Egeria Densa Planch Research Articles

Inhibitory Effect of Aqueous Extracts from Egeria densa Planch. on Cyanobacteria Microcystis aeruginosa (Kützing) Lemmermann Growth

This study aimed to investigate the allelopathic effect of live Egeria densa Planch. and its aqueous extracts in inhibiting the cyanobacterium Microcystis aeruginosa (Kützing) Lemmermann through a possible growth inhibition pathway. Under coexistence, the presence of live E. densa reduced the growth of M. aeruginosa by 48% compared to when M. aeruginosa was alone. Consequently, we prepared two separate aqueous extracts with distilled water: one from E. densa plants collected from monocultures, and the other from E. densa plants collected from co-cultivation with M. aeruginosa. At a concentration of 0.5 g/L, both extracts successfully suppressed the growth of M. aeruginosa throughout the 5-day exposure period. The extracts obtained from E. densa plants grown in a combined culture with M. aeruginosa showed significant growth-inhibiting capabilities compared to the extracts obtained from E. densa monoculture (p < 0.05). They showed 22% more growth inhibition compared to cultures exposed to monoculture extracts at day 5, indicating that the production of allelochemicals in E. densa was induced during co-existence with cyanobacteria. However, the higher concentrations (2 and 4 g/L) of both extracts did not effectively exhibit a successful inhibitive ability, possibly due to the presence of high nutrient concentrations, specifically PO43−, which may be potentially suppressing the activity of allelochemicals. Further studies are recommended in identifying the specific allelochemicals and exploring their practical implementation in the field.

The effect of lead (Pb) on the growth rates of two aquatic macrophyte species; Limnobium laevigatum (Humb & Bonpl. ex Willd) Heine and Egeria densa Planch. grown in different experimental media

Anthropogenic causes contribute to toxic pollutants in aquatic environments and heavy metal pollution. As a heavy metal, Lead (Pb), is one of the most common causes of pollution in water. Heavy metals must be removed from the aquatic environment because they adversely affect health and all living things in each environment. In this study we aimed to determine the effects of lead (Pb) exposure on the growth rates and biomass of two aquatic macrophyte species, E. densa and L. laevigatum. Plants grown in in two different experimental media. For this purpose, both plants were exposed to 3 different concentrations of lead (1 ppm, 5 ppm, 15 ppm). Samples were measured on the 1st, 4th and 7th days, and the first and last weights of the plants were compared. Bioexperiments were run in triplicate. Positive values were observed in the growth rates of both plants, except for the negative growth rates observed on the 1st day at 1 ppm and 5 ppm lead concentrations in the pond water environment. Both plants showed positive growth in 25% Hoagland medium at all concentrations and days, except for the 1 ppm lead concentration, being observed for E. densa. As a result of our study, lead exposure did not significantly alter the growth rates of E. densa and L. laevigatum in the experimental media used for short-term (up tp 7 days) durations.

Susceptibility of native and invasive submersed plants in New Zealand to florpyrauxifen-benzyl in growth chamber exposure studies

AbstractInvasive aquatic plants constantly threaten freshwaters and associated environs globally. Water resource managers frequently seek new control tactics to combat invasive macrophytes, especially when the availability of herbicides registered for submersed plant control is limited. The synthetic auxin herbicide, florpyrauxifen-benzyl, recently registered (2018) for aquatic site applications in the United States, has shown success in controlling several invasive aquatic weeds. Studies were conducted to evaluate responses of native and invasive submersed plants to florpyrauxifen-benzyl under growth chamber conditions to provide insight on the selectivity of varying herbicide concentrations in New Zealand. Florpyrauxifen-benzyl concentrations evaluated ranged from 0.01 to 107.86 µg ai L−1, encompassing the maximum use concentration (48 µg L−1) for submersed plant applications. Dose–response metrics indicated the New Zealand native species watermilfoil [Myriophyllum triphyllum Orchard] was highly sensitive to florpyrauxifen-benzyl following a 21-d static exposure, having a dry weight 50% effective concentration (EC50) value of 1.2 µg L−1. The invasive species oxygen-weed [Lagarosiphon major (Ridley) Moss] and Canadian waterweed (Elodea canadensis Michx.) were less sensitive, with dry weight EC50 values of 35.4 and >107.86 µg L−1, respectively. Brazilian waterweed (Egeria densa Planch.) was most tolerant to the tested concentrations, as EC50 values were not achieved. Overall, results indicate florpyrauxifen-benzyl demonstrates potential for controlling L. major, with further large-scale screening required to confirm control among field site applications. As the native species (M. triphyllum) was most sensitive to florpyrauxifen-benzyl compared with the invasive plant evaluated (I/N ratio indicated >31.3 times more sensitive), any targeted concentration used for invasive plant control for field applications would likely injure the native M. triphyllum plants. Future studies should investigate additional native and invasive species for management guidance and consider how exposure times influence plant response using similar florpyrauxifen-benzyl concentrations tested in the present study.

