Effect of light and nutrients on interspecific interactions between submerged macrophytes:implications for restoration of multispecies aquatic vegetation in eutrophic lakes

Constructing multispecies submerged vegetation systems and maintaining stable seasonal succession is crucial for restoring shallow eutrophic lakes.However,little is known about the interactions between successional and existing species of different growth forms,particularly under the low light and high nutrient conditions of eutrophic lakes.We measured the functional traits of mature Vallisneria natans(Lour.)Hara plants and Potamogeton crispus L.shoots in monoculture and mosaic patterns under different light and nutrient conditions.The effect of light on functional traits of the submerged macrophyte species was more significant than that of nutrients,but the reverse was true for P.crispus biomass allocation.Moreover,interspecific interactions affected only the submerged macrophytes under the low light condition and varied with species.Specifically,the interaction of P.crispus to V.natans was biased towards competition,while the interaction of V.natans to P.crispus was converted from facilitation to competition by eutrophication,particularly in the homogenous mosaic growth pattern.This study demonstrates that sufficient light is a prerequisite and patch planting is an effective means to form a multispecies submerged vegetation system.In addition,we emphasize that the coexistence of eutrophication and low light will likely result in a competition between submerged macrophytes thus simplifying the vegetation,even if their growth forms and growing seasons are different.These findings help explain the collapse of multispecies submerged vegetation and guide the restoration of aquatic plants in eutrophic lakes.

Temporal characteristics of algae-denitrifying bacteria co-occurrence patterns and denitrifier assembly in epiphytic biofilms on submerged macrophytes in Caohai Lake,SW China

Denitrifying bacteria in epiphytic biofilms play a crucial role in nitrogen cycle in aquatic habitats.However,little is known about the connection between algae and denitrifying bacteria and their assembly processes in epiphytic biofilms.Epiphytic biofilms were collected from submerged macrophytes(Patamogeton lucens and Najas marina L.)in the Caohai Lake,Guizhou,SW China,from July to November 2020 to:(1)investigate the impact of abiotic and biotic variables on denitrifying bacterial communities;(2)investigate the temporal variation of the algae-denitrifying bacteria co-occurrence networks;and(3)determine the contribution of deterministic and stochastic processes to the formation of denitrifying bacterial communities.Abiotic and biotic factors influenced the variation in the denitrifying bacterial community,as shown in the Mantel test.The co-occurrence network analysis unveiled intricate interactions among algae to denitrifying bacteria.Denitrifying bacterial community co-occurrence network complexity(larger average degrees representing stronger network complexity)increased continuously from July to September and decreased in October before increasing in November.The co-occurrence network complexity of the algae and nirS-encoding denitrifying bacteria tended to increase from July to November.The co-occurrence network complexity of the algal and denitrifying bacterial communities was modified by ammonia nitrogen(NH_(4)^(+)-N)and total phosphorus(TP),pH,and water temperature(WT),according to the ordinary least-squares(OLS)model.The modified stochasticity ratio(MST)results reveal that deterministic selection dominated the assembly of denitrifying bacterial communities.The influence of environmental variables to denitrifying bacterial communities,as well as characteristics of algal-bacterial co-occurrence networks and the assembly process of denitrifying bacterial communities,were discovered in epiphytic biofilms in this study.The findings could aid in the appropriate understanding and use of epiphytic biofilms denitrification function,as well as the enhancement of water quality.

Epiphytic bacterial communities on two common submerged macrophytes in Taihu Lake: diversity and host-specificity

