses were based on recordings in more than 40000 quadrats (25×25 cm) in 208 unshaded sites in the predominantly small and shallow (<1.6 m) Danish streams. Also, the distribution of plant abundance and richness from the source to the outlet of the stream system was determined.The submerged plants in Danish streams include 87 terrestrial, 22 amphibious and 30 obligate submerged taxa. The distribution of plant types was mainly related to water depth and distance to the bank among the physical conditions, while the type of bottom substrate had no significant influence. Terrestrial plants and amphibious plants (excluding Sparganium emersum) dominated in shallow water near the bank, but declined rapidly with increasing depth and distance to the bank, reflecting the importance of dispersal by ingrowth from populations on the banks to the water among these plants. Accordingly, these two plant groups constituted a higher proportion of total plant abundance in small streams than large streams. S. emersum dominated on great depth and distance to the banks, probably reflecting the lengthwise dispersal of this species from upstream to downstream parts of the stream system, the tolerance of the species to weed cutting and the adaptation to grow at low light intensities. Obligate submerged plants dominated at intermediate depths and at all distances from the bank except from 0 to 50 cm. This distribution reflects the ability of these species to disperse lengthwise in streams and live permanently submerged.The species number of all species and obligate submerged plants was lower in the smallest stream sites compared to larger downstream sites, while there was no difference for terrestrial and amphibious plants. The downstream increase of submerged species can be explained by the increase of habitat area and the dispersal of plants with the current, implying that the species pool accumulates with distance from the source. This result is at variance with a maximum richness of submerged plant species predicted for intermediate-sized streams according to the River Continuum Concept developed for large North American streams having forested shallow upstream sections and unshaded, deep downstream sections both unsuitable to submerged plant growth. The results for Danish streams imply that both the longitudinal connection through the flowing water and the transversal connection to the local species on the banks are important for plant distribution in the streams.     

CO2 is the main inorganic C species entering photosynthetically active leaf protoplasts of the freshwater macrophyte Ranunculus penicillatus ssp. pseudofluitans

Submerged aquatic macrophytes growing in water where free CO₂ is unavailable (above pH 8·2) must use mechanisms to supply external dissolved inorganic carbon in a form available to chloroplasts (CO₂). Active transport of HCO₃^– across the plasmalemma has not been proven to be widespread in aquatic macrophytes and catalytic conversion of HCO₃^– to CO₂ is the usual supply mechanism in submerged macrophytes. The interaction of leaf form and function in this respect was investigated in the linear, submerged leaves of Ranunculus penicillatus (Dumort.) Bab ssp. pseudofluitans (Syme) S.Webster. Viable protoplasts were isolated using a mixture of cell wall degrading enzymes optimized for this species. Protoplast viabilities greater than 80% after 5 h of isolation were achieved. Photosynthetic rates of isolated protoplasts were comparable with that of intact plant tissue. Results of carbon isotopic disequilibrium experiments showed that CO₂ was the preferred species of dissolved inorganic carbon for photosynthesis by protoplasts and that HCO₃^– which predominates in the plant’s natural environment mainly contributes by supplying CO₂ outside the cells.     

Why the free floating macrophyte Stratiotes aloides mainly grows in highly CO2-supersaturated waters

