An experimental system to study ecophysiological responses of submerged macrophytes to temperature and light

An experimental growth system was devised to study the ecophysiological responses of submerged macrophytes to temperature and light. Using the system, a pilot study was conducted to compare responses to light of three representative species common to the littoral zone of Lake Biwa: Vallisneria asiatica Miki var. biwaensis Miki, which is endemic to Japan, Potamogeton maackianus A. Benn., a dominant submerged macrophyte in Lake Biwa, and Elodea nuttallii (Planch.) St. John, one of the most influential submerged exotic macrophytes naturalized in Japan. Different relative growth rate responses and other growth parameters among the species were observed. We ascertained the experimental system to be useful for the comparison of ecophysiological responses of submerged macrophytes.     

An alternative mechanism for shade adaptation: implication of allometric responses of three submersed macrophytes to water depth

Allometric scaling models describing size-dependent biological relationships are important for understanding the adaptive responses of plants to environmental variation. In this study, allometric analysis was used to investigate the biomass allocation and morphology of three submerged macrophytes (Potamogeton maackianus, Potamogeton malaianus and Vallisneria natans) in response to water depth (1.0 and 2.5?m) in an in situ experiment. The three macrophytes exhibited different allometric strategies associated with distinct adjustments in morphology and biomass allocation in response to varying water depths. In deeper water, after accounting for the effects of plant size, P. maackianus and P. malaianus tended to enhance light harvesting by allocating more biomass to the stem, increasing shoot height and specific leaf area. V. natans tended to allocate more biomass to the leaf than to the basal stem (rosette), showing a higher leaf mass ratio and shoot height in deeper water. The three species decreased biomass allocation to roots as water depth increased. The main effect of water depth treatments was reduced light availability, which induced plastic shoot or leaf elongation. This shows that macrophytes have evolved responses to light limitation similar to those of terrestrial plants.     

Biology and ecology of Brazilian elodea (Egeria densa) and its specific herbivore, Hydrellia sp., in Argentina

Egeria densa (Hydrocharitaceae) is a submerged macrophyte from South America that is a weed in several countries. It crowds out native plants and hinders water use, causing economic and environmental damage. The leafminer fly Hydrellia sp. 1 (Diptera: Ephydridae), was found feeding in E. densa throughout its Argentine distribution, and is currently the only known specialist herbivore of E. densa. It was reared in the laboratory and tested on 25 plant species. This herbivore can cause heavy defoliation in the laboratory and in the field. Hydrellia sp. 1 was found only on E. densa, but in the laboratory it also developed on two other Hydrocharitaceae species in the same family; Egeria naias, and Elodea callitrichoides. Significant oviposition and feeding were only observed on its primary natural host, and to a lesser degree on E. naias. Field studies indicate Hydrellia sp. 1 is present in the field year round, unless the host plant is prostrate for long periods, or covered by floating macrophytes. These results indicate Hydrellia sp. 1 may be a suitable biocontrol candidate for E. densa.     

Effects of Spirogyra arcta on biomass and structure of submerged macrophyte communities

The presence of algae can greatly reduce the amount of light that reaches submerged macrophytes, but few experimental studies have been conducted to examine the effects of algae on biomass and structure of submerged macrophyte communities. We constructed communities with four submerged macrophytes (Hydrilla verticillata, Egeria densa, Ceratophyllum demersum, and Chara vulgaris) in three environments in which 0 (control), 50 and 100% of the water surface was covered by Spirogyra arcta. Compared to the control treatment, the 100% spirogyra treatment decreased biomass of the submerged macrophyte communities and of all the four macrophytes except C. demersum. Compared to the control and 50% treatments, the 100% treatment significantly increased relative abundance of C. demersum and decreased that of E. densa. Therefore, the presence of S. arcta can greatly affect the productivity and alter the structure of submerged macrophyte communities. To restore submerged macrophyte communities in conditions with abundant algae, assembling communities consisting of C. demersum or similar species may be a good practice.     

The effect of spring water on the growth of a submerged macrophyte Egeria densa

We elucidated the effect of spring water on the growth of Egeria densa Planch., a widespread submerged macrophyte in Japan. We observed the longitudinal distributions of physical (water temperature, particle diameter of the bed sediment, sediment layer thickness, etc.), chemical [pH, dissolved oxygen (DO), dissolved inorganic carbon (DIC), phosphate (PO₄-P), total nitrogen (TN), and total phosphorus (TP) content of the sediment, etc.], and biological (species composition, biomass, and growth rate) factors related to E. densa in the Kurohashi River, a spring-fed stream flowing into the Lake Biwa. It was found that E. densa growth rate from summer to autumn was negatively correlated to pH and DO, which implies that the low pH spring water increases the growth rate of the species. The growth rate was also positively correlated to the free carbon dioxide (CO₂) concentration (r = 0.67, p = 0.02). These results indicate that the low pH spring water increases E. densa growth rate by affecting free CO₂ concentration in water.     