Leaves of terrestrial and aquatic plants are home to a wide diversity of bacterial species. However, the diversity and variability of epiphytic bacteria on their submerged plant hosts remains poorly understood. We investigated the diversity and composition of epiphytic bacteria from two common submerged macrophytes: Vallisneria natans and Hydrilla verticillata in Taihu Lake, Jiangsu, China, using methods of terminal restriction fragment length polymorphisms (T-RFLP) and clone library analyses targeted at bacterial 16S rRNA genes. The results show that: (1) the libraries of the two waterweeds contain wide phylogenetic distribution of bacteria, and that the sequences of the two libraries can be separated into 93 OTUs (at 97% similar value); (2) Betaproteobacteria, including Burkholderiales, was the most abundant bacterial group on both plants. Cyanobacteria and Gammaproteobacteria were the second largest groups on V. natans and H. verticillata, respectively. Both clone libraries included some sequences related to those of methanotrophs and nitrogen-fixing bacteria; (3) Cluster analysis of the T-RFLP profiles showed two distinct clusters corresponding to the two plant populations. Both ANOSIM of the T-RFLP data and Libshuff analysis of the two clone libraries indicated a significant difference in epiphytic bacterial communities between the two plants. Therefore, the epiphytic bacterial communities on submerged macrophytes appear to be diverse and host-specific, which may aid in understanding the ecological functions of submerged macrophytes in general.

The response of structure and nitrogen removal function of the biofilm on submerged macrophytes to high ammonium in constructed wetlands

The ammonium exceedance discharge from sewage treatment plants has a great risk to the stable operation of subsequent constructed wetlands(CWs).The effects of high ammonium shocks on submerged macrophytes and epiphytic biofilms on the leaves of submerged macrophytes in CWs were rarely mentioned in previous studies.In this paper,the 16S rRNA sequencing method was used to investigate the variation of the microbial communities in biofilms on the leaves of Vallisneria natans plants while the growth characteristics of V.natans plants were measured at different initial ammonium concentrations.The results demonstrated that the total chlorophyll and soluble sugar synthesis of V.natans plants decreased by 51.45%and 57.16%,respectively,and malondialdehyde content increased threefold after8 days if the initial NH_(4)^(+)-N concentration was more than 5 mg/L.Algal density,bacterial quantity,dissolved oxygen,and pH increased with high ammonium shocks.The average removal efficiencies of total nitrogen and NH_(4)^(+)-N reached 73.26%and 83.94%,respectively.The heat map and relative abundance analysis represented that the relative abundances of phyla Proteobacteria,Cyanobacteria,and Bacteroidetes increased.The numbers of autotrophic nitrifiers and heterotrophic nitrification aerobic denitrification(HNAD)bacteria expanded in biofilms.In particular,HNAD bacteria of Flavobacterium,Hydrogenophaga,Acidovorax,Acinetobacter,Pseudomonas,Aeromonas,and Azospira had higher abundances than autotrophic nitrifiers because there were organic matters secreted from declining leaves of V.natans plants.The analysis of the nitrogen metabolic pathway showed aerobic denitrification was the main nitrogen removal pathway.Thus,the nitrification and denitrification bacterial communities increased in epiphytic biofilms on submerged macrophytes in constructed wetlands while submerged macrophytes declined under ammonium shock loading.

Fluorescence Properties of Submerged Macrophytes in Nanjishan Wetland,Southern Poyang Lake