We examined how the freely floating macrophyte, Stratiotes aloides L., sampled from a CO₂-supersaturated pond, changes leaf morphology, photosynthesis and inorganic carbon acquisition during its different submerged and emerged life stages in order to evaluate whether S. aloides requires consistently supersaturated CO₂ conditions to grow and complete its life cycle. Submerged rosettes formed from over-wintering turions had typical traits of submerged plants with high specific leaf area and low chlorophyll a concentrations. Emergent leaf parts of mature, floating specimens had typical terrestrial traits with stomata, low specific leaf area and high chlorophyll a content, while offsets formed vegetatively and basal, submerged parts of mature plants showed traits in between. All submerged leaf types exhibited some ability to use HCO₃⁻ but only rosettes formed from turions had efficient HCO₃⁻ use. Rosettes also had the highest CO₂ affinity and maximum CO₂-saturated photosynthesis in water. Half-saturation constants for CO₂ (21–74 μM CO₂) were for all submerged leaf parts 5–140 times lower than the concentrations of free CO₂ in the pond (350–2800 μM CO₂). Emergent leaves were less efficient in water but had significantly higher photosynthesis than submerged, mature leaf parts in air, and rates of photosynthesis of emergent leaves in air were three to five times higher than rates of CO₂-saturated photosynthesis of the three submerged leaf types in water. Underwater photosynthetic rates estimated at CO₂ concentrations corresponding to air equilibrium were not sufficiently high to support any noticeable growth except for rosettes, in which bicarbonate utilization combined with high CO₂ affinity resulted in photosynthetic rates corresponding to almost 34% of maximum rates at high free CO₂. We conclude that S. aloides requires consistently high CO₂-supersaturation to support high growth and to complete its life cycle, and we infer that this requirement explains why S. aloides mainly grows in ponds, ditches and reed zones that are characterized by strong CO₂-supersaturation.     

Induction of reduced photorespiratory activity in submersed and amphibious aquatic macrophytes

Incubation under water in a 30 C/14-hour or 12 C/10-hour photoperiod caused the CO₂ compensation points of 10 aquatic macrophytes to decrease below 25 or increase above 50 microliters CO₂ per liter, respectively. Submerged and aerial leaves of two amphibious angiosperms (Myriophyllum brasiliense and Proserpinaca palustris) maintained high compensation points when incubated in air but, when the submerged or aerial leaves of Proserpinaca were incubated under water, the compensation points dropped as low as 10. This suggests that, in addition to temperature and photoperiod, some factor associated with submergence regulates the compensation point of aquatic plants. In the high-compensation point plants, photorespiration, as a percentage of net photosynthesis, was equivalent to that in terrestrial C₃ plants. For Hydrilla verticillata, the decreasing CO₂ compensation points (110, 40, and 10) were associated with reduced photorespiration, as indicated by decreased O₂ inhibition, decreased rates of CO₂ evolution into CO₂-free air, and increased net photosynthetic rates.     

Why the free floating macrophyte Stratiotes aloides mainly grows in highly CO2-supersaturated waters

We examined how the freely floating macrophyte, Stratiotes aloides L., sampled from a CO₂-supersaturated pond, changes leaf morphology, photosynthesis and inorganic carbon acquisition during its different submerged and emerged life stages in order to evaluate whether S. aloides requires consistently supersaturated CO₂ conditions to grow and complete its life cycle. Submerged rosettes formed from over-wintering turions had typical traits of submerged plants with high specific leaf area and low chlorophyll a concentrations. Emergent leaf parts of mature, floating specimens had typical terrestrial traits with stomata, low specific leaf area and high chlorophyll a content, while offsets formed vegetatively and basal, submerged parts of mature plants showed traits in between. All submerged leaf types exhibited some ability to use HCO₃⁻ but only rosettes formed from turions had efficient HCO₃⁻ use. Rosettes also had the highest CO₂ affinity and maximum CO₂-saturated photosynthesis in water. Half-saturation constants for CO₂ (21–74 μM CO₂) were for all submerged leaf parts 5–140 times lower than the concentrations of free CO₂ in the pond (350–2800 μM CO₂). Emergent leaves were less efficient in water but had significantly higher photosynthesis than submerged, mature leaf parts in air, and rates of photosynthesis of emergent leaves in air were three to five times higher than rates of CO₂-saturated photosynthesis of the three submerged leaf types in water. Underwater photosynthetic rates estimated at CO₂ concentrations corresponding to air equilibrium were not sufficiently high to support any noticeable growth except for rosettes, in which bicarbonate utilization combined with high CO₂ affinity resulted in photosynthetic rates corresponding to almost 34% of maximum rates at high free CO₂. We conclude that S. aloides requires consistently high CO₂-supersaturation to support high growth and to complete its life cycle, and we infer that this requirement explains why S. aloides mainly grows in ponds, ditches and reed zones that are characterized by strong CO₂-supersaturation.     

