Relationship between epipelon,epiphyton and phytoplankton in two limnological phases in a shallow tropical reservoir with high Nymphaea coverage

Macrophytes and phytoplankton are recognized as having roles in determining alternative stable states in shallow lakes and reservoirs, while the role of periphyton has been poorly investigated. Temporal and spatial variation of phytoplankton, epipelon and epiphyton was examined in a shallow reservoir with high abundance of aquatic macrophytes. The relationships between algae communities and abiotic factors, macrophyte coverage and zooplankton density were also analyzed. Monthly sampling was performed in three zones of the depth gradient of the reservoir. Two phases of algal dominance were found: a phytoplankton phase and epipelon phase. The phase of phytoplankton dominance was characterized by high macrophyte coverage. Rotifera was the dominant zooplankton group in all the zones. Flagellate algae were dominant in phytoplankton, epipelon and epiphyton. Macrophyte coverage was found to be a predictor for algal biomass. Changes in biomass and species composition were associated with macrophyte cover variation, mainly the Nymphaea. In addition to the abiotic factors, the macrophyte coverage was a determining factor for changes to the algal community, contributing to the alternation between dominance phases of phytoplankton and epipelon. The macrophyte–phytoplankton–periphyton relationship needs to be further known in shallow reservoirs, especially the role of epipelon as an alternate stable state.

     

Regulation of submersed macrophyte biomass in a temperate lowland river: Interactions between shading by bank vegetation, epiphyton and water turbidity

Growth of aquatic vegetation is often controlled by light supply, which is potentially decreased by bank vegetation, water turbidity and epiphytic biofilm. To understand the relative importance of these shading factors and the interactions between them we analysed the seasonal course of macrophyte biomass, shading by bank vegetation, turbidity of the water column and epiphytic light absorption in shaded and sunny sections of a temperate eutrophic lowland river. At a shaded site, bank vegetation decreased the light supply by 79%, 0.5 m water column by 45% and 2-week-old epiphyton by 28% during the vegetation period. Growth of submersed macrophytes, but not of epiphyton, was light-limited in the shaded sections. We found a saturation-type correlation between light supply and macrophyte biomass. Therefore, the additional light absorption of the water column or epiphyton only shortened the period of optimum light supply at the sunny site, but was crucial for macrophyte development at the shaded site. Light absorption of phytoplankton was most important in spring and that of epiphyton in late summer. Submersed macrophytes effectively retained particles and thus improved light supply of downstream stands, but this positive feedback effect was only relevant for shaded sections in summer.     

Fish induced macrophyte loss in shallow lakes: top–down and bottom–up processes in mesocosm experiments

SUMMARY 1. Macrophyte loss from Sites of Special Scientific Interest in England has become widespread over the last 20 years. One reason for this may be changing trends in angling, a multimillion pound industry that has an enormous impact on aquatic ecosystems. Stocking with cyprinid fish is a common angling management practice but the particular fish species and distribution of their biomass may be crucial to the ecosystem. 2. Carp (Cyprinus carpio), roach (Rutilus rutilus), bream (Abramis brama) and tench (Tinca tinca) at biomasses ranging from 0 to 800 kg ha^?1 and at various sizes were placed into experimental mesocosms in Little Mere, a shallow, fertile lake in Cheshire, U.K. The effects these treatments had on the aquatic ecosystem were studied over two summers. Specifically the effects of the treatments on macrophyte growth, benthic and macrophytic macro‐invertebrate populations, water chemistry, epiphyton production and plankton survival were investigated. 3. Carp had a greater detrimental effect on the macrophytes than bream, tench and in particular roach. A biomass of fish > 200 kg ha^?1 adversely affected the extent of macrophyte growth. 4. The decline in macrophyte growth was most likely as a result of increased epiphyton growth that probably reduced the amount of light and carbon dioxide available to the plant. There were no observed direct fish impacts on macrophytes. 5. The chemical data suggested that inorganic nitrogen levels were low and it is possible that release of nitrogen, from fish excreta, followed by immediate uptake, could have been a major factor stimulating epiphyton growth and subsequently macrophyte loss. Phosphorus concentrations increased even in the controls and substantial amounts were available. Phosphorus stimulation can therefore be discounted. Macrophyte‐associated macro‐invertebrates were positively correlated with epiphyton load but had no impact on the extent of epiphytic growth. Shading from disturbed sediment or phytoplankton was also unimportant.     