We used a FluorCam portable chlorophyll fluorescence imaging system to measure QY-max （the maximum light quantum yield, Fv/Fm, the largest light quantum efficiency of PS Ⅱ） of submerged plants in wetlands of Baisha Lake and Changhu Lake, Jiangxi Nanjishan Wetland National Nature Reserve, in winter 2013. Specifically, we measuredΦPS Ⅱ （PS Ⅱ actual quantum efficiency）, qP （photochemical quenching） and corresponding fluorescence images. Using the visual method and sampling sites method to obtain coverage, richness and abundance of submerged plants, and determined nutrient levels in water. The results show that the QY-max ofHydrilla verticillata andVallisneria natans in Baisha Lake ranged from 0.48 to 0.68 and 0.52 to 0.71, respectively; theΦPS Ⅱ of these two species ranged from 0.32 to 0.58 and 0.20 to 0.46, respectively. The two plants had similar photosynthetic efifciency. The QY-max ofNymphoides peltatum andV. natans in Changhu Lake ranged from 0.66 to 0.77 and 0.19 to 0.68, respectively; theΦPS Ⅱ of these two species ranged from 0.26 to 0.48 and 0.22 to 0.43, respectively. The observed higher photosynthetic efifciency of N peltatum suggests it is more likely to become the dominant species. In Baisha Lake, the frequency of occurrence of plants was：H. verticilata, 90%;V. natans, 93.3%;Najas minor26.7%, andPotamogeton francheti10%. In Changhu Lake, the frequency of N. pel-tatum was 86.7%,V. natans was 16.7%, andN. minor was 56.7%. The overal frequency of submerged plants living in Baisha Lake was much higher than that of submerged plants living in Changhu Lake, with different species dominating the two lakes. According to comprehensive analysis and comparison of trophic levels, biodiversity and photosynthetic fluorescence characteristics in the two lakes, eutrophication of Baisha Lake was higher than for Changhu Lake; andH. verticillata andV. natans were the dominant species, with similar photosynthetic activity. Conversely, in Changhu Lake,N. peltatum andV. natans were the dominant species, but the photosynthetic activity of N. peltatum was higher thanV. natans. Differences in eutrophication levels in different water bodies in the Nanjishan Wetland and differing ecological niches of submerged plant species are characteristic of this system.

EFFECTS OF SUBMERGED MACROPHYTES ON SEDIMENT SUSPENSION AND NH_4-N RELEASE UNDER HYDRODYNAMIC CONDITIONS

Aquatic vegetation affects sediment suspension and nutrient release by changing the flow structure. Experiments on the influence of submerged vegetation on flow structure, sediment suspension, and NH4-N release were carried in a flume with natural submerged vegetation. Turbulence characteristics in the vegetation section were measured using a three-dimensional acoustic Doppler velocimeter. The effects of submerged vegetation on bed shear stress （ τb ）, sediment suspension, and NH4-N release were analyzed. Results show that with vegetation, bed shear stress is reduced by about 20% - 80%, which, in turn, reduces sediment suspension. The impact of submerged vegetation on sediment suspension and NH4-N release should be considered along with flow intensity. When the flow Reynolds number is relatively small, the submerged vegetation is quite capable of inhibiting sediment suspension and reducing NH4-N release, but when the Reynolds number reaches a certain value, the presence of aquatic plants exacerbates sediment suspension and promotes NH4-N release. Results also reveal that a highly significant positive correlation exists between NH4-N concentration and water turbidity in both vegetated and non-vegetated channels.

The effects of biodiversity gradient on plant mass and metabolism of individual submerged macrophytes

Background:The effects of biodiversity on community function and services are frequently studied in the history of ecology,while the response of individual species to biodiversity remains great elusive.In this study,we determined the biodiversity effects on community productivity as well as species level plant mass and carbon(C)and nitrogen(N)metabolism of eight submerged plants.These macrophytes in Lake Erhai were selected and planted in a water depth of one meter along a diversity gradient of 1,2,4 or 8 species.Then,the community productivity or species level plant mass,soluble protein,free amino acid and soluble carbohydrate were correlated to species richness to determine the biodiversity effects on community and single species.Results:The results showed that the community level biomass was positively correlated to plant species richness although the species level plant mass of individual species responded differently to the overall plant species richness.Namely,only one plant mass positively correlated to species richness and the others decreased or showed no significant correlation with the increase of species richness.The soluble proteins of most macrophytes were positively correlated to species richness;however,both the free amino acid and soluble carbohydrate of the plants were negatively or not significantly correlated to species richness.Conclusions:These results indicated that the selection effects might dominate in our aquatic communities and the negative impacts of biodiversity on C and N metabolism of the macrophytes increased with the increase of species richness,which might result from the strong competition among the studied species.The biodiversity effects on the plant mass,and C and N metabolism of individual submerged species were first reported in this study,while more such field and control experiments deserve further research.