Morphological and anatomical changes of Melaleuca cajuputi under submergence

Melaleuca cajuputi is a woody plant of the Myrtaceae which is a dominant species in tropical peat swamps in southern Thailand, where the groundwater level fluctuates greatly. Although the current year seedlings are likely submerged, their adaptive responses have never been studied. The objective of the present study was to examine their responses to submergence, and especially their morphological and anatomical changes. Not only did the seedlings of M. cajuputi survive submergence for 56 days, but they could also increase their dry weight, shoot length, and leaf number during submergence. These growth responses to submergence indicate that the seedlings of M. cajuputi could make photosynthetic production under water. The leaves that developed under water were heterophyllous “aquatic leaves” that appear to represent adaptations to improve the uptake of gases from the water. Intercellular spaces in the stems and leaves were more strongly developed in the submerged seedlings than in non-submerged seedlings with the shoot and leaves in the air. The intercellular spaces appear to be schizogenous aerenchyma that facilitates gas exchange. The growth responses and anatomical responses in stems and leaves to submergence, which were found in M. cajuputi, are commonly known in herbaceous plants with amphibious characteristics, but had not been reported in woody plants. And our results suggest that M. cajuputi adapts to submergence similarly to other amphibious plants, thereby ensuring continuing biomass production.     

Photosynthetic consequences of phenotypic plasticity in response to submergence: Rumex palustris as a case study

Survival and growth of terrestrial plants is negatively affected by complete submergence. This is mainly the result of hampered gas exchange between plants and their environment, since gas diffusion is severely reduced in water compared with air, resulting in O2 deficits which limit aerobic respiration. The continuation of photosynthesis could probably alleviate submergence-stress in terrestrial plants, but its potential under water will be limited as the availability of CO2 is hampered. Several submerged terrestrial plant species, however, express plastic responses of the shoot which may reduce gas diffusion resistance and enhance benefits from underwater photosynthesis. In particular, the plasticity of the flooding-tolerant terrestrial species Rumex palustris turned out to be remarkable, making it a model species suitable for the study of these responses. During submergence, the morphology and anatomy of newly developed leaves changed: 'aquatic' leaves were thinner and had thinner cuticles. As a consequence, internal O2 concentrations and underwater CO2 assimilation rates were higher at the prevailing low CO2 concentrations in water. Compared with heterophyllous amphibious plant species, underwater photosynthesis rates of terrestrial plants may be very limited, but the effects of underwater photosynthesis on underwater survival are impressive. A combination of recently published data allowed quantification of the magnitude of the acclimation response in this species. Gas diffusion resistance in terrestrial leaves underwater was about 15,000 times higher than in air. Strikingly, acclimation to submergence reduced this factor to 400, indicating that acclimated leaves of R. palustris had an approximately 40 times lower gas diffusion resistance than non-acclimated ones.     

Acclimation of photosynthesis to supersaturated CO2 in aquatic plant bicarbonate users

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Photosynthesis of Ulva sp. II. utilization of CO2 and HCO−3 when submerged

The photosynthetic capacity of submerged Ulva sp. when utilizing CO₂ and HCO^?₃ as exogenous carbon forms has been investigated and compared with ambient carbon concentrations in sea water. Saturating concentrations of HCO^{? ₃} and CO₂ were 1200 and 100 μM, respectively at saturating light, and photosynthetic rates under such conditions averaged 700 μmolO₂·gDW^?1 ·h^?1. The HCO^?₃ concentration of sea water (≈2500μM), was thus found to be saturating for photosynthesis of Ulva. At the CO₂ concentration of sea water (≈ 10 μM), the contribution of this carbon form to photosynthesis could be 27% at the most. Under conditions of slow water movement, the relative importance of CO₂ utilization would probably be minimized in favour of HCO^?₃ utilization. It is concluded that HCO^?₃ uptake is not limiting photosynthesis for Ulva under natural conditions.     