Seasonal succession of lotic epiphyton: effects of macrophyte cover and grazing by chironomid larvae

The epiphyton is a community of periphyton on the aquatic plants. The seasonal dynamics of biomass, production and community structure of epiphytic algae on Potamogeton perfoliatus was studied at the Moscow River reach with low flow velocity from May to October 2000. The relative importance of physical-chemical environmental variables and herbivorous pressure in structuring the epiphyton community was accessed using multivariate redundancy analysis. Algal composition was mainly determined by the macrophyte cover, suggestion the overriding importance of the underwater light availability in seasonal development of epiphyton. There was a significant positive relationship between biomass of attached diatom Cocconeis placentula and abundance of chironomid larvae (r = 0.57; p < 0.01). This indicates that increase in herbivorous pressure caused the shift in epiphyton community structure. However, the intensity of herbivorous impact on epiphyton was ultimately determined by light availability in macrophyte stands. At low irradiance level and consequent slow growth of algae the upper layer of epiphyton was seriously destroyed by chironomids that resulted in dominance of C. placentula. As irradiance level and epiphyton productivity increased chironomids were unable to prevent biomass accumulation of the upper layer thus inhibiting the growth of C. placentula. The high concentrations of nutrients and relatively stable discharge determined the crucial role of light regime in the regulation of trophic interactions between epiphyton and chironomid larvae.     

Development of submerged macrophyte and epiphyton in a flow-through system: Assessment and modelling predictions in interconnected reservoirs

Every approach to lake restoration requires the reestablishment of submerged macrophytes. However, macrophyte overgrowth in shallow lakes may lead to deterioration and a consequent necessity for restoration treatments. We assumed that a major threat to the increased trophic level in the Jankovac flow-through system arises from the sediment, where the accumulation of deciduous leaf litter and decayed macrophyte fragments could generate anoxic conditions. The integrated Water Quality Model (WQM) and the Submerged Aquatic Vegetation Model (SAVM) were combined in the Jankovac Model (JanM) and applied during the vegetated season in 2008 and 2014, with the aim to offer a possible approach to the maintenance of good water quality. The impacts of flow velocity and epiphyton growth on submerged macrophyte coverage and biomass were simulated. Biocenotic analyses suggested that epiphyton growth was more extensive in 2014 in comparison to 2008. The results of JanM indicated that increased flow velocities enhanced macrophyte growth and dissolved oxygen concentrations concurrently with the decline of epiphyton biomass. Furthermore, results suggested that epiphyton growth rate of 0.4 d⁻¹ maintained macrophyte coverage and biomass at a satisfactory level of 70% reservoir coverage. Considering the proposed scenarios hydraulic treatment could be applied to regulate submerged macrophytes in shallow reservoirs, as an efficient and less invasive approach than sediment removal, especially in sensitive karst areas.     