Physiological responses and phytoremediation capacity of floating and submerged aquatic macrophytes exposed to ciprofloxacin.

Ciprofloxacin (Cipro) water contamination is a global concern, having reached disturbing concentrations and threatening the aquatic ecosystems. We investigated the physiological responses and Cipro-phytoremediation capacity of one floating (Salvinia molesta D.S. Mitchell) and one submerged (Egeria densa Planch.) species of aquatic macrophytes. The plants were exposed to increased concentrations of Cipro (0, 1, 10, and 100µg.Cipro.L-1) in artificially contaminated water for 96 and 168 h. Although the antibiotic affected the activities of mitochondrial electron transport chain enzymes, the resulting increases in H2O2 concentrations were not associated with oxidative damage or growth reductions, mainly due to the activation of antioxidant systems for both species. In addition to being tolerant to Cipro, after only 96h, plants were able to reclaim more than 58% of that from the media. The phytoremediation capacity did not differ between the species, however, while S. molesta bioaccumulate, E. densa appears to metabolize Cipro in their tissues. Both macrophytes are indicated for Cipro-phytoremediation projects.

Effect of glyphosate-based herbicides on the photosynthetic responses of the macrophyte Egeria densa Planch. from topical lotic ecosystems

Effect of glyphosate-based herbicides on the photosynthetic responses of the macrophyte Egeria densa Planch. from topical lotic ecosystems

A Review of the Biocontrol Programmes Against Aquatic Weeds in South Africa

Biological control (biocontrol) against invasive macrophytes is one of the longest standing programmes in South Africa, initiated in the 1970s against water hyacinth, Pontederia crassipes Mart. (Pontederiaceae). Since then, 15 agent species (13 insects, one mite and one pathogen) have been released against six weeds, most of which are floating macrophytes, with excellent levels of success. The release of the water hyacinth planthopper Megamelus scutellaris Berg (Hemiptera: Delphacidae) in particular, has improved biocontrol prospects for water hyacinth since 2018. In the last decade, however, a new suite of submerged and rooted emergent invasive macrophytes has been targeted. The first release against a submerged macrophyte in South Africa, and the first release against Brazilian waterweed, Egeria densa Planch. (Hydrocharitaceae), anywhere in the world, was achieved with the release of a leafmining fly, Hydrellia egeriae Rodrigues-Júnior, Mathis & Hauser (Diptera: Ephydridae). Yellow flag, Iris pseudacorus L. (Iridaceae) and Mexican waterlily, Nymphaea mexicana Zucc. (Nymphaeaceae), have also been targeted for biocontrol for the first time worldwide, and are in the early stages of agent development. Post-release evaluations, long term monitoring and controlled experiments have highlighted the need for a more holistic approach to managing aquatic invasive plants in South Africa, whose presence is largely driven by eutrophication, resulting in regime shifts between floating and submerged invaded states.

Characterization of the fate and distribution of methoxyfenozide in a water-plant-fish-sediment microcosm using a multimedia fugacity model

Methoxyfenozide is widely employed in paddy land and can flow out into the aquatic environment. The present study combines two approaches, namely, an aquatic microcosm simulation experiment and a multimedia fugacity model, to study the fate and distribution of methoxyfenozide in an aquatic microcosm containing water, sediment, aquatic plants, and zebrafish. The model results indicated that the simulated concentrations agreed with the observed values within one order of magnitude. The degradation rate was less than 7.0% in the three types of aquatic microcosms at 740 h in the model. Methoxyfenozide exhibited very high persistence in the aquatic microcosm. Water played a key role in the fate of methoxyfenozide, acting as a sink in the simulated aquatic environment, followed by sediment. Only approximately 2% of methoxyfenozide entered the organisms (zebrafish and Egeria densa Planch). Methoxyfenozide underwent a significant transport process between the water and sediment. The applications of multimedia fugacity models are useful for understanding the behaviors, fate, and transport of pesticides after their release into the environment and to facilitate risk assessment and management activities.

