Long‐term effects of liming in Norwegian softwater lakes: the rise and fall of bulbous rush (Juncus bulbosus) and decline of isoetid vegetation

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Could rising aquatic carbon dioxide concentrations favour the invasion of elodeids in isoetid‐dominated softwater lakes?

1. During the past century, isoetid vegetation types in softwater lakes have often been invaded by faster‐growing elodeids. In these C‐limited systems, this may be related to rising aquatic CO₂ levels. 2. In a laboratory experiment we tested the growth response of two elodeid species, Myriophyllum alterniflorum and Callitriche hamulata, at four different CO₂ levels, ranging from 20 to 230 μmol L⁻¹. In addition, we tested the effect of the nutrient status of the sediment on the growth of C. hamulata at the different CO₂ levels. 3. Shoot and root growth increased with rising CO₂ availability. Irrespective of sediment type, growth was minimal to negative at the lowest CO₂ treatment level, while becoming positive at CO₂ levels around 40–50 μmol L⁻¹. Substantial growth was only obtained when the macrophytes were growing on mesotrophic sediments. The plants reached close to maximal growth at CO₂ levels of c. 100 μmol L⁻¹. 4. Within this experiment, the growth of C. hamulata at CO₂ levels above 90 μmol L⁻¹ may have been limited by N and P availability in both sediment types. The growth rate of M. alterniflorum did not seem to be limited by N and P availability, most likely due to its much higher relative root production. 5. The experimental results show that neither M. alterniflorum nor C. hamulata is able to invade isoetid‐dominated softwater lakes at very low aquatic CO₂ concentrations. However, if the sediments contain enough nutrients, a rise in aquatic CO₂ could allow the invasion of elodeid species leading to the subsequent disappearance of slow‐growing isoetids.     

Competition between isoetids and invading elodeids at different concentrations of aquatic carbon dioxide

1. The growth of submerged macrophytes in softwater lakes is often assumed to be carbon limited. Isoetid species are well adapted to grow at low carbon availability and therefore commonly dominate the submerged macrophyte vegetation in softwater lakes. In many such lakes, however, large‐scale invasions of fast‐growing elodeid species, replacing the isoetid vegetation, have been observed. 2. In a laboratory experiment, we tested how rising aquatic carbon availability, in interaction with different densities of the isoetid Littorella uniflora, affected the growth (and thereby the potential invasion success) of the elodeid Myriophyllum alterniflorum. For this purpose, the growth of M. alterniflorum was determined at a combination of three concentrations of dissolved CO₂ (15, 90, 200 μmol L^?1) and three densities of L. uniflora (0, 553, 1775 plants m^?2). 3. At an ambient CO₂ of 15 μmol L^?1, M. alterniflorum could not sustain itself, whereas at raised CO₂ concentrations, growth became positive and increased with higher CO₂ availability. 4. The presence of L. uniflora, independent of its density, reduced the growth of M. alterniflorum by 50%. Whether this is related to nutrient availability or other factors is not clear. 5. Despite the growth reduction of M. alterniflorum by L. uniflora, at CO₂ ≥90 μmol L^?1, L. uniflora was still overgrown by M. alterniflorum. This may imply that, in field situations, M. alterniflorum can invade softwater systems with relatively high CO₂ availability, even in the presence of dense stands of L. uniflora.     

Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.     