Seasonality of chlorophyll and nutrients in Lake Erken – effects of weather conditions

The effect of submerged macrophytes on interactions among epilimnetic phosphorus, phytoplankton, and heterotrophic bacterioplankton has been acknowledged, but remains poorly understood. Here, we test the hypotheses that the mean summer phytoplankton biomass (chlorophyll a): phosphorus ratios decrease with increased macrophyte cover in a series of nine lakes. Further, we test that both planktonic respiration and bacterioplankton production increase with respect to phytoplankton biomass along the same gradient of increasing macrophyte cover. Increased macrophyte cover was associated with a lower fraction of particulate phosphorus in epilimnia, with total particulate phosphorus declining from over 80% of total phosphorus in a macrophyte free lake to less than 50% in a macrophyte rich lake. Phytoplankton biomass (chlorophyll a) too was lower in macrophyte dominated lakes, despite relatively high levels of total dissolved phosphorus. Planktonic respiration and bacterioplankton production were higher in macrophyte rich lakes than would be expected from phytoplankton biomass alone, pointing to a subsidy of bacterioplankton metabolism by macrophyte beds at the whole lake scale. The results suggest that the classical view of pelagic interactions, which proposes phosphorus determines phytoplankton abundance, which in turn determines bacterial abundance through the production of organic carbon, becomes less relevant as macrophyte cover increases.     

Physical variables driving epiphytic algal biomass in a dense macrophyte bed of the St. Lawrence River (Quebec, Canada)

The variables affecting epiphyton biomass were examined in a sheltered, multispecies macrophyte bed in the St. Lawrence River. Alteration of light penetration, resulting from the presence of dense macrophytes forming a thick subsurface canopy, primarily determined epiphyton biomass. Seasonal decrease of water levels also coincided with major increases in biomass. Plant morphology was the next important variable influencing epiphytic biomass, whereas the contribution of other variables (sampling depth, macrophyte species, relative abundance of macrophytes, and temperature) was low. Groups of lowest epiphyte biomass (0.1–0.6 mg Chla g^–1 DW) were defined by the combination of a low percentage of incident light (<13% surface light) and simple macrophyte stem types found below the macrophyte canopy. Highest epiphyte biomass (0.7–1.8 mg Chla g^–1 DW) corresponded to samples collected in mid-July and August, under high irradiance (>20% surface light) and supported by ramified stems. Our results suggest that epiphyton sampling should be stratified according to the fraction of surface light intensity, macrophyte architecture, and seasonal water level variations, in decreasing order of influence.     

Wave exposure related growth of epiphyton: implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.     

Patch- and reach-scale dynamics of a macrophyte-invertebrate system in a New Zealand lowland stream

Abundant growths of macrophytes are a common feature of streams in open lowland areas of New Zealand during summer, but the values of these to aquatic biota are poorly understood. We studied the temporal dynamics of, and associations amongst, elements of a macrophyte-invertebrate system to provide an improved information base for lowland stream management. The biomass of macrophytes increased significantly over the four quarterly sampling occasions from 43.8 g m^-2 in June to 370.8 g m^-2 in March; biomass was dominated by Egeria densa on all dates, except in December when Potamogeton crispus was dominant. We did not detect strong associations between epiphyton biomass and invertebrate abundance in our study, but this may reflect the fact that we sampled loosely-adhering epiphyton on young, surface-reaching shoots whereas invertebrates were collected from macrophytes growing through the water column. Density of some invertebrate species per gram dry weight of plant material varied by macrophyte type, with the chironomids Tanytarsus vespertinus and Naonella forsythi displaying positive correlations with Egeria and Potamogeton biomass, respectively. The shrimp Paratya curvirostris accounted for 50% of phytophilous invertebrate biomass, with Chironomidae the only other group to comprise more than 9%. Abundance of total phytophilous invertebrates displayed a positive linear relationship with macrophyte biomass in a sample (0.1 m²), and a humped relationship with species richness, such that highest numbers of taxa occurred at macrophyte biomass levels around 400 g dw m^-2. Our study suggests that intermediate macrophyte biomass levels are likely to enhance macroinvertebrate biodiversity in sandy-bottomed lowland streams. This revised version was published online in July 2006 with corrections to the Cover Date.     