First record of Egeria densa Planch. (Hydrocharitaceae) for the aquatic Flora of Egypt

Egeria densa Planch. is reported for the first time for the aquatic flora of Egypt. This species is a South American aquatic plant species considered highly invasive outside of its original range. In Egypt, the species may have been introduced as an aquarium plant and has long been wrongly identified as Elodea canadensis. However, it was collected from wild habitat at the northern lake of Egypt (Idku Lake). The article presents data on the comprehensive morphological and palynological features, distribution, ecological notes and conservation status in Egypt of the newly recorded taxon, as well as photographs of herbarium samples, were also provided. This species has not been reported yet in North Africa, except for Algeria. The new record of its presence in Egypt fills the gap in its general distribution in the warm temperate region of the world. Based on the present study Hydrocharitaceae in Egypt, are now represented by 7 genera viz. Halophila, Thalassia, Ottelia, Vallisneria, Elodea, Najas and Egeria. The present data indicate that there are insufficient collections of Egeria densa in Egypt considering that the aquatic flora of Egypt is poorly known.

Invasive Aquatic Plants as Ecosystem Engineers in an Oligo-Mesotrophic Shallow Lake.

Exotic hydrophytes are often considered as aquatic weeds, especially when forming dense mats on an originally poorly colonized environment. While management efforts and research are focused on the control and on the impacts of aquatic weeds on biodiversity, their influence on shallow lakes’ biogeochemical cycles is still unwell explored. The aim of the present study is to understand whether invasive aquatic plants may affect the biogeochemistry of shallow lakes and act as ecosystem engineers. We performed a multi-year investigation (2013–2015) of dissolved biogeochemical parameters in an oligo-mesotrophic shallow lake of south-west of France (Lacanau Lake), where wind-sheltered bays are colonized by dense mats of exotic Egeria densa Planch. and Lagarosiphon major (Ridl.) Moss. We collected seasonal samples at densely vegetated and plant-free areas, in order to extrapolate and quantify the role of the presence of invasive plants on the biogeochemistry, at the macrophyte stand scale and at the lake scale. Results revealed that elevated plant biomass triggers oxygen (O2), dissolved inorganic carbon (DIC) and nitrogen (DIN) stratification, with hypoxia events frequently occurring at the bottom of the water column. Within plants bed, elevated respiration rates generated important amounts of carbon dioxide (CO2), methane (CH4) and ammonium (NH4+). The balance between benthic nutrients regeneration and fixation into biomass results strictly connected to the seasonal lifecycle of the plants. Indeed, during summer, DIC and DIN regenerated from the sediment are quickly fixed into plant biomass and sustain elevated growth rates. On the opposite, in spring and autumn, bacterial and plant respiration overcome nutrients fixation, resulting in an excess of nutrients in the water and in the increase of carbon emission toward the atmosphere. Our study suggests that aquatic weeds may perform as ecosystem engineers, by negatively affecting local oxygenation and by stimulating nutrients regeneration.

Resilience of an aquatic macrophyte to an anthropogenically induced environmental stressor in a Ramsar wetland of southern Chile.

In mid-2004, anthropogenically induced changes in water quality of the Río Cruces wetland, a Ramsar site located in southern Chile (ca. 40°S), enhanced the resuspension of iron-enriched sediments, which were subsequently deposited over the most abundant aquatic macrophyte of the wetland (Egeria densa Planch. 1849). This event triggered the formation of brownish, necrotic patches and increased iron contents in the leaves and stems of E. densa, which contributed to a significant demise of the plant within the wetland. In this study, we estimate the recovery time as a proxy for resilience of this macrophyte at organismal and population levels. Macro- and micro-optical characteristics, as well as iron contents in tissues of E. densa, were documented in four time windows (2004, 2008, 2012, and 2014). In addition, the size of the macrophyte population and its spatial occurrence were monitored from 2008 to 2016 across 36 study sites within the wetland. Our results suggest necrotic patches and high iron contents in E. densa persisted at least until 2008. After 2013, a significant increase in the spatial occurrence of E. densa was observed within the wetland, reaching full recovery of the population during 2015. The health of plant tissues and iron contents in leaves and stems showed recovery period close to 4years, while the recovery of the spatial occurrence of E. densa took approximately 9years. While the monitoring of plant health was not performed on a strict annual basis, the recovery rates estimated here are slower than those described for other macrophytes. This finding might reflect the long-lasting effects of the disturbance from 2004 and the interaction with biotic processes, such as foraging by waterbirds recolonizing the Río Cruces wetland. These results show that full recovery of E. densa was achieved through a cascade of effects starting with abiotic factors (water quality) and passing through physiological and individual levels, to finally reach the population level. A key aspect of this response is the invasive nature of the macrophyte, which likely contributed to its recovery as a consequence of improved water quality. Less successful macrophyte species in other systems may not reach the specific population recovery, and become subdominant species instead, or even be eradicated from the wetland either as the result of herbivory or due to competition with other macrophytes.

