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.     

Sulfate reduction in lake sediments inhabited by the isoetid macrophytes Littorella uniflora and Isoetes lacustris

Sulfur cycling was examined in sediments inhabited with the isoetids Littorella uniflora and Isoetes lacustris in the oligotrophic soft-water Lake Kalgaard, Denmark. Based on short-term tracer incubations sulfate reduction was measured along a transect from the shore (0.6 m) to profundal sediments (4.6 m). The sulfate reduction rates were low (0.008–0.8 mmol m⁻² d⁻¹) in the sandy shallow sediments with low organic content (<1.3 mmol C g⁻¹ sed DW) and high redox potentials (>100 mV), whereas sulfate reduction was higher at the deeper sites (2.7–4.6 mmol m⁻² d⁻¹) with high organic content (max. 11.5 mmol C g⁻¹ sed DW) and lower redox potentials (<100 mV). High concentrations of dissolved organic carbon (DOC) were found in the low particulate organic sediments (up to 18.4 mM), and most of the DOC pool consisted of acetate (40–77%). Reoxidation of sulfides due to root oxygen release was probably important at all sites and a positive efflux of sulfate across the sediment–water interface was measured, attaining rates (up to 4.8 mmol m⁻² d⁻¹) similar to the sulfate reduction rates. Reoxidation of sulfides was also manifested by high fraction (>80%) of reduced sulfides being accumulated as elemental sulfur or pyrite (chromium reducible sulfur, CRS). The largest pools of CRS were found in high organic sediment with vertical distributions resembling those of the sulfate reduction rates. The overall effect of isoetid growth on sulfur cycling in the rhizosphere is a suppression of sulfate reduction in low organic sediments and the governing of sulfide reoxidation in sediments with higher organic content.     

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.     

Nutrient cycling in shallow,oligotrophic Lake Kvie,Denmark

The importance of isoetids for the exchange of dissolved inorganic nitrogen (DIN) between sediment and water was studied in shallow Lake Kvie, Denmark. Vegetated sediments from the littoral zone (55% of lake area) were compared to unvegetated sediments from the littoral and profundal zone. Clear effects of the isoetids were found on DIN in the porewater. At the vegetated station, NH₄ ⁺ showed the highest concentrations just below the surface (< 40 µM) whereas NO₃ ^- was dominating below 5 cm depth with concentrations up to 100 µM during the spring. The unvegetated littoral sediment showed a distinct change between winter where NH₄ ⁺ dominated and summer where NO₃ ^- was most abundant. NH₄ ⁺ dominated in the profundal sediment and showed increasing concentration with depth. The E_h was high (> 400 mV) in the vegetated sediment, indicating isoetid release of O₂ in the rhizosphere. A low DIN uptake was observed at the vegetated station while, based on porewater data, a diffusive release from the sediment was expected. This difference was due to plant assimilation. In general a release of NH₄ ⁺ and an uptake of NO₃ ^- was seen in all sediments. The denitrification rate calculated from the mass balance for the entire lake was 0.4 mmol m - 2 d^-1 and accounted for removal of 77% of the annual N-input to Lake Kvie.     

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.     

Reduced root anchorage of freshwater plants in sandy sediments enriched with fine organic matter

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

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Effects of arbuscular mycorrhizae on biomass and nutrients in the aquatic plant Littorella uniflora

1. It has been hypothesised that the symbiosis with arbuscular mycorrhizal fungi (AMF) leads to a higher uptake of phosphorus (P) and nitrogen (N) in aquatic plants, but it has never been shown experimentally without the use of fungicides. In particular, the symbiosis may be important for nutrient uptake by isoetids in oligotrophic lakes, where low concentrations of inorganic N and P both in the water and in the sediment limit the growth of plants and where symbiosis facilitates the uptake of nutrients from the sediment. 2. Plants of the isoetid Littorella uniflora were propagated under the sterile conditions without an AMF infection. The plants were then grown for 60 days with and without re‐infection by AMF, and with either high (150 μm ) or low (ambient concentration approximately 15 μm ) CO₂ concentration. 3. The study proved that the symbiosis between AMF and L. uniflora had a positive impact on the retention of N and P in the plants at very low nutrient concentrations in the water and on biomass development. Shoot biomass and standing stocks of both P and N were significantly higher in re‐infected plants. 4. Raised CO₂ concentration resulted in a fivefold increase in hyphal infection, but had no impact on the number of arbuscules and vesicles in the cross sections. There were significantly higher biomass and lower tissue P and N concentrations in the plants from high CO₂ treatments. This resulted in similar standing stocks of P and N in plants from low and high CO₂ treatments. 5. The results from this study showed that the symbiosis between AMF and L. uniflora is an important adaptation enabling isoetids to grow on nutrient‐poor sediments in oligotrophic lakes.     