Inorganic Carbon Source for Photosynthesis in the Seagrass Thalassia hemprichii (Ehrenb.) Aschers

Photosynthetic carbon uptake of the tropical seagrass Thalassia hemprichii (Ehrenb.) Aschers was studied by several methods. Photosynthesis in buffered seawater in media in the range of pH 6 to pH 9 showed an exponentially increasing rate with decreasing pH, thus indicating that free CO₂ was a photosynthetic substrate. However, these experiments were unable to determine whether photosynthesis at alkaline pH also contained some component due to HCO₃⁻ uptake. This aspect was further investigated by studying photosynthetic rates in a number of media of varying pH (7.8-8.61) and total inorganic carbon (0.75-13.17 millimolar). In these media, photosynthetic rate was correlated with free CO₂ concentration and was independent of the HCO₃⁻ concentration in the medium. Short time-course experiments were conducted during equilibration of free CO₂ and HCO₃⁻ after injection of ¹⁴C labeled solution at acid or alkaline pH. High initial photosynthetic rates were observed when acidic solutions (largely free CO₂) were used but not with alkaline solutions. The concentration of free CO₂ was found to be a limiting factor for photosynthesis in this plant.     

EFFECTS OF CO2 ENRICHMENT ON THE BLOOM‐FORMING CYANOBACTERIUM MICROCYSTIS AERUGINOSA (CYANOPHYCEAE): PHYSIOLOGICAL RESPONSES AND RELATIONSHIPS WITH THE AVAILABILITY OF DISSOLVED INORGANIC CARBON1

Microcystis aeruginosa Kütz. 7820 was cultured at 350 and 700 μL·L ^{? 1} CO₂ to assess the impacts of doubled atmospheric CO₂ concentration on this bloom‐forming cyanobacterium. Doubling of CO₂ concentration in the airflow enhanced its growth by 52%–77%, with pH values decreased and dissolved inorganic carbon (DIC) increased in the medium. Photosynthetic efficiencies and dark respiratory rates expressed per unit chl a tended to increase with the doubling of CO₂. However, saturating irradiances for photosynthesis and light‐saturated photosynthetic rates normalized to cell number tended to decrease with the increase of DIC in the medium. Doubling of CO₂ concentration in the airflow had less effect on DIC‐saturated photosynthetic rates and apparent photosynthetic affinities for DIC. In the exponential phase, CO₂ and HCO₃ ^? levels in the medium were higher than those required to saturate photosynthesis. Cultures with surface aeration were DIC limited in the stationary phase. The rate of CO₂ dissolution into the liquid increased proportionally when CO₂ in air was raised from 350 to 700 μL·L ^{? 1}, thus increasing the availability of DIC in the medium and enhancing the rate of photosynthesis. Doubled CO₂ could enhance CO₂ dissolution, lower pH values, and influence the ionization fractions of various DIC species even when the photosynthesis was not DIC limited. Consequently, HCO₃ ^? concentrations in cultures were significantly higher than in controls, and the photosynthetic energy cost for the operation of CO₂ concentrating mechanism might decrease.     