Heavy metals contamination and accumulation in submerged macrophytes in an urban river in China

Deteriorating urban water quality has attracted considerable attention in China. We investigated the contamination levels and distribution of heavy metals (As, Cd, Cu, Ni, Pb, and Zn) in Yuxi River water and sediments, and assessed the heavy metal accumulation capability of five species of submerged macrophytes: Vallisneria natans (Lour.) Hara, Potamogeton pectinatus L., Hydrilla verticillata (L. f.) Royle, Myriophyllum spicatum L., and Potamogeton crispus L. Samples were collected from upstream and downstream locations in different season. The results showed that the levels of heavy metals in the downstream areas were higher than in the upstream areas. Heavy metal concentrations in the river water during the dry seasons were higher than those during the rainy seasons, and the opposite results appeared in sediments and submerged macrophytes. In general, the river was slightly contaminated by heavy metals, and the concentrations of Pb and Ni in this river should serve as a warning, while Cd and Zn pollution in the sediments desperately needs to be removed. Furthermore, Potamogeton pectinatus L. showed a higher accumulation capacity for these metals among the five native submerged macrophytes and could be defined as a hyperaccumulator for Cd. Therefore, the potential use of native aquatic plants in contaminated rivers is worth further exploration.     

Prediction of Egeria najas and Egeria densa occurrence in a large subtropical reservoir (Itaipu Reservoir,Brazil-Paraguay)

Incidence data of two native submerged aquatic macrophytes (Egeria najas Planch. and Egeria densa Planch.) were obtained in eight arms of a large (1350 km²) subtropical reservoir (Itaipu Binacional Reservoir, Brazil-Paraguay). Environmental variables were measured simultaneously. Two large-scale surveys in the same localities identified by a global positioning system were carried out in April 1999 (n = 235) and January 2001 (n = 230). Logistic regressions were used to test the effect of environmental variables on the likelihood of E. najas and E. densa presence or absence. The two species were found under different environmental conditions: conductivity, light attenuation coefficient (k) and fetch were, in this order, the most important environmental variables in predicting the probability of occurrence of E. najas, whereas light attenuation coefficient was the main predictor of the probability of occurrence of E. densa. Thus, both species were negatively affected by the light attenuation coefficient. However, this effect was stronger in E. densa. The small area occupied by these species may be accounted for by the permanent high turbidity of Itaipu Reservoir. Additionally, the dominance of E. najas over of E. densa can be explained by the probably higher light requirements of E. densa. In other reservoirs worldwide, with higher water transparency, the opposite is frequently true. Between 1999 and 2001, an episodic water-level drawdown (5 m) caused the disappearance of submerged vegetation. After water-level normalization, previous vegetation presence (in 1999) was an important predictor of the probability of occurrence of E. najas in 2001.     

The effect of different submerged macrophyte species and biomass on sediment resuspension in a shallow freshwater lake

Our study aim was to elucidate the effects of different species of submerged macrophytes and biomass levels on sediment resuspension. For this purpose experiments were conducted in four different enclosures (Potamogeton maackianus enclosure-PE, Vallisneria spinulosa enclosure-VE, manipulated enclosure-ME and aquaculture enclosure-AE). A sediment trap method was employed and the experiments were conducted from summer to winter in a shallow freshwater lake located in central China. A total of 813, 1277, 613 and 693 g DW m⁻² of sediment was resuspended in VE, AE, ME and PE, respectively. Our results showed that P. maackianus was more effective than V. spinulosa in restraining sediment resuspension. Macrophytes reached their maximum effectiveness of reducing resuspension at a certain species-specific biomass threshold above which biomass effects on resuspension were negligible. The threshold biomass was estimated as 300 g m⁻² for P. maackianus. Accordingly, within a lake management and aquaculture aspect, we conclude that as long as biomass does not fall below this threshold its consumption will not influence sediment resuspension. In the mid-lower reaches of the Yangtze River macrophyte coverage protects the lake sediment against adverse effects of monsoon wind; if the vegetation is eroded aquaculture sediment resuspension increases significantly.     