Distinguishable root plaque on root surface of Potamogeton crispus grown in two sediments with different nutrient status

The properties of plaques were different on the root surface of Potamogeton crispus planted in sediments from two different shallow lakes. Lake Tangxunhu sediment, with low pH, contained low organic matter, whereas Lake Yuehu sediment, with high pH, had high calcium deposits mixed with high organic matter. The contents of mineral elements in sediment of Lake Tangxunhu was lower than that of Lake Yuehu, except for iron (Fe) content, but the contents of mineral elements extracted by sodium dithionite–sodium citrate–sodium bicarbonate (DCB) from root plaques were higher in Lake Tangxunhu than those in Lake Yuehu, except for Fe. These element distributions on P. crispus root plaques were characterized by scanning electron microscope combined with energy-dispersive X-ray spectrometer and were consistent with the contents of mineral elements in sediment. The root plaque of P. crispus planted in Lake Tangxunhu sediment mainly contained silicon (Si) and Fe, and the content of Si was greater than Fe, which may be contributed to the formation of poly-silicic-ferric in the natural conditions. However, the root plaque of P. crispus planted in the sediment with higher calcium content of Lake Yuehu was rich in Fe, Si, phosphorus (P), and calcium (Ca). Due to oxygen secretion by plant roots, the root plaque has more Fe₃(PO₄)₂ and a certain amount of Ca₃(PO₄)₂. The ratio of magnesium (Mn) to Fe extracted by DCB from root plaque in Lake Tangxunhu sediment was 0.031 and 0.010 in Lake Yuehu sediment. In Lake Tangxunhu sediment, lower content of organic matter results in weak reducibility. Enhanced oxidation ability by oxygen secretion of P. crispus root could oxidize low-valent Fe and Mn into iron–manganese oxide, which leads to formation of iron–manganese plaque on the root surface. However, this case is different in Lake Yuehu sediment, where Fe and Mn can be reduced in high organic sediment and low-valent Mn can precipitate in the sediment in which pH is >8. Thus, low-valent Fe in Lake Yuehu sediment moves to the root surface of P. crispus, where it oxidizes into Fe oxide, i.e., Fe plaque.     

Binding of phosphate in sediment accumulation areas of the eastern Gulf of Finland,Baltic Sea

The relationships between P and components binding P were studied by analysing the concentrations of N, P, Fe, Mn, Ca and Al in sediments and pore water along the estuarine transect of the River Neva in August 1995. The high sediment organic matter concentration resulted in low surface redox potential and high pore-water o-P concentration, whereas the abundance of amphipods resulted in high surface redox potentials and low pore-water o-P concentration. However, despite the variation in sediment organic matter and the abundance of amphipods, very reduced conditions and slightly variable concentrations of Tot-P (0.7–1.1 mg g^–1 DW) were observed in the 10–15 cm sediment depth along the estuarine gradient, indicating that the pools of mobile P were largely depleted within the depth of 0–15 cm. Multiple regression analysis demonstrated that organic matter and Tot-Fe concentration of the sediment were closely related to the variation in Tot-P concentration of the sediments (r ² = 0.817, n=32). In addition, the high total Fe:P ratio suggested that there is enough Fe to bind P in sediments along the estuarine gradient. However, low Fe_diss concentrations in the pore water of reduced sediment (redox-potential <–50 mV) indicated efficient precipitation of FeS (FeS and FeS₂), incapable to efficiently bind P. Consequently, the low Fe_diss:o-P ratio (< 1) recorded in pore water in late summer implied that Fe³⁺ oxides formed by diffusing Fe_diss in the oxic zone of the sediments were insufficient to bind the diffusing o-P completely. The measured high o-P concentrations in the near-bottom water are consistent with this conclusion. However, there was enough Fe_diss in pore water to form Fe³⁺ oxides to bind upwards diffusing P in the oxic sediment layer of the innermost Neva estuary and the areas bioturbated by abundant amphipods.     