Alterations in Rubisco activity and in stomatal behavior induce a daily rhythm in photosynthesis of aerial leaves in the amphibious-plant Nuphar lutea

Nuphar lutea is an amphibious plant with submerged and aerial foliage, which raises the question how do both leaf types perform photosynthetically in two different environments. We found that the aerial leaves function like terrestrial sun-leaves in that their photosynthetic capability was high and saturated under high irradiance (ca. 1,500 μmol photons m⁻² s⁻¹). We show that stomatal opening and Rubisco activity in these leaves co-limited photosynthesis at saturating irradiance fluctuating in a daily rhythm. In the morning, sunlight stimulated stomatal opening, Rubisco synthesis, and the neutralization of a night-accumulated Rubisco inhibitor. Consequently, the light-saturated quantum efficiency and rate of photosynthesis increased 10-fold by midday. During the afternoon, gradual closure of the stomata and a decrease in Rubisco content reduced the light-saturated photosynthetic rate. However, at limited irradiance, stomatal behavior and Rubisco content had only a marginal effect on the photosynthetic rate, which did not change during the day. In contrast to the aerial leaves, the photosynthesis rate of the submerged leaves, adapted to a shaded environment, was saturated under lower irradiance. The light-saturated quantum efficiency of these leaves was much lower and did not change during the day. Due to their low photosynthetic affinity for CO₂ (35 μM) and inability to utilize other inorganic carbon species, their photosynthetic rate at air-equilibrated water was CO₂-limited. These results reveal differences in the photosynthetic performance of the two types of Nuphar leaves and unravel how photosynthetic daily rhythm in the aerial leaves is controlled.     

Photosynthetic inorganic carbon utilization and growth of Porphyra linearis (Rhodophyta)

Photosynthetic (oxygen evolution) and growth (biomass increase) responses to ambient pH and inorganic carbon (Ci) supply were determined for Porphyralinearis grown in 0.5 L glass cylinders in the laboratory, or in 40 L fibreglass outdoor tanks with running seawater. While net photosynthetic rates were uniform at pH 6.0–8.0, dropping only at pH 8.7, growth rates were significantly affected by pH levels other than that of seawater (c. pH 8.3). In glass cylinders, weekly growth rates averaged 76% at external pH 8.0, 13% at pH 8.7 and 26% at pH 7.0. Photosynthetic O₂ evolution on a daily basis(i.e. total O₂ evolved during day time less total O₂ consumed during night time) was similar to the growth responses at all experimental pH levels, apparently due to high dark respiration rates measured at acidic pH. Weekly growth rates averaged 53% in algae grown in fibreglass tanks aerated with regular air (360 mg L_-1 CO₂) and 28% in algae grown in tanks aerated with CO₂-enriched air (750 mg L^-1 CO₂). The pH of the seawater medium in which P. linear is was grown increased slightly during the day and only rarely reached 9.0. The pH at the boundary layer of algae submerged in seawater increased in response to light reaching, about pH 8.9 within minutes, or remained unchanged for algae submerged in a CO₂-free artificial sea water medium. Photosynthesis of P. linearissaturated at Ci concentrations of seawater (K_0.5560 μM at pH 8.2) and showed low photosynthetic affinity for CO₂(K_0.5 61 μM) at pH 6.0. It is therefore concluded that P. linearisuses primarily CO₂ with HCO₃ ^- being an alternative source of Ci for photosynthesis. Its fast growth could be related to the enzyme carbonic anhydrase whose activity was detected intra- and extracellularly. This revised version was published online in August 2006 with corrections to the Cover Date.     

CO2 uptake and transport in leaf mesophyll cells

Abstract The acquisition of inorganic carbon for photosynthetic assimilation by leaf mesophyll cells and chloroplasts is discussed with particular reference to membrane permeation of CO₂ and HCO^?₃. Experimental evidence indicates that at the apoplast pH normally experienced by leaf mesophyll cells (pH 6–7) CO₂ is the principal species of inorganic carbon taken up. Uptake of HCO^?₃ may also occur under certain circumstances (i.e. pH 8.5), but its contribution to the net flux of inorganic carbon is small and HCO^?₃ uptake does not function as a CO₂-concentrating mechanism. Similarly, CO₂ rather than HCO^?₃ appears to be the species of inorganic carbon which permeates the chloroplast envelope. In contrast to many C₃ aquatic plants and C₄ plants, C₃ terrestrial plants lack specialized mechanisms for the acquisition and transport of inorganic carbon from the intercellular environment to the site of photosynthetic carboxylation, but rely upon the diffusive uptake of CO₂.     