Recovery of black-necked swans, macrophytes and water quality in a Ramsar wetland of southern Chile: Assessing resilience following sudden anthropogenic disturbances

In 2004 migration and mortality for unknown reasons of the herbivorous Black necked swan (Cygnus melancorhyphus (Molina, 1782)) occurred within the Río Cruces wetland (southern Chile), a Ramsar Site and nature sanctuary. Before 2004, this wetland hosted the largest breeding population of this water bird in the Neotropic Realm. The concurrent decrease in the spatial occurrence of the aquatic plant Egeria densa Planch. 1849 - the main food source of swans - was proposed as a cause for swan migration and mortality. Additionally, post-mortem analyses carried out on swans during 2004 showed diminished body weight, high iron loads and histopathological abnormalities in their livers, suggesting iron storage disease. Various hypotheses were postulated to describe those changes; the most plausible related to variations in water quality after a pulp mill located upstream the wetland started to operate in February 2004. Those changes cascaded throughout the stands of E. densa whose remnants had high iron contents in their tissues. Here we present results of a long-term monitoring program of the wetland components, which show that swan population abundance, body weights and histological liver conditions recovered to pre-disturbance levels in 2012. The recovery of E. densa and iron content in plants throughout the wetland, also returned to pre-disturbance levels in the same 8-year time period. These results show the temporal scale over which resilience and natural restoring processes occur in wetland ecosystems of temperate regions such as southern Chile.

Evidence for a new regime shift between floating and submerged invasive plant dominance in South Africa

Classical biological control for the management of floating invasive plants has been highly successful in South Africa. However, restoring ecosystem services has been compromised by a new suite of submerged invasive plants. This study proposes that biological control of floating invasive macrophytes acts as a catalyst in a regime shift between floating and submerged invasive plant dominance. Regime shifts are large and sudden changes in the structure and functioning of ecosystems. The proposed shift is driven by the rapid decomposition of floating plants and subsequent increase in availability of nutrients and light. A mesocosm experiment explored the effect of biological control on floating Pistia stratiotes L. (Araceae) upon the growth of invasive submerged Egeria densa Planch. (Hydrocharitaceae), and native submerged plant species of the same family; Lagarosiphon major (Ridl.) Moss (Hydrocharitaceae). The results revealed a cascade effect of biological control of P. stratiotes on the availability of nitrogen, resulting in increased relative growth rates and invasive capacity for E. densa. In contrast, the native L. major could not compete with healthy or damaged P. stratiotes. These findings highlight the vulnerability of South African freshwater systems to submerged plant invasions and demonstrate the importance of a more holistic approach to invasive plant management.

The Brazilian elodea (Egeria densa Planch.) invasion reaches Southeast Europe

The Brazilian elodea (Egeria densa Planch.) invasion reaches Southeast Europe

Effects of wind-induced sediment resuspension on distribution and morphological traits of aquatic weeds in shallow lakes