Morphological responses of a submerged macrophyte to epiphyton

Epiphyton might have distinctive influence on the morphology of substrate macrophyte. In this article, we evaluate the influence of epiphyton on the morphological characteristics of their substrate submerged macrophyte, Potamogeton perfoliatus under two light intensities. The experiment was carried out for a period of 84 days in 12 glass aquaria under laboratory conditions. It was based on a 2 × 2 factorial design with epiphyton status (present or absent) and light intensity (200 or 80 μE m⁻² s⁻¹). Both epiphyton and light intensity had significant effects on the morphology and biomass allocation of the experimental plants. The average number of leaves, total length of newly recruited shoots and diameter of stems were greater in the epiphyton-free control plants than in the epiphyton-colonized plants under low light conditions. The plants with epiphyton allocated more biomass in their rhizomes and roots (% relative to total biomass basis) when compared to the control plants in both light intensities. There were also significant epiphyton–light interactions. The control plants under low light intensity showed higher internodal elongation in their main shoots when compared to the plants under high light intensity as an adaptation mechanism. Whereas the plants with epiphyton did not show such an adaptation. The new shoots of the control plants under low light intensity did not show any internodal elongation as observed in the main shoots. Furthermore, the length of the leaves of main shoots was larger in control plants with epiphyton and high light intensity than in plants with epiphyton and low light intensity, but such a variation was absent in the new shoots. We conclude that the long-term colonization by epiphyton and their shading effects induced the observed morphological changes in plants.     

The contribution of epiphyton to the primary production of tropical floodplain wetlands

Tropical floodplains are one of the most productive ecosystems on earth. Studies on floodplain productivity have primarily focused on trees and macrophytes, rather than algae, due to their greater biomass. However, epiphyton—algae and bacteria attached to the submerged portion of aquatic macrophytes—is a major source of energy in many tropical floodplains. Epiphyton productivity rates are unknown for most tropical floodplain wetlands, and spatial variability is not well understood. In this study, we measured primary productivity of epiphyton in Kakadu National Park in northern Australia. We estimated the relative contribution of epiphyton to aquatic production (epiphyton, + phytoplankton + macrophytes). We sampled sites dominated by different macrophyte structural types: vertical emerging grasses, horizontal emerging grasses, submerged macrophytes, and macrophytes with floating leaves. Epiphyton productivity was highly influenced by the structural type of the macrophyte. Highest potential productivity per weight was measured from epiphyton growing on macrophytes with floating leaves and horizontal grasses (1.52 ± 0.53 and 1.82 ± 0.61 mgC/dw g epiphyton/h, respectively) and lowest in submerged macrophytes and vertical grasses (0.57 ± 0.26 and 0.66 ± 0.47 mgC/dw g epiphyton/h, respectively). When considering the areal biomass of the macrophyte and the amount of epiphyton attached, epiphyton on horizontal grasses and submerged macrophytes had productivity values approximately ten times higher (45–219 mgC/m²/d) compared to those on vertical grasses and macrophytes with floating leaves (2–18 mgC/m²/d). Epiphyton contributed between 2 to 13 percent to the aquatic production of these tropical floodplain wetlands.     

Wetland macrophyte diversity, trophic status and phytoplankton： a case study of the cascading effects of an invasive herbivore

Shallow water bodies can exist in alternative stable states, a clear water state with high coverage of macrophytes or a turbid state with high phytoplankton biomass. The alternative equilibria hypothesis has been proposed to explain the occurrence of the alternative stable states (Scheffer et al., 1993)[1], which assumes that: 1),     

Floating meadow epiphyton: biological and chemical features of epiphytic material in an Amazon floodplain lake