Degraded Softwater Lakes: Possibilities for Restoration

In the Netherlands, the characteristic flora of shallow softwater lakes has declined rapidly as a consequence of eutrophication, alkalization and acidification. The sediment of most lakes has become nutrient rich and anaerobic. We expected that, if a vital seed bank was still present, restoration of the original water quality and sediment conditions would lead to the return of softwater macrophytes. The restoration of 15 degraded, shallow, softwater lakes in the Netherlands was monitored from 1983 to 1998. In eutrophied as well as in acidified lakes, removal of accumulated organic matter from the sediment and shores was followed by rapid recolonization of softwater macrophytes present in the seedbank. After isolation from alkaline water and subsequent mud removal, this recovery was also observed in alkalized lakes. Further development of softwater vegetation correlated strongly with the water quality. When renewed eutrophication was successfully prevented, softwater macrophytes could expand. However, in acidified lakes, Juncus bulbosus and Sphagnum species became dominant after restoration. Liming of an acidified lake was followed by re‐acidification within 3 years. Recolonization by softwater macrophytes was inhibited by high turbidity of the water column and spreading of large helophytes on the shore. As an alternative, controlled inlet of alkaline, nutrient‐poor groundwater was studied in a few lakes. The pH of those lakes increased, the carbon and nitrogen availability decreased and softwater macrophytes returned. Successful restoration has contributed considerably to maintaining biodiversity in softwater lakes in the Netherlands.     

Water‐level fluctuations affect sediment properties,carbon flux and growth of the isoetid Littorella uniflora in oligotrophic lakes

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Elevated alkalinity and sulfate adversely affect the aquatic macrophyte Lobelia dortmanna

The increase in alkalinity and SO₄ ^2? in softwater lakes can negatively affect pristine isoetid population because the increase in alkalinity and SO₄ ^2? can stimulate sediment mineralization and consequently cause anoxia. The consequences of increased sediment mineralization depend on the ability of isoetids such as Lobelia dortmanna to oxidize the rhizosphere via radial O₂ loss. To study how alkalinity and SO₄ ^2? affect the isoetid L. dortmanna, and if negative effects could be alleviated by neighboring plants, three densities of L. dortmanna (“Low”?=?64 plants m^?2, “Medium”?=?256 plants m^?2 and “High”?=?1,024 plants m^?2) were exposed to elevated alkalinity in the water column, or a combination of both elevated alkalinity and SO₄ ^2?, and compared to a control situation. The combination of SO₄ ^2? and alkalinity significantly increased mortality, lowered areal biomass and reduced actual photosynthetic efficiency. Plant density did not significantly alleviate the negative effects caused by SO₄ ^2? and alkalinity. However, actual photosynthetic efficiency was significantly positively correlated to redox potential in the sediment, indicating a positive relationship between plant performance and sediment oxidation. The negative effects on L. dortmanna were probably caused by long periods of tissue anoxia by itself or in combination with H₂S intrusion. Therefore, increase in both SO₄ ^2? and alkalinity surface water can dramatically affect L. dortmanna populations, causing reduction or even disappearance of this icon species.     

Influence of sediment organic enrichment and water alkalinity on growth of aquatic isoetid and elodeid plants

1. Lake eutrophication has increased phytoplankton blooms and sediment organic matter. Among higher plants, small, oligotrophic rosette species (isoetids) have disappeared, while a few tall, eutrophic species (elodeids) may have persisted. Despite recent reduction of nutrient loading in restored lakes, the vegetation has rarely regained its former composition and coverage. Patterns of recovery may depend on local alkalinity because HCO₃^? stimulates photosynthesis of elodeids and not of isoetids. In laboratory growth experiments with two isoetids (Lobelia dortmanna and Littorella uniflora) and two elodeids (Potamogeton crispus and P. perfoliatus), we test whether organic enrichment of lake sediments has a long‐lasting influence by: (i) reducing plant growth because of oxygen stress on plant roots and (ii) inhibiting growth more for isoetids than elodeids. We also test whether (iii) increasing alkalinity (from 0.17 to 3.20 meq. L^?1) enhances growth and reduces inhibition of organic sediment enrichment for elodeids but not for isoetids. 2. In low organic sediments, higher oxygen release from roots of isoetids than elodeids generated oxic conditions to greater sediment depth for Lobelia (4.3 cm) and Littorella (3.0 cm) than for Potamogeton species (1.6–2.2 cm). Sediment oxygen penetration depth fell rapidly to 0.4–1.0 cm for all four species at even modest organic enrichment and oxygen consumption in the sediments. Roots became shorter and isoetid roots became thicker to better supply oxygen to apical meristems. 3. Growth of elodeids was strongly inhibited across all levels of organic enrichment of sediments being eight‐fold lower at the highest enrichment compared to the unenriched control. Leaf biomass of isoetids increased three‐fold by moderate organic enrichment presumably because of greater CO₂ supply from sediments being their main CO₂ source. At higher organic enrichment, isoetid biomass was reduced, leaf chlorophyll declined up to 10‐fold, root length declined from 7 to <2 cm and mortality rose (up to 50%) signalling high plant stress. 4. Lobelia was not affected by HCO₃^? addition in accordance with its use of sediment CO₂. Biomass of elodeids increased severalfold by rising alkalinity from 0.17 to 3.20 meq. L^?1 in accordance with their use of HCO₃^? for photosynthesis, while the negative impact of organically enriched sediments remained. 5. Overall, root development of all four species was so strongly restricted in sediments enriched with labile organic matter that plants if growing in situ may lose root anchorage. Other experiments demonstrate that this risk is enhanced by greater water content and reduced consolidation in organically rich sediments. Therefore, formation of more muddy and oxygen‐demanding sediments during eutrophication will impede plant recovery in restored lakes while high local alkalinity will help elodeid recovery.     