The Structure of a Macroinvertebrate Community in a Northern German Lake Outlet (Lake Belau,Schleswig-Holstein) with Special Emphasis on Abundance,Biomass and Secondary Production

The structure of macroinvertebrate communities was studied at I I sampling sites of the outlet of Lake Belau in the lowlands of northern Germany. To describe the structures of macrobenthic animal communities three different units were examined: abundance, biomass, and secondary production. 112 taxa were collected from the entire stream. The numbers of species ranged from 31 (fine sand) to 70 (submerged macrophytes). For the stream, average macroinvertebrate density was 18.400 ind. M⁻¹. Density was highest at the macrophytes amounting to 35,630 individuals per m², and lowest in the pure sand with only 3,900 ind. M⁻². Average biomass (dry mass) was 194 g DM m⁻² varying from 9.8 (peat) to 381 g DM m⁻² (gravel with mollusk shells near the upstream lake). For the stream, average annual production was 129 g DM m⁻² varying from 15 (peat) to 286 g DM m⁻² (macrophytes). The highest values for each unit were found in stream sections with gravel and submerged macrophytes. Lower values occured in sections that contained peat and sand. Usually, a single structure of the macroinvertebrate community was dominated by less than ten taxa, which varied at each sampling site depending on the units observed.     

Mn accumulation in a submerged plant Egeria densa (Hydrocharitaceae) is mediated by epiphytic bacteria

Many aquatic plants act as biosorbents, removing and recovering metals from the environment. To assess the biosorbent activity of Egeria densa, a submerged freshwater macrophyte, plants were collected monthly from a circular drainage area in Lake Biwa basin and the Mn concentrations of the plants were analysed. Mn concentrations in these plants were generally above those of terrestrial hyperaccumulators, and were markedly higher in spring and summer than in autumn. Mn concentrations were much lower in plants incubated in hydroponic medium at various pH levels with and without Mn supplementation than in field‐collected plants. The precipitation of Mn oxides on the leaves was determined by variable pressure scanning electron microscopy‐energy dispersive X‐ray analysis and Leucoberbelin blue staining. Several strains of epiphytic bacteria were isolated from the field‐collected E. densa plants, with many of these strains, including those of the genera Acidovorax, Comamonas, Pseudomonas and Rhizobium, found to have Mn‐oxidizing activity. High Mn concentrations in E. densa were mediated by the production of biogenic Mn oxide in biofilms on leaf surfaces. These findings provide new insights into plant epidermal bacterial flora that affect metal accumulation in plants and suggest that these aquatic plants may have use in Mn phytomining.     

Adaptation of submerged macrophytes to both water depth and flood intensity as revealed by their mechanical resistance

Little is known about the mechanical resistance response of submerged macrophytes to floods. An experiment was conducted to investigate the plant growth, root anchorage strength, and stem tensile properties of five submerged macrophytes under three initial water levels (1.0, 2.5, and 4.0 m) with four water level fluctuation speeds (0, 5, 15, and 25 cm d⁻¹). Our results demonstrate that the biomass, relative growth rate, root anchorage strength, and stem tensile properties of the five species decreased with increasing initial water level, suggesting that deep water can inhibit plant growth and decrease their mechanical resistance. Floods weakened the stem tensile properties and strengthened the root performances of Myriophyllum spicatum, Hydrilla verticillata, and Potamogeton malaianus in shallow water. However, floods induced opposite mechanical resistance responses from plants in deep water, indicating a possible trade-off between stem breakage and uprooting under flooding conditions. M. spicatum, Ceratophyllum demersum, and P. malaianus were more tolerant of deep water and flood intensity than Potamogeton maackianus and H. verticillata, as indicated by their larger biomass, plant heights, stem tensile properties, and root anchorage strength. This is the first article that mechanically explains the competitive capability and survival potential of submerged macrophytes to water depth and flood intensity.     

Biological control of phytoplankton by the subtropical submerged macrophytes Egeria densa and Potamogeton illinoensis: a mesocosm study