Photosynthesis of Submerged and Surface Leaves of the Dwarf Water Lily(Nymphoides aquatica)Using PAM Fluorometry

Dwarf Water Lilies Nymphoides aquatica(J.F.Gmel)Kuntze have floating and submerged leaves.Some submerged aquatic vascular plants have a form of CAM(Crassulacean Acid Metabolism)called Submerged Aquatic Macrophyte(SAM)metabolism.Blue-diode based PAM technology was used to measure the Photosynthetic Oxygen Evolution Rate(POER:1O_(2)≡4e^(-)).Optimum Irradiance(E_(opt)),maximum POER(POER_(max))and quantum efficiency(α_(0))all vary on a diurnal cycle.The shape of the POER vs.E curves is different in seedling,submerged and surface leaves.Both E_(opt) and POER_(max) are very low in seedling leaves(E_(opt)≈104μmol photon m^(-2) s^(-1),PPFD;POER_(max)≈4.95µmol O_(2) g^(-1) Chl a s^(-1)),intermediate in mature submerged leaves(E_(opt)≈419µmol photon m^(-2) s^(-1) PPFD,POER_(max)≈38.1µmol O_(2) g^(-1) Chl a s^(-1))and very high in surface leaves(E_(opt)≈923µmol photon m^(-2) s^(-1) PPFD,POER_(max)≈76.1µmol O_(2) g^(-1) Chl a s^(-1)).Leaf titratable acid(C4 acid pool)is too small(≈20 to 50 mol H+m^(-3))to support substantial SAM metabolism.Gross daily photosynthesis of surface leaves is≈3.71 g C m^(-2) d^(-1) in full sun and as much as 1.4 gC m^(-2) d^(-1) in shaded submerged leaves.There is midday inhibition of photosynthesis.

Phytotoxicity of four herbicides on Ceratophyllum demersum,Vallisneria natans and Elodea nuttallii

The physiological effects of 4 herbicides （butachlor, quinclorac, bensulfuron-methyl and atrazine） on 3 submerged macrophytes （Ceratophyllum demersum, Vallisneria natans and Elodea nuttallii） were tested in laboratory. The variables of the relative growth rate and the photosynthetic pigment content showed that all of the tested herbicides affected the growth of the plants obviously, even at the lowest concentration （0.0001 mg/L）. Except for the C. demersum treated with quinclorac at 0.005 and 0.01 mg/L, the relative growth rates of the plants were inhibited significantly （p 〈 0.01）. Statistical analysis of chlorophyll a （Chl-a） contents was carded out with both the t-test and one-way ANOVA to determine the difference between the treatment and control. The results showed that Chl-a contents of the plants in all treatment groups were affected by herbicides significantly, except for the C. demersum treated with bensulfuron-methyl at 0.0005 mg/L. The decrease in Chl-a content was positively correlated to the dosage of the herbicides in most treatment groups. It was suggested that herbicides in water bodies might potentially affect the growth of aquatic macrophytes. Since the Chl-a content of submerged macrophytes responded to the stress of herbicides sensitively and directly, it could be used as a biomaker in environmental monitoring or in the ecological risk assessment of herbicide contamination.

Patterns and trajectories of macrophyte change in East China’s shallow lakes over the past one century

Macrophytes play important roles in maintaining ecosystem health and stability of shallow lakes.Better understanding of their long-term dynamics has important theoretical and practical significance for both lake ecosystem restoration and eutrophication control.However,the knowledge about the historical status and changing patterns of macrophytes in China’s shallow lakes is still controversial and lacks systematic research.Here,we reviewed and synthesized the published records of submerged macrophytes from 14 typical shallow lakes in the eastern plain covering the past 100 years.The results suggest that submerged macrophytes have experienced three clear stages of change:rare period(the 1900s–the 1950s),growth period(the 1950s–the 1980s),and recession period(the 1980s–now).This finding is different from the traditional understanding that submerged macrophytes were abundant in the early 20th century and have been degrading since then.On this basis,we proposed the possible evolution pattern(less-more-less)of submerged macrophytes in the eastern plain lake region over the past 100 years,which provides new perspectives about the long-term evolution process of macrophytes in shallow lakes.Furthermore,we found that the decline of submerged macrophytes during the regime shift shows a gradual process at the interdecadal scale;this finding contradicts the classical regime shift theory that macrophytes decline sharply during the critical transition.This study has important theoretical value for the restoration of the eastern plain lakes in China from“turbid lake”to“clear lake”,especially for establishing the historical reference condition and restoration path of macrophytes.