Organic enrichment of sediments reduces arbuscular mycorrhizal fungi in oligotrophic lake plants

1. Arbuscular mycorrhizal fungi (AMF) commonly colonise isoetid species inhabiting oxygenated sediments in oligotrophic lakes but are usually absent in other submerged plants. We hypothesised that organic enrichment of oligotrophic lake sediments reduces AMF colonisation and hyphal growth because of sediment O₂ depletion and low carbon supply from stressed host plants. 2. We added organic matter to sediments inhabited by isoetids and measured pore‐water chemistry (dissolved O₂, inorganic carbon, Fe²⁺ and ), colonisation intensity of roots and hyphal density after 135 days of exposure. 3. Addition of organic matter reduced AMF colonisation of roots of both Lobelia dortmanna and Littorella uniflora, and high additions stressed the plants. Even small additions of organic matter almost stopped AMF colonisation of initially un‐colonised L. uniflora, though without reducing plant growth. Mean hyphal density in sediments was high (6 and 15 m cm^?3) and comparable with that in terrestrial soils (2–40 m cm^?3). Hyphal density was low in the upper 1 cm of isoetid sediments, high in the main root zone between 1 and 8 cm and positively related to root density. Hyphal surface area exceeded root surface area by 1.7–3.2 times. 4. We conclude that AMF efficiently colonise isoetids in oligotrophic sediments and form extensive hyphal networks. Small additions of organic matter to sediments induce sediment anoxia and reduce AMF colonisation of roots but cause no apparent plant stress. High organic addition induces night‐time anoxia in both the sediment and the plant tissue. Tissue anoxia reduces root growth and AMF colonisation, probably because of restricted translocation of nutrient ions and organic solutes between roots and leaves. Isoetids should rely on AMF for P uptake on nutrient‐poor mineral sediments but are capable of growing without AMF on organic sediments.     

Positive feedback favors invasion by a submersed freshwater plant

The submersed macrophyte Utricularia inflata has invaded lakes in northern New York State, thereby threatening native isoetids such as Eriocaulon aquaticum. Isoetids often dominate and modify softwater lakes due to their capacity to oxidize sediment and thus influence solute mobilization. Greenhouse experiments tested the hypotheses that U. inflata invasion could result in higher porewater iron (Fe) concentrations and greater ammonium (NH₄ ⁺) and Fe release from the sediment into the water column, and that this mobilization would stimulate further U. inflata growth. In the first experiment, three levels of U. inflata impact on E. aquaticum were imposed using sediment cores overlain by lake water: E. aquaticum alone, E. aquaticum with a cover of U. inflata, and bare sediment—the latter to simulate local extirpation of the isoetid by the invasive. After 16 weeks, sediment porewater NH₄ ⁺ and total dissolved Fe concentrations were significantly higher (P < 0.05) for the U. inflata and bare sediment treatments. Water column concentrations of these solutes were five-fold higher (P < 0.05) for the bare sediment treatment than E. aquaticum alone, indicating that isoetid extirpation by U. inflata can compromise water quality. A second experiment demonstrated that U. inflata grew faster over bare sediment than over sediment with E. aquaticum (P < 0.05), likely due to greater solute mobilization in the absence of E. aquaticum. Where U. inflata causes a decline of native isoetids in Adirondack Mountain lakes, changes to lake sediment and water chemistry can create a positive feedback loop further escalating the impact of this invasive species.     