1. In temperate regions, submerged macrophytes can hamper phytoplankton blooms. Such an effect could arise directly, for instance via allelopathy, or indirectly, via competition for nutrients or the positive interaction between submerged macrophytes and zooplankton grazing. However, there is some evidence that the positive interaction between submerged macrophytes and zooplankton grazing is less marked in warmer regions, where the interaction is less well studied, and that negative effects of higher water plants on phytoplankton biomass are weaker. 2. We carried out two consecutive mesocosm experiments in Uruguay (subtropical South America) to study the effects of two common submerged macrophytes from this region (Egeria densa and Potamogeton illinoensis) on phytoplankton biomass, in the absence of zooplankton grazing. We compared phytoplankton development between different macrophyte treatments (no macrophytes, artificial macrophytes, real Egeria and real Potamogeton). We used artificial macrophytes to differentiate between physical effects (i.e. shading, sedimentation and competition with periphyton) and biological effects (i.e. nutrient competition and allelopathy). 3. In Experiment 1, we found no evidence for physical effects of macrophytes on phytoplankton biomass, but both macrophyte species seemed to exert strong biological effects on phytoplankton biomass. Only Egeria affected phytoplankton community structure, particularly tempering the dominance of Scenedesmus. Nutrient addition assays revealed that only Egeria suppressed phytoplankton through nutrient competition. 4. We performed a second mesocosm experiment with the same design, but applying saturating nutrient conditions as a way of excluding the effects of competition for nutrients. This experiment showed that both macrophytes were still able to suppress phytoplankton through biological mechanisms, providing evidence for allelopathic effects. Our results indicate that both common macrophytes are able to keep phytoplankton biomass low, even in the absence of zooplankton grazing.     

The species composition,occurrence and temporal stability of submerged aquatic macrophyte patches along the main channel border of pool 5A,Upper Mississippi River

Species composition, relative abundance, distribution and physical habitat associations of submerged aquatic macrophytes in the main channel border (MCB) habitat of Pool 5A, Upper Mississippi River (UMR) were investigated during the summers of 1980 and 1983. The submerged aquatic macrophytes in Pool .5A MCB were a small and stable component of the river ecosystem. Submerged plants occurred primarily in small, monospecific clumps. Clumps in close proximity to each other formed plant patches. Plant patches were stable in location and number between 1980 and 1983; 82.5% of the patches first observed in 1980 were present in 1983. Submerged macrophytes covered about 10–12 ha of the 201 ha MCB in Pool 5A. Submerged plants were most common in the lower two-thirds of the pool. Ten species of aquatic macrophytes occurred on rock channel-training structures and eleven occurred on non-rock substrates in the MCB. The most common submerged plants, in order of abundance, were Vallisneria americana Michx., Heteranthra dubia Jacq., Potamogeton pectinatus L., Ceratophyllum demersum L. and Potamogeton americanus C. & S.     

Snail communities increase submerged macrophyte growth by grazing epiphytic algae and phytoplankton in a mesocosm experiment

The relationships between producers (e.g., macrophytes, phytoplankton and epiphytic algae) and snails play an important role in maintaining the function and stability of shallow ecosystems. Complex relationships exist among macrophytes, epiphytic algae, phytoplankton, and snails. We studied the effects of snail communities (consisting of Radix swinhoei, Hippeutis cantori, Bellamya aeruginosa, and Parafossarulus striatulus) on the biomass of phytoplankton and epiphytic algae as well as on the growth of three species of submerged macrophytes (Hydrilla verticillata, Vallisneria natans, and one exotic submerged plant, Elodea nuttallii) in a 90‐day outdoor mesocosm experiment conducted on the shore of subtropical Lake Liangzihu, China. A structural equation model showed that the snail communities affected the submerged macrophytes by grazing phytoplankton and epiphytic algae (reduction in phytoplankton Chl‐a and epiphytic algal abundance), enhancing the biomass of submerged macrophytes. Highly branched macrophytes with high surfaces and morphologies and many microhabitats supported the most snails and epiphytic algae (the biomass of the snail communities and epiphytic algae on H. verticillata was greater than that on V. natans), and snails preferred to feed on native plants. Competition drove the snails to change their grazing preferences to achieve coexistence.     

Temporal changes and vertical distribution of macrophytes in Lake Kawaguchi

 The distribution of macrophytes in Lake Kawaguchi, Japan, was surveyed in August 1999 using a sampling anchor from a boat. The survey revealed that the present aquatic vegetation was composed of 17 submerged species, and no floating-leaved plants were present. The diversity of submerged plants and their vertical growth limits decreased progressively from the west to the east end of the lake. At the time of the survey, the dominant species was Elodea nuttallii (Planch.) St. John, which had once grown explosively in the mid-1980s in the lake, and except for Vallisneria asiatica Miki, all of the species commonly observed in the lake seem to have diminished their growth greatly. Among them, Potamogeton compressus L., which was the most dominant species in the lake before the invasion of Elodea nuttallii, had decreased most severely. It is suggested that the present state of aquatic vegetation in this lake was much influenced by the persistent growth of Elodea nuttallii as well as by environmental factors. Received: August 7, 2001 / Accepted: February 9, 2002     