Oxygen stress and reduced growth of Lobelia dortmanna in sandy lake sediments subject to organic enrichment

1. Lobelia dortmanna is a common representative of the small isoetid plants dominating the vegetation in nutrient‐poor lakes in Europe and North America. Because of large permeable root surfaces and continuous air lacunae Lobelia exchanges the majority of O₂ and CO₂ during photosynthesis across the roots. This leads to profound diel pulses of O₂ and CO₂ in sandy sediments with low microbial O₂ consumption rates. The ready radial root loss of O₂ may, however, make Lobelia very susceptible to more reducing sediments. Therefore, we grew Lobelia for 6 months on natural and organically enriched sandy sediments to test how: (i) root oxygenation influenced degradation of organic matter and depth profiles of N and C; (ii) Lobelia and microbial O₂ consumption rates influenced pool size and depth penetration of O₂ in the sediments; and (iii) sediment enrichment influenced growth and mineral nutrition of Lobelia. 2. Naturally low‐organic sediments (0.32% DW) accumulated organic C and N during the experiment as a result of growth of Lobelia and surface micro‐algae. In contrast, surface layers of enriched sediments (0.58, 0.87 and 2.46% DW) lost organic C and N because of enhanced mineralisation rates because of oxygen availability. In deeper layers of enriched sediments no significant differences in organic C and N pools were found between plant‐covered and plant‐free sediments probably because faster organic degradation because of root oxygenation was balanced by release of organic matter from the plants and because short roots with dense Fe‐Mn coatings in the most enriched sediments constrained O₂ release. 3. Depth‐integrated O₂ pools were much higher in light than darkness, higher in plant‐covered than plant‐free sediments and higher in sandy than in organically enriched sediments. All sediments had a primary O₂ maximum 1–2 mm below the sediment surface in light because of photosynthesis of micro‐algae. Plant‐covered sediments of low organic content (0.32 and 0.58% DW) also had a secondary deep maximum (2–4 cm) because of higher O₂ release from Lobelia roots than microbial O₂ consumption. Nitrification occurred here resulting in depletion of NH and accumulation of NO. In low organic sediments, oxygen pools increased with higher plant biomass both in light and darkness. The deep O₂ and NO₃ maxima disappeared in high organic sediments of greater O₂ consumption rates and smaller O₂ release rates. 4. Lobelia was stressed by increasing O₂ consumption rate of the sediments. Plant weight and leaf number declined twofold and maximum root length declined fourfold suggesting severe problems maintaining sufficient axial O₂ transport to the root tips because of rapid radial O₂ loss. Despite markedly higher nutrient concentrations in the enriched sediments, leaf‐N declined twofold and leaf‐P declined fourfold to growth‐limiting levels. These responses can be explained by constrains on mycorrhisal activity, root metabolism and vascular transport because of O₂ depletion. Management efforts to stop the decline and ensure the recovery of the isoetid vegetation should therefore focus on improving water quality as well as sediment suitability for growth.     

Effects of added organic matter on iron and manganese redox systems in sediment

Addition of five types of organic matter to Lake Washington sediments resulted in release of high concentrations of iron, organic carbon, and manganese into the interstitial water, and caused an increase in observed sediment oxygen consumption rates. The depressed electrode potentials (E_h < —150 mV) that should accompany such reduction processes did not occur, indicating that E_h was being poised by redox systems present in the sediment. Iron redox systems [Fe(OH)₃‐Fe²⁺, Fe₃(OH)₈‐Fe²⁺, and Fe(OH)₃‐Fe₃(OH)₈] were shown to be poising the E_h of control sediments throughout 13 weeks of incubation and dominating the potential of several of the organically amended sediments following the first three weeks of incubation. Depression of calculated iron system E_o values relative to that of the control sediment early in the incubation appeared to be due to the decreased pH and non‐equilibrium conditions in the organic matter‐amended sediment during the first weeks of incubation. Manganese redox systems exerted no discernable impact on the E_h of the sediment.     

Phosphorus dynamics during decomposition of mangrove (Rhizophora apiculata) leaves in sediments

Rhizophora apiculata leaf litter decomposition and the influence of this process on phosphorus (P) dynamics were studied in mangrove and sand flat sediments at the Bangrong mangrove forest, Phuket, Thailand. The remaining P in the mangrove leaf litter increased with time of decomposition to 174% and 220% of the initial amount in the litter in sand flat and mangrove sediment, respectively, although about 50% of the dry weight had been lost. The incorporation of P into the litter was probably associated with humic acids and metal bridging, especially caused by iron (Fe), which also accumulated in considerable amounts in the litter (5-10 times initial concentration). The addition of leaves to the sediment caused increased concentrations of dissolved reactive phosphate (DRP) in the porewater, especially in sand flat sediment. The DRP probably originated from Fe-bound P in the sediment, because decomposition of buried leaf litter caused increased respiration and reduced the redox potential (E_h) in the sediments. Binding of P to refractory organic material and oxidized Fe at the sediment-water interface explains the low release of DRP from the sediment. This mechanism also explains the generally low DRP concentration in the mangrove porewater, the low nutrient content of the R. apiculata leaves, but also the higher total sediment P concentration of the mangrove sediment as compared to sediments outside the mangrove. Both the low release rates for DRP from the sediment and the accumulation of P associated with leaf litter decomposition tend to preserve P in the sediments.     