Submersed macrophyte growth at low pH: carbon source influences response to dissolved inorganic carbon enrichment

1. Twenty‐one submersed macrophyte species were grown in the greenhouse at low and high dissolved inorganic carbon (DIC) concentrations and at pH 6 to test the hypothesis that growth response to DIC enrichment in the water column would be greatest for species able to use both bicarbonate (HCO) and free CO₂ from the water column, intermediate for species restricted to free CO₂ in the water column, and least for those species able to use free CO₂ from both sediment porewater and the water column. 2. Relative growth rates (RGR) increased significantly with DIC enrichment for 12 of the 21 species. Bicarbonate users as a group averaged a 4.6‐fold increase in RGR with DIC enrichment, significantly greater than the 2.5‐ and 1.7‐fold increases of the water column CO₂ users and sediment CO₂ users, respectively. 3. The ability to use HCO, as measured by a pH drift technique, was positively correlated with the ratio of RGR at high DIC to RGR at low DIC for the 21 species. 4. Bicarbonate users also exhibited significantly lower belowground : total biomass (R : T) ratios than the other two groups, and alone showed a significant increase in R : T ratio with DIC enrichment. 5. Our results indicate that DIC availability may influence community structure in acidic and circumneutral lakes.     

Influence of quantity and lability of sediment organic matter on the biomass of two isoetids,Littorella uniflora and Echinodorus repens

1. Despite real improvement in the water quality of many previously eutrophic lakes, the recovery of submerged vegetation has been poor. This lack of recovery is possibly caused by the accumulation of organic matter on the top layer of the sediment, which is produced under eutrophic conditions. Hence, our objective was to study the combined effects of quantity and lability of sediment organic matter on the biomass of Echinodorus repens and Littorella uniflora and on the force required to uproot plants of L. uniflora. 2. Lake sediments, rich in organic matter, were collected from four lakes, two with healthy populations of isoetids and two from which isoetids had disappeared. The four lake sediments were mixed with sand to prepare a range of experimental sediments that differed in quantity and lability of sediment organic matter. Two isoetid species, E. repens and L. uniflora, were grown in these sediments for 8 weeks. Sediment quality parameters, including elemental composition, nutrient availability and mineralisation rates, were determined on the raw sources of sediment from the lakes. Porewater and surface water were analysed for the chemical composition in all mixtures. At the end of the experiment, plants were harvested and their biomass, tissue nutrient concentration and (for L. uniflora) uprooting force were measured. 3. For both species, all plants survived and showed no signs of stress on all types of sediment. The biomass of E. repens increased as the fraction of organic matter was increased (from 6 to 39% of organic content, depending upon sediment type). However, in some of the sediment types, a higher fraction of organic matter led to a decline in biomass. The biomass of L. uniflora was less responsive to organic content and was decreased significantly only when the least labile sediment source was used to create the gradient of organic matter. The increase in shoot biomass for both species was closely related to higher CO₂ concentrations in the porewater of the sediment. The force required to uproot L. uniflora plants over a range of sediment organic matter fitted a Gaussian model; it reached a maximum at around 15% organic matter and declined significantly above that. 4. Increasing organic matter content of the sediment increased the biomass of isoetid plants, as the positive effects of higher CO₂ production outweighed the negative effects of low oxygen concentration in more (labile) organic sediments. However, sediment organic matter can adversely affect isoetid survival by promoting the uprooting of plants.     