The spread of invasive aquatic plants (i.e., aquatic weeds) requires a broader knowledge of the factors determining their settlement at the lake scale, in order to improve management practices and biomonitoring. Among hydrodynamic pressures, wave action might influence submerged vegetation distribution in shallow lakes and potentially engender modifications of plant morphological traits. We here report a field survey conducted between 2014 and 2015 in French Atlantic Lakes to assess the spatial distribution and biomass production of two rooted aquatic weeds, Egeria densa Planch. and Lagarosiphon major (Ridl.) Moss, in relation to wind-induced sediment resuspension, water depth and sedimentary features. Moreover, we explored the relation between plant morphological traits and hydrodynamic disturbance under in situ conditions. At the lake scale, E. densa and L. major formed extensive monospecific stands, and occurred in the same areas only at low biomass. Both monospecific and mixed stands preferentially developed in shallow-sheltered or deep-exposed areas. Plant morphological traits showed different patterns in function of sediment resuspension according to the species and the lake. The influence of resuspension was however not systematic, with many cases where morphological traits were not affected at all. Maximum stem length appeared to be the most correlated trait towards sediment resuspension. Moreover, we found a significant correlation between the biomass and the sedimentary organic matter, indicating an interaction between the organic substrate preference of the plants and the effect of the canopy on fine particles sedimentation. On the whole, we highlighted the link between plant distribution, phenotypic plasticity and sediment resuspension, with calm-water zones favouring the settlement of dense vegetated stands. Our study could thus contribute to improve prediction models for identifying suitable areas for potential colonization by aquatic weeds. Further research is needed to better understand the role played by hydraulic forces in structuring the habitats in shallow lakes.

Real-time in-situ Simultaneous Monitoring of Dissolved Oxygen and Materials Movements at a Vicinity of Micrometers from an Aquatic Plant by Combining Deflection of a Probe Beam and Fluorescence Quenching.

It is desirable to be able to monitor the intake or release of the components at different organs of aquatic plants in real time and in-situ. Here, we report a novel optical detection system that allows for real-time in-situ simultaneous monitoring of the dissolved oxygen and material movements at a vicinity of micrometers from the aquatic plant surface. A blue semiconductor diode-laser was used as the light source of both the probe beam and excitation light for fluorescence. The laser light reflected by a dichroic mirror was focused to a vicinity of the plant/water interface in a culture dish by an objective lens. The distance between the focused laser beam and the plant surface was adjusted by an X-Y-Z micro-stage. Deflection of the probe beam was detected by a position sensor, and fluorescence from the vicinity was monitored by a PMT. A commercial fluorescent DO sensor, which simultaneously monitored temperature, was immersed into the culture dish at about 1 cm away from the aquatic plants. A white-light LED was used to illuminate the aquatic plants in the dish in photosynthesis process. A Ru-complex (tris (2,2'-bipyridyl)ruthenium(II) chloride) was used as a fluorescent probe, and Egeria densa Planch. was used as a model aquatic plant. The DO-quenched fluorescence and material movement-induced deflection signals are compared at different distances from the aquatic plant surface. The results show that the optical detection system can monitor DO and the material movements at a vicinity of the aquatic plants not only much more sensitively, but also much more closely to real time than analytical methods that monitor concentration changes at a bulk solution.

High dose of urea enhances the nickel and copper toxicity in Brazilian elodea (Egeria densa Planch. Casp.)

Selected cellular responses for urea (5 mM), Ni (100 μM), and Cu (100 μM) treatments and their combined effects on the leaves of a submerged macrophyte Brazilian elodea (Egeria densa Planch. Casp.) were studied for 4 days. It was observed that the high dose of urea (5 mM) had a significant toxic effect on some physiological and biochemical characteristics of E. densa such as the content of chlorophyll a and the activities of catalase, ascorbate peroxidase, and guaiacol peroxidase. The elevated level of lipid peroxidation was accompanied by the accumulation of proline and ascorbate. A substantial increase in the amount of proline and urease activity was observed in Urea + Ni-treated leaves. The ascorbate content significantly increased in all the treated plants which correlated with ascorbate peroxidase activity. Urea with Cu ions affected the E. densa photosynthetic pigment system and urease activity adversely. The catalase activity was inhibited by the action of all the pollutants studied, especially in Urea + Cu-treated plants, whereas the peroxidases activity (APX and GPX) increased under the Cu and Urea + Cu action. The results presented indicate that the addition of the high dose of urea (5 mM) to the heavy metals studied may enhance the toxic effects in E. densa leaves, especially in the case of copper.

Влияние повышенных концентраций мочевины и тяжелых металлов (Ni2+ и Cu2+) на водные макрофиты (на примере Egeria densa Planch)

Urea and heavy metals (Ni2+ and Cu2+) higher concentration infl uence on the photosynthetic pigments’ content, lipids peroxidation intensity and urease ferments activity in the submerged aquatic macrophyte – Egeria densa Planch. have been studied. It has been shown that addition of metals to the medium with urea tends to strengthen its toxic effect upon plants. It has been found that four-day long hatching of plants in the medium with urea and nickel lead to the oxidative stress development in Canada water weed leaves. Copper concentration equal to 100 mkmol/l appeared to be lethal for plants.