SUMMARY. 1. Quantities and the chemical composition of epiphyton on the roots of floating aquatic macrophytes were measured in Lake Calado, an Amazon floodplain lake. Growth of epiphytic algae following physical disturbance and losses of epiphyton due to grazing and storms were investigated.
2. Deposition of silt from invading river water decreased chlorophyll and nutrient content (%C, %N, %P) of epiphyton during rising water. N:P ratios of epiphyton indicated that proximity to the river increased supplies of phosphorus. Attached algal biomass per unit root tissue was higher overall during the falling water period, when light was greater, storms less frequent, and new host plant tissue produced more slowly.
3. Epiphytic algal biomass at the margins of floating meadows exceeded that of the phytoplankton in the open water on a per unit area basis. Increases in attached algal chlorophyll ranged from two- to ten-fold over 1 week. Artificial denudation of roots was followed by rapid regrowth of attached algae, leading, after I week, to four-fold increases in chlorophyll over the pre-denuded state.
4. Wind-blown macrophytes experienced an episodic loss of 70% of epiphytic material in less than 1 h. Particulate material lost from roots grazed by snails included root tissue and contained significantly more carbon than material lost from ungrazed roots.     

Influence of nutrients, submerged macrophytes and zooplankton grazing on phytoplankton biomass and diversity along a latitudinal gradient in Europe

In order to evaluate latitudinal differences in the relationship of phytoplankton biomass and diversity with environmental conditions in shallow lakes, we sampled 98 shallow lakes from three European regions: Denmark (DK), Belgium/The Netherlands (BNL) and southern Spain (SP). Phytoplankton biomass increased with total phosphorus (TP) concentrations and decreased with submerged macrophyte cover across the three regions. Generic richness was significantly negatively related to submerged macrophyte cover and related environmental variables. Zooplankton:phytoplankton biomass ratios were positively related to submerged macrophyte cover and negatively to phytoplankton generic richness in DK and BNL, suggesting that the low generic richness in lakes with submerged macrophytes was due to a higher zooplankton grazing pressure in these regions. In SP, phytoplankton generic richness was not influenced by zooplankton grazing pressure but related to conductivity. We observed no relationship between phytoplankton generic richness and TP concentration in any of the three regions. The three regions differed significantly with respect to mean local and regional generic richness, with BNL being more diverse than the other two regions. Our observations suggest that phytoplankton diversity in European shallow lakes is influenced by submerged macrophyte cover indirectly by modulating zooplankton grazing. This influence of submerged macrophytes and zooplankton grazing on phytoplankton diversity decreases from north to south.     

不同初始螺类密度对沉水植物刺苦草Vallisneria spinulosa Yan及其附着藻类的影响

大型水生植物及其附着藻类是浅水湖泊中的重要初级生产者。淡水螺类作为重要的初级消费者,其密度对沉水植物及其附着藻类的影响存在争议。本研究设置4种初始螺类密度（0、40、80、240 ind·m^-2）,研究淡水螺类（椭圆萝卜螺Radix swinhoei H.Adams）对刺苦草（Vallisneria spinulosa Yan）及其附着藻类的直接牧食作用和螺类种群的变化。结果显示,在添加螺处理中,刺苦草和人工基质表面附着藻类的生物量显著降低,同时沉水植物的生长显著增加,在较高初始密度螺类处理中刺苦草产生更多的分株。到实验结束时,螺类的死亡率较高,但3个有螺处理间螺类鲜重无显著差异,而高初始密度螺类条件下的最终密度仍较高,同时个体重量（均重）也较小。在中富营养条件下淡水螺类可以直接牧食沉水植物叶片,但对植物生长的抑制作用不显著,有可能是因为沉水植物并不能作为唯一的食物来源维持螺类种群,同时螺类的种群结构受到水体营养水平等因素的制约。     

Phytoplankton as a factor in the decline of the submersed macrophyte Myriophyllum spicatum L. in Lake Wingra,Wisconsin, U.S.A.