Contribution of Sediment Respiration to Summer CO2 Emission from Low Productive Boreal and Subarctic Lakes

We measured sediment production of carbon dioxide (CO₂) and methane (CH₄) and the net flux of CO₂ across the surfaces of 15 boreal and subarctic lakes of different humic contents. Sediment respiration measurements were made in situ under ambient light conditions. The flux of CO₂ between sediment and water varied between an uptake of 53 and an efflux of 182 mg C m⁻² day⁻¹ from the sediments. The mean respiration rate for sediments in contact with the upper mixed layer (SedR) was positively correlated to dissolved organic carbon (DOC) concentration in the water (r² = 0.61). The net flux of CO₂ across the lake surface [net ecosystem exchange (NEE)] was also closely correlated to DOC concentration in the upper mixed layer (r² = 0.73). The respiration in the water column was generally 10-fold higher per unit lake area compared to sediment respiration. Lakes with DOC concentrations <5.6 mg L⁻¹ had net consumption of CO₂ in the sediments, which we ascribe to benthic primary production. Only lakes with very low DOC concentrations were net autotrophic (<2.6 mg L⁻¹) due to the dominance of dissolved allochthonous organic carbon in the water as an energy source for aquatic organisms. In addition to previous findings of allochthonous organic matter as an important driver of heterotrophic metabolism in the water column of lakes, this study suggests that sediment metabolism is also highly dependent on allochthonous carbon sources.     

Is the tetrasporophyte of Asparagopsis armata (Bonnemaisoniales) limited by inorganic carbon in integrated aquaculture?1

Seaweeds cultivated in traditional land‐based tank systems usually grow under carbon‐limited conditions and consequently have low production rates, if no costly artificial source of inorganic carbon is supplied. In integrated aquaculture, the fish effluents provide an extra source of dissolved inorganic carbon (DIC) to seaweeds due to fish respiration. To evaluate if the tetrasporophyte of Asparagopsis armata (Harv.) F. Schmitz (the Falkenbergia stage) is carbon limited when cultivated with effluents of a fish (Sparus aurata) farm in southern Portugal, we characterized the DIC forms in the water, assessed the species photosynthetic response to the different DIC concentrations and pH values, and inferred for the presence of a carbonic anhydrase (CA)–mediated mechanism. Results showed that A. armata relies mainly on CO₂ to meet photosynthetic needs. Nevertheless, from pH 7.5 upward, the CO₂ supply to RUBISCO seems to derive also from the external dehydration of HCO₃^– mediated by CA. The contribution of this mechanism was essential for A. armata to attain fully saturated O₂‐evolution rates at the natural seawater DIC concentration (2–2.2 mM) and pH values (～8.0). We revealed in this study that seaweeds cultivated in fish‐farm effluents benefit not only from a rich source of ammonia but also from an important and free source of DIC for their photosynthesis. If supplied at relatively high turnover rates (～100 vol · d^?1), fish‐farm effluents provide enough carbon to maximize the photosynthesis and growth even for species with low affinity for HCO₃^–, avoiding the artificial and costly supply of inorganic carbon to seaweed cultures.     

Massive uprooting of Littorella uniflora (L.) Asch. during a storm event and its relation to sediment and plant characteristics

During spring storms massive uprooting of Littorella uniflora occurred in a shallow Dutch softwater lake. The aim of this study was to test whether changes in plant morphology and sediment characteristics could explain the observed phenomenon. Uprooting was expected to occur in plants having a high shoot biomass and low root to shoot ratio (R:S), growing on sediments with a high organic matter content. Normally, uprooting of the relative buoyant L. uniflora is prevented by an extensive root system, expressed as a high R:S. This was studied by sampling floating and still rooted L. uniflora plants, as well as sediment and sediment pore water, along a gradient of increasing sediment organic matter content. Increasing organic matter content was related to increasing L. uniflora shoot biomass and consequently decreasing R:S. Furthermore, the results indicated that uprooting indeed occurred in plants growing on very organic sediments and was related to a low R:S. The increased shoot biomass on more organic sediments could be related to increased sediment pore water total inorganic carbon (TIC; mainly CO₂) availability. Additionally, increased phosphorus availability could also have played a role. The disappearance of L. uniflora might lead to higher nutrient availability in the sediments. It is suggested that this could eventually promote the expansion of faster‐growing macrophytes.     