Distribution and quantitative development of aquatic macrophytes in relation to sediment characteristics in oligotrophic Lake Kalgaard, Denmark

SUMMARY. The distribution and quantitative development of aquatic macrophytes have been studied in oligotrophic Lake Kalgaard, Denmark. The vegetation is dominated by isoetid species, which are widely distributed (about 40% of the lake bottom) compared to emergent and floating-leaved macrophytes (about 4%). Littorella uniflora dominates at depths of 0–2 m and Isoetes lacustris from 2.0 to 4.5m. Within the colonization area the mean midsummer biomass of Littorella is 112g organic dry weight m⁻² and that of Isoetes, 66 gm⁻². The total biomass of these two species constitutes 99% of the biomass of submerged macrophytes.
The perennial Littorella shows only small seasonal biomass variations. The vegetational biomass, the above-ground fraction of the biomass, and the weight of individual plants all increased with the organic content of the sediment at water depths from 0 to 0.75 m. At the same time the interstitial concentrations of carbon dioxide, extractable inorganic nitrogen, and exchangeable inorganic phosphorus increased, thus supporting the hypothesis that an increasing organic content of the sediments at this low level creates a physiologically richer medium for the plants.
The isoetid growth form is discussed in relation to the chemical environment of oligotrophic, softwater lakes.     

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.     

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.     

Influence of outbreak of macroalgal blooms on phosphorus release from the sediments in Swan Lake Wetland,China

Macroalgal blooms have occurred worldwide frequently in coastal areas in recent decades, which dramatically modify phosphorus (P) cycle in water column and the sediments. Rongcheng Swan Lake Wetland, a coastal wetland in China, is suffering from extensive macroalgal blooms. In order to verify the influence of macroalgal growth on sediment P release, the sediments and filamentous Chaetomorpha spp. were incubated in the laboratory to investigate the changes of water quality parameters, P levels in overlying water, and sediments during the growth period. In addition, algal biomass and tissue P concentration were determined. In general, Chaetomorpha biomasses were much higher in high P treatments than in low P treatments. Compared with algae+low P water treatment, the addition of sediments increased the algal growth rate and P accumulation amount. During the algal growth, water pH increased greatly, which showed significant correlation with algal biomass in treatments with high P (P < 0.05). P fractions in the sediments showed that Fe/Al–P and organic P concentrations declined during the algal growth, and great changes were observed in algae+low P water+sediment treatment for both. As a whole, the sediments can supply P for Chaetomorpha growth when water P level was low, and the probable mechanism was the release of Fe/Al–P at high pH condition induced by intensive Chaetomorpha blooms.     

Shading by an invasive macrophyte has cascading effects on sediment chemistry

The submersed freshwater macrophyte Utricularia inflata is a recent invader of Adirondack Mountain lakes (NY, USA). Previous experiments suggested that U. inflata can indirectly change nutrient cycling in Adirondack lake ecosystems by reducing the growth of native isoetid macrophytes, which in turn affects sediment chemistry. A 13-week greenhouse experiment was conducted to test the hypothesis that shading can explain the detrimental effect of U. inflata on the native short-statured isoetid, Eriocaulon aquaticum. Eriocaulon aquaticum has a dense root system that oxidizes sediment by releasing oxygen; it also takes up carbon dioxide from sediment. Growth and asexual reproduction of E. aquaticum grown under shaded conditions was reduced significantly compared to an unshaded control (P < 0.001). Shading resulted in sediment changes: redox potential fell from 216 mV in the absence of shading to 76 mV under four layers of shade cloth (P < 0.0001). Shading also increased the concentration of extractable sediment ammonium (P < 0.01), as well as carbon dioxide concentrations (P < 0.0001) and pH of porewater (P < 0.05). The effect of U. inflata on the native isoetids and consequently on sediment chemistry closely matched the impact of shade cloth with similar light attenuation. Our results indicate that the principal mechanism by which U. inflata affects native isoetids and sediment chemistry is shading.     

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.