PHOTOSYNTHETIC CARBON METABOLISM IN ENTEROMORPHA COMPRESSA (CHLOROPHYTA)1

The intertidal macroalga Enteromorpha compressa showed the ability to use HCO₃^? as an exogenous inorganic carbon (Ci) source for photosynthesis. However, although the natural seawater concentration of this carbon form was saturating, additional CO₂ above ambient Ci levels doubled net photosynthetic rates. Therefore, the productivity of this alga, when submerged, is likely to be limited by Ci. When plants were exposed to air, photosynthetic rates saturated at air-levels of CO₂ during mild desiccation. Based on carbon fixing enzyme activities and Ci pulsechase incorporation patterns, it was found that Enteromorpha is a C₃ plant. However, this alga did not show O₂ inhibited photosynthetic rates at natural seawater Ci conditions. It is suggested that such a C₄- like gas exchange response is due to the HCO₃^? utilization system concentrating CO₂ intracellularly, thus alleviating apparent photorespiration.     

<Emphasis Type="Italic">Lotus corniculatus</Emphasis> L. response to carbon dioxide concentration and radiation level variations

Carbon dioxide concentration and light conditions may greatly vary between mountainous and lowland areas determining the photosynthetic performance of plants species. This paper aimed to evaluate the photosynthetic responses of Lotus corniculatus, growing in a mountain and a lowland grassland, under low and high radiation and CO₂ concentration. Net photosynthetic rate, stomatal conductance, transpiration rate, and intercellular CO₂ concentration were measured while the water-use efficiency and the ratio of variable to maximal fluorescence were calculated. Photosynthetic response curves to different levels of radiation and intercellular CO₂ partial pressure were estimated. Our results showed that high radiation and CO₂ concentration enhanced water-use efficiency of plants at both sites, enabling them to use more efficiently the available water reserves under drought conditions. The increase of radiation and CO₂ concentration would enhance the photosynthetic performance of the mountainous population of L. corniculatus, which overall seems to express higher phenotypic plasticity.     

CO2 dynamics along Danish lowland streams: water–air gradients, piston velocities and evasion rates

We measured CO₂ concentration and determined evasion rate and piston velocity across the water–air interface in flow-through chambers at eight stations along two 20 km long streams in agricultural landscapes in Zealand, Denmark. Both streams were 9–18-fold supersaturated in CO₂ with daily means of 240 and 340 μM in January–March and 130 and 180 μM in June–August. Annual CO₂ medians were 212 μM in six other streams and 460 μM in four groundwater wells, while seven lakes were weakly supersaturated (29 μM). Air concentrations immediately above stream surfaces were close to mean atmospheric conditions except during calm summer nights. Piston velocity from 0.4 to 21.6 cm h^?1 was closely related to current velocity permitting calculation of evasion rates for entire streams. CO₂ evasion rates were highest in midstream reaches (170–1,200 mmol m^?2 day^?1) where CO₂-rich soil water entered fast stream flow, while rates were tenfold lower (25–100 mmol m^?2 day^?1) in slow-flowing lower reaches. CO₂ evasion mainly derived from the input of CO₂ in soil water. The variability of CO₂ evasion along the two lowland streams covered much of the range in sub-Arctic and temperate streams reported previously. In budgets for the two stream catchments, loss of carbon from soils via the hydrological cycle was substantial (3.2–5.7 mmol m^?2 day^?1) and dominated by CO₂ consumed to form HCO₃ ^? by mineral dissolution (69–76%) and export of organic carbon (15–23%) relative to dissolved CO₂ export (7–9%).