Photosynthetic Uptake of Free CO2, by the Roots of Lobelia dortmanna

Lobelia dortmanna L. is probably unable to utilize HCO₃^?and uses only free CO₂ for photosynthesis as shown by the Winkler method. A ¹⁴C technique was used to show that if CO₂ is added to the water around roots, photosynthesis increases 3–5 times more than when the corresponding amount of CO₂ is added to the water surroundings the leaves. As the CO₂ content in lakes where Lobelia grows is very limited, Lobelia must absorb CO₂ from the sediment, and the carbon dioxide will have to diffuse from the roots up into the leaves through the intercellular system of the plant. In conjunction with Lobelia's CO₂ uptake from the sediment O₂ is liberated. The plant, therefore, acts as an oxygen pump, which oxidizes the sediment down to a depth of 20 cm.     

CO2 supersaturation along the aquatic conduit in Swedish watersheds as constrained by terrestrial respiration,aquatic respiration and weathering

We tested the hypothesis that CO₂ supersaturation along the aquatic conduit over Sweden can be explained by processes other than aquatic respiration. A first generalized‐additive model (GAM) analysis evaluating the relationships between single water chemistry variables and pCO₂ in lakes and streams revealed that water chemistry variables typical for groundwater input, e.g., dissolved silicate (DSi) and Mg²⁺ had explanatory power similar to total organic carbon (TOC). Further GAM analyses on various lake size classes and stream orders corroborated the slightly higher explanatory power for DSi in lakes and Mg²⁺ for streams compared with TOC. Both DSi and TOC explained 22–46% of the pCO₂ variability in various lake classes (0.01–>100 km²) and Mg²⁺ and TOC explained 11–41% of the pCO₂ variability in the various stream orders. This suggests that aquatic pCO₂ has a strong groundwater signature. Terrestrial respiration is a significant source of the observed supersaturation and we may assume that both terrestrial respiration and aquatic respiration contributed equally to pCO₂ efflux. pCO₂ and TOC concentrations decreased with lake size suggesting that the longer water residence time allow greater equilibration of CO₂ with the atmosphere and in‐lake mineralization of TOC. For streams, we observed a decreasing trend in pCO₂ with stream orders between 3 and 6. We calculated the total CO₂ efflux from all Swedish lakes and streams to be 2.58 Tg C yr^?1. Our analyses also demonstrated that 0.70 Tg C yr^?1 are exported to the ocean by Swedish watersheds as HCO₃^? and CO₃^2? of which about 0.56 Tg C yr^?1 is also a residual from terrestrial respiration and constitute a long‐term sink for atmospheric CO₂. Taking all dissolved inorganic carbon (DIC) fluxes along the aquatic conduit into account will lower the estimated net ecosystem C exchange (NEE) by 2.02 Tg C yr^?1, which corresponds to 10% of the NEE in Sweden.     

Regulation of the induction of bicarbonate uptake by dissolved CO2 in the marine diatom, Phaeodactylum tricornutum