Dissolved gas and nutrient dynamics within an Egeria densa Planch. bed

Submerged aquatic weeds may colonize freshwater shallow lakes by forming dense mats: when this occurs, their invasion interferes with human activities and generates several impacts on aquatic processes. In this paper, we present the preliminary results of an investigation concerning the dissolved gas and nutrient dynamics within a bed of Egeria densa Planch. in a bay of Lacanau Lake (southwestern France). Two 24-h cycles were carried out during summer and autumn 2013, for the measurement of dissolved oxygen (O2), carbon dioxide (CO2), methane (CH4) and dissolved inorganic nitrogen (DIN). Our results show that Egeria densa seasonal dynamics might strongly influence the colonized area: during summer, steady levels of O2 (O2 > 0.15 mm) were coupled to the macrophyte photosynthetic activity, while during autumn respiration processes were mostly pronounced (0.02 mm < O2 < 0.25 mm). High concentrations of CH4 (0.6–53.9 μm) and CO2 (0.03–0.83 mm) were measured within the bed during both sampling campaigns, that indicating a potential contribution to greenhouse gas emissions towards the atmosphere. This study suggests that a shallow environment colonized by an elevated biomass of submerged aquatic weeds is most likely subjected to water hypoxia and nutrient mobilization.

Polluting macrophytes Colombian lake Fúquene used as substrate by edible fungus Pleurotus ostreatus

Invasive aquatic plants from Lake Fúquene (Cundinamarca, Colombia), water hyacinth (Eichhornia crassipes C. Mart.) and Brazilian elodea (Egeria densa Planch.) have been removed mechanically from the lake and can be used for edible mushrooms production. The growth of the oyster mushroom (Pleurotus ostreatus) on these aquatic macrophytes was investigated in order to evaluate the possible use of fruiting bodies and spent biomass in food production for human and animal nutrition, respectively. Treatments included: water hyacinth, Brazilian elodea, sawdust, rice hulls and their combinations, inoculated with P. ostreatus at 3%. Water hyacinth mixed with sawdust stimulated significantly fruiting bodies production (P = 3.3 × 10(-7)) with 71% biological efficacy, followed by water hyacinth with rice husk (55%) and elodea with rice husk (48%), all of these have protein contents between 26 and 47%. Loss of lignin (0.9-21.6%), cellulose (3.7-58.3%) and hemicellulose (1.9-53.8%) and increment in vitro digestibility (16.7-139.3%) and reducing sugars (73.4-838.4%) were observed in most treatments. Treatments spent biomass presented Relative Forage Values (RFV) from 46.1 to 232.4%. The results demonstrated the fungus degrading ability and its potential use in aquatic macrophytes conversion biomass into digestible ruminant feed as added value to the fruiting bodies production for human nutrition.

Hydrodynamically mediated macrophyte silica dynamics

In most aquatic ecosystems, hydrodynamic conditions are a key abiotic factor determining species distributions and abundance of aquatic plants. Resisting stress and keeping an upright position often relies on investment in tissue reinforcement, which is costly to produce. Silica could provide a more economical alternative. Two laboratory experiments were conducted to measure the response of two submerged species, Egeria densa Planch. and Limnophila heterophylla (Roxb.) Benth., to dissolved silicic acid availability and exposure to hydrodynamic stress. The results were verified with a third species in a field study (Nuphar lutea (L.) Smith). Biogenic silica (BSi) concentration in both stems and leaves increases with increasing dissolved silica availability but also with the presence of hydrodynamic stress. We suggest that the inclusion of extra silica enables the plant to alternatively invest its energy in the production of lignin and cellulose. Although we found no significant effects of hydrodynamic stress on cellulose or lignin concentrations either in the laboratory or in the field, BSi was negatively correlated with cellulose concentration and positively correlated with lignin concentration in samples collected in the field study. This implies that the plant might perform with equal energy efficiency in both standing and running water environments. This could provide submerged species with a tool to respond to abiotic factors, to adapt to new ecological conditions and hence potentially colonise new environments.