Biomass distribution,growth and decay of Egeria densa in a tropical high-mountain reservoir (NEUSA,Colombia)

In tropical settings, freshwater macrophytes can reach abundant biomasses. The combination of high temperatures, high irradiances and a continuous growing season are commonly held responsible for such prolific growth. Most research in tropical systems has been done in low altitudes. High-altitude systems, though, are common settings for macrophytes in the tropics, and constitute a unique environment as temperatures can reach values comparable with sub-tropical winter ranges on a daily basis. We sought to study the biomass dynamics, and distribution of Egeria densa P., a dominant submerged weed in Neusa Reservoir (Colombian Andes, 2970 m.a.s.l.). The specific objectives were: (1) to assess the spatial distribution of standing biomass of E. densa in relation to some characteristics of the littoral zone, (2) to study its growth and decay rates as a function of water depth and (3) to compare standing biomass, growth and decay rates in Neusa with those found for other laboratory and natural systems.     

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

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

Lacunar gas discharge: a valid estimate of photosynthetic rates in submerged macrophytes?

Abstract Photosynthetic oxygen produced by the submerged aquatic plant Egeria densa Planch, is partitioned into lacunar gas spaces and to the external medium. Despite an apparently linear relationship between volume of lacunar gas discharge and dissolved oxygen release under certain conditions, the partitioning is, as expected from theoretical considerations, not necessarily constant in different conditions. On this basis we believe that lacunar gas discharge is an inconsistent and therefore unreliable method of measuring photosynthesis in submerged macrophytes.     

Oxygen diffusion and dark respiration in aquatic macrophytes

Abstract. The supply of oxygen to respiring shoot tissue was investigated for three submerged macrophytes (Potamogeton crispus L., Egeria densa Planch, and Myriophyllum triphyllum Orchard). For all species, the response of oxygen uptake rates to the external O₂ concentration was a rectangular hyperbola over the range 0–5.0 × 10^?3m³ m^?3. However, the response pattern for material with water-infiltrated lacunar airspaces was non-hyperbolic over this range. The change in response was interpreted as an increased substrate (O₂) limitation, resulting from lower radial diffusion rates within the infiltrated material. Neither the uninfiltrated nor the infiltrated responses obeyed the linear and logarithmic formulae of the type observed for submerged macrophytes by earlier authors. These results suggest that the responses observed are affected by factors such as water velocity, internal restrictions to diffusion and the range of oxygen tensions investigated. Therefore, it is unlikely that one response formula can adequately account for the effects of oxygen concentration on submerged macrophyte oxygen uptake. The lacunar airspaces also represent a possible oxygen source for dark respiration. The consumption of oxygen from the airspaces was investigated by displacing the gas from the lacunae and measuring the subsequent increase in the rate of oxygen assimilation from the external liquid. Approximately 30% of the oxygen consumed by E. densa and P. crispus, and more than 40% of that consumed by M. triphyllum, was derived from the lacunar system. This O₂ supply is a consequence of the higher oxygen concentration in the lacunae than in the external medium, due to the low solubility of oxygen in water. Storage of photosynthetically-produced oxygen in the lacunae could not be identified during a light/dark transient, due to rate changes caused by the effects of light on the respiratory metabolism. However, O₂ partial pressure gradients artificially set up between the lacunae and water equilibrated within an hour, suggesting that excess oxygen would be lost to the water within this time.     

Community collocation of four submerged macrophytes on two kinds of sediments in Lake Taihu, China

To explore a method for rapid restoration and artificial regulation of submerged macrophytes in large-scale restoration of eutrophic lakes, the succession and the biodiversity changes of four communities composed of four native, common submerged macrophytes, Hydrilla verticillata, Potamogeton malaianus, Vallisneria spiralis and Najas marina, on two kinds of sediments were investigated. Under low light intensity (reduced by 99%), the plant biomass changed with seasonal changes, plant competition, and environmental stress. The competitive capability for light differed in the four species due to different shoot height and tiller number. After 405 days of transplantation, H. verticillata became dominant in all communities. The biomass of H. verticillata, with strong ability to endure low water light environment, accounted for more than 90% of the total community biomass, and P. malaianus had only weak growth, while V. spiralis and N. marina almost disappeared. Based on livability and biomass of submerged macrophytes on two sediment types, brown clay sediment appeared to be more favorable for the settlement of the plants, while fertile sludge sediment was suitable for vegetative growth. In conclusion, the improvement of habitats and the selection of appropriate plant species are of the greatest importance for ecological restoration of the aquatic ecosystem.