Physiological properties of photosynthesis were determined in the marine diatom, Phaeodactylum tricornutum UTEX640, during acclimation from 5% CO₂ to air and related to H₂CO₃ dissociation kinetics and equilibria in artificial seawater. The concentration of dissolved inorganic carbon at half maximum rate of photosynthesis (K_0·5[DIC]) value in high CO₂‐grown cells was 1009 mmol m ^{? 3} but was reduced three‐fold by the addition of bovine carbonic anhydrase (CA), whereas in air‐grown cells K_0·5[DIC] was 71 mmol m ^{? 3}, irrespective of the presence of CA. The maximum rate of photosynthesis (P_max) values varied between 300 and 500 μ mol O₂ mg Chl ^{? 1} h ^{? 1} regardless of growth pCO₂. Bicarbonate dehydration kinetics in artificial seawater were re‐examined to evaluate the direct HCO₃ ^? uptake as a substrate for photosynthesis. The uncatalysed CO₂ formation rate in artificial seawater of 31·65°/_oo of salinity at pH 8·2 and 25 °C was found to be 0·6 mmol m ^{? 3} min ^{? 1} at 100 mmol m ^{? 3} DIC, which is 53·5 and 7·3 times slower than the rates of photosynthesis exhibited in air‐ and high CO₂‐grown cells, respectively. These data indicate that even high CO₂‐grown cells of P. tricornutum can take up both CO₂ and HCO₃ ^? as substrates for photosynthesis and HCO₃ ^? use improves dramatically when the cells are grown in air. Detailed time courses were obtained of changes in affinity for DIC during the acclimation of high CO₂‐grown cells to air. The development of high‐affinity photosynthesis started after a 2–5 h lag period, followed by a steady increase over the next 15 h. This acclimation time course is the slowest to be described so far. High CO₂‐grown cells were transferred to controlled DIC conditions, at which the concentrations of each DIC species could be defined, and were allowed to acclimate for more than 36 h. The K_0·5[DIC] values in acclimated cells appeared to be correlated only with [CO_2(aq)] in the medium but not to HCO₃ ^? , CO₃^{2 ?} , total [DIC] or the pH of the medium and indicate that the critical signal regulating the affinity of cells for DIC in the marine diatom, P. tricornutum, is [CO_2(aq)] in the medium.     

Utilization of Sediment CO2 by Selected North American Isoetids

The photosynthetic uptake of root-zone CO₂ was determined forEriocaulon septangulare, Gratiola aurea, Isoetes macrospora,Littorella uniflora var. americana and Lobelia dortmanna aspart of a study of the photosynthetic carbon economy of submergedaquatic isoetids. The pH and dissolved inorganic carbon (DIC)of the sediment interstitial water in four Wisconsin lakes reflectedthe water column character, where the DIC increased with depthin the sediment to concentrations five to ten times those ofthe water column. Sediment free CO₂ concentrations were 5–50times those in the water column and were similar at all sites(about 05–1.0mM CO₂ in the root-zone). In ‘pH-drift’studies these plants were unable to take up HCO₂^–. Laboratory determinations of the carbon uptake from the rootand shoot-zones were made for all five species. These experimentsshowed that CO₂ in the root-zone accounted for 65–95 percent of external carbon uptake for the five species. For G.aurea and E. septangulare, root-zone CO₂ was > 85 per centof carbon uptake. Carbon, CO₂, photosynthesis, sediment, isoetid, Eriocaulon septangulare, Gratiola aurea, Isoetes macrospora, Littorella uniflora, Lobelia dortmanna     

Inhibition of Chara vulgaris oospore germination by sulfidic sediments

The germinability of Chara vulgaris oospores collected from the sediments of four Ontario lakes varies considerably, ranging in germination percentage from 7% to 54%. Chemical analysis of the interstitial water of the sediments indicated that oospores with low germination occurred in lakes which have high acid volatile sulfides (H₂S, FeS, HS⁻) and high soluble Fe²⁺. The inhibitory effects of sediment on oospore germination were demonstrated by transplant experiments, and suggested that sulfide was the toxic agent. Exposure of high-germinating sedimentary oospores to free sulfide concentrations greater than 2.0 mM caused a greater than 30% reduction in oospore germination. The presence of sulfide in sediments was shown to result from sulfate reduction by bacteria in sediment pore water of those lakes where oospore viability was lowest. Differences in oospore germination percentage appear, therefore, to be due to the toxicity of H₂S produced in the sediment, either by a direct effect on the oospore, or on the parent plants.