A dramatic decline in biomass and areal coverage of the submersed macrophyte Myriophyllum spicatum in Lake Wingra, Wisconsin, USA during the mid-1970's is documented using aerial photography, vegetation surveys, and quadrat biomass sampling. Over the same period, light penetration as measured by Secchi disc transparency and extinction coefficient decreased substantially. During this period, extinction coefficient was closely correlated with chlorophyll a levels implicating phytoplankton as the major source of decreased light penetration. A growth model for M. spicatum predicts a substantial decline in macrophyte biomass when extinction coefficient is increased to the levels reported since 1977. Available data do not show whether the phytoplankton increase preceeded or followed the macrophyte decline. Nonetheless it is clear that phytoplankton growth can account for a substantial portion of the decline and that macrophyte recovery will be difficult given the phytoplankton-induced decrease in water clarity.     

Eutrophication and black swan (Cygnus atratus Latham) populations: tests of two simple relationships

In an earlier study, regression models were developed which related black swan populations to macrophyte biomass and to phytoplankton productivity in Tomahawk Lagoon No 2, a small South Island, New Zealand, lake in which phytoplankton and macrophytes vary widely from year to year, in an irregular inverse cycle. Two years of further study confirmed that winter swan populations are directly correlated with macrophytes, for samples taken on a single day in winter each year. They also confirmed that winter maximum swan populations can be predicted from measurements of phytoplankton productivity in the previous summer. This relationship is inverse. For 6 other lakes, 9 of the 11 data points lay within the 95% confidence intervals of the macrophyte-swan relationship, with the only outlier being a lake in which the macrophytic vegetation is dominated by filamentous algae. Further analysis, giving equal weight to each of the 7 lakes in the sample, confirmed that there is a significant correlation in winter between swan populations and macrophyte biomass for these lakes which show a wide range of size and trophic status. There was no such significant correlation in summer. All of the 8 data points from 7 other lakes lay within the rather broad 95% confidence intervals of the phytoplankton- swan relationship. The 4 lakes which fitted most closely to the predictions of this relationship have silty sediment. The other 3, which all had lower swan populations than predicted, have sediment predominantly of sand or fine clay, and macrophyte biomass and swan populations may be constrained by the nature of the sediment, rather than by shading effects from phytoplankton.     

Origin of phytoplankton and the environmental factors governing the structure of microalgal communities in lowland streams

Information on the structure of microalgal assemblages in the epiphyton and epilithon is necessary to understand the origin of phytoplankton in lowland rivers. To this end, we carried out concurrent investigations on phytoplankton, epiphyton and epilithon in 18 reaches of three Estonian rivers during the midsummers of 2002 and 2003. A total of 251 taxa was recorded, of which 192 were epiphyton species, 158 were epilithon species and 150 were phytoplankton species. Canonical correspondence analysis (CCA), based on the 31 most abundant taxa, indicated differences in the structure of the algal assemblages between the different biotopes (phytoplankton, epiphyton and epilithon) as well as between the studied rivers. The composition of the phytoplankton clearly differed from that of the other biotopes, with prevailing small flagellates, a chrysophyte (Synura uvella) and cryptophytes (Rhodomonas lacustris and Cryptomonas erosa). The epiphyton was characterized by a large number of diatoms, while the epilithic community contained filamentous cyanobacteria (Phormidium tergestinum and Planktolyngya sp.) and a green alga (Stigeoclonium tenue) in addition to diatoms. Based on redundancy analysis (RDA), phosphorous was the most relevant parameter determining the distribution of species in the phytoplankton assemblages. Shading by trees on the river bank, dissolved oxygen concentration and water temperature as well as river width determined the distribution of species in the epiphyton. The data set on the epilithon did not reveal any significant relationships between species distribution and the measured environmental parameters.     

Epiphyte shading: Its role in resulting depth distribution of submerged aquatic macrophytes

The biomass of epiphytic algae increased more than the biomass of phytoplankton with increased nutrient availability (N, P, Si) in four lakes. Large epiphyte biomasses on the submerged macrophyte,Littorella uniflora, reduce light attenuation more than phytoplankton did. High values of shading by epiphytes correspond with a reduction in the maximum depth penetration ofLittorella by more than 1 meter.