A restoration-promoting integrated floating bed and its experimental performance in eutrophication remediation

Numerous studies on eutrophication remediation have mainly focused on purifying water first, then restoring submerged macrophytes. A restoration-promoting integrated floating bed （RPIFB） was designed to combine the processes of water purification and macrophyte restoration simultaneously. Two outdoor experiments were conducted to evaluate the ecological functions of the RP1FB. Trial 1 was conducted to compare the eutrophication purification among floating bed, gradual-submerging bed （GSB） and RPIFB technologies. The results illustrated that RPIFB has the best purification capacity. Removal efficiencies of RPIFB for TN, TP,NH4＋-N, NO3-N, CODcr, Chlorophyll-a and turbidity were 74.45%, 98.31%, 74.71%, 88.81%, 71.42%, 90.17% and 85%, respectively. In trial 2, influences of depth of GSB and photic area in RPIFB on biota were investigated. When the depth of GSB decreased and the photic area of RPIFB grew, the height of Potamogeton crispus Linn. increased, but the biomass of Canna indica Linn. was reduced. The mortalities of Misgurnus anguillicaudatus and Bellamya aeruginosa in each group were all less than 7%. All results indicated that when the RPIFB was embedded into the eutrophic water, the regime shift from phytoplankton-dominated to macrophyte-dominated state could be promoted. Thus, the RPIFB is a promising remediation technology for eutrophication and submerged macrophyte restoration.

Interactions between invasive Eurasian watermilfoil and native water stargrass in Cayuga Lake, NY, USA

Aims Eurasian watermilfoil(Myriophyllum spicatum L.)is a common invasive plant in American lakes and has many negative impacts on invaded ecosystems.Drastic decline of this plant at the north-ern end of Cayuga Lake in the New York State has occurred since the 1980s,with a much smaller magnitude of decline of Eurasian watermilfoil at the southern end(Johnson et al.2000)During the same period,increases in the abundance of native plants,partic-ularly water stargrass(Heteranthera dubia(Jacq.)MacMill.),have been observed(Johnson et al.(1998))We aimed to investigate the mechanisms responsible for the decline of Eurasian watermilfoil and evaluate the responses of co-occurring plants at the two ends of Cayuga Lake over time.We hypothesized that plant interactions might have contributed to the drastic decline of Eurasian watermil-foil,particularly allelopathy by native water stargrass.Methods A lake survey was conducted to assess distribution and abundance of plant communities at the northern end and the southern end of Cayuga Lake.Additionally,two sets of greenhouse experiments were conducted to investigate the interactions between invasive Eurasian watermilfoil and native water stargrass.A competition experiment evaluated intra-versus inter-specific competition among plants grown together;an allelopathy experiment examined the responses of plants to each other’s extracts.Important Findings The lake survey showed that water stargrass was extremely abun-dant at the northern end,whereas Eurasian watermilfoil was sparse at the northern end but one of the most common species at the southern end.The survey also revealed that water stargrass was more abundant than Eurasian watermilfoil at sites where the two species coexisted in the lake.Results from greenhouse experi-ments revealed no effects of Eurasian watermilfoil on water star-grass growth.However,Eurasian watermilfoil biomass was reduced by 46%when treated with high concentration of water stargrass extracts.This is likely due to osmotic effects rather than allelopathic effects of water stargrass.We proposed several possible reasons for the drastic decline of Eurasian watermilfoil and the increase in water stargrass abundance at the northern end of Cayuga Lake,including space competition,nutrients,substrates,wind exposure and water clarity in addition to insect herbivory and mechanic harvesting.