Periphyton responds differentially to nutrients recycled in dissolved or faecal pellet form by the snail grazer Theodoxus fluviatilis

1. We aimed to separate the effects of grazers on periphyton via grazing from that of nutrient recycling from their faecal pellets. 2. We set up three different experimental treatments (snails/no snails/faecal pellets) and sampled over 16 days. The ‘snail’ treatment contained a low density (snail biomass c. 14 g^?2) of the gastropod grazer Theodoxus fluviatilis and the ‘faecal pellet’ treatment received the same amount of faecal pellets as were produced in the ‘snail’ treatment. Whereas the ‘faecal pellet’ treatment provided extra nutrients to periphyton from the faeces, the ‘snail’ treatment provided nutrients in the form of both faeces and in excreta. There was also direct grazing on periphyton in the ‘snail’ treatment. The ‘no snail’ was not grazed and received no nutrients in faeces or excreta. 3. We measured periphyton C, N and P content, chlorophyll‐a (chl‐a), primary production, bacterial biomass, bacterial production and bacterial respiratory activity. In the water column we measured dissolved inorganic N and soluble reactive P. 4. Snails increased the amount of dissolved inorganic N in the water. On day 16, the periphyton N : P ratio in the ‘faecal pellet’ treatment was lower, and periphyton P content was higher, than in the other two treatments. N : P ratios decreased over time in the ‘faecal pellet’ treatment. Primary and bacterial production were positively correlated in all treatments. 5. Algal chl‐a and primary production of periphyton per unit area and periphyton chl‐a : C ratios increased over the 16 day in the ‘snail’ treatment, and thus excretion of dissolved N by snails had a stronger positive effect on the periphyton community than N and P in faecal pellets. 6. Our data show that excretion and egestion can have different effects on periphyton, probably because of the higher proportion of dissolved N in excreta and the higher proportion of P recycled in faecal pellets. The relative effect of nutrients recycled in egesta or in excretions, probably depends on the form of nutrient limitation of the periphyton. Further, the different components of the periphyton matrix could react differently to the different forms of nutrient recycling. 7. We conclude that direct grazing effects are less important than nutrient effects when nutrients are limiting and grazing pressure is low. Further, the spatial separation of different grazing effects can lead to differences in periphyton production and nutrient stoichiometry. This might be an explanation for the patchiness of periphyton in nature.     

Diatom Succession and Interaction in Littoral Periphyton and Plankton

Periphyton and plankton samples were collected at four littoral stations in a relatively shallow, eutrophic lake (Elk Lake, B.C., Canada) over a six month period from August 1967 to January 1968. The most abundant planktonic diatoms demonstrated a pattern of seasonal succession typical of temperate lakes, and all were present in the periphyton. This pattern was identical at all stations:Fragilaria crotonensis was dominant from August to October,Asterionella formosa in November and December, andMelosira italica (plusM. varians) in January.F. virescens, although never dominant, peaked in October. Periphyton communities were dominated byAchnanthes minutissima, Cocconeis placentula, F. crolonensis andF. virescens. Considerable station variation in successional patterns occurred over different exposure periods; station differences were least in those samples immersed for monthly intervals, and greatest in those immersed for a cumulative four-month period. Interaction between the phytoplankton and periphyton was illustrated by the occurrence of species common to both habitats. A decrease in cell numbers and percent abundance of these species in planktonic populations coincided with their increase in the periphyton, a relationship which appeared dependent on the breakdown of thermal stratification in November. For example, following turnover,F. crotonensis andA. formosa settled out of the plankton and correspondingly increased in percent abundance in the periphyton. This interdependence was less evident in the four-month samples, whereA. minutissima andC. placentula dominated throughout the entire period and appeared to out-compete the more typical planktonic components for diminishing substrate area. Species interaction or competition was accentuated as exposure duration and periphyton total cell standing crops increased and species diversity decreased, and appeared to account in part, for station differences in successional patterns.     

How protective are refuges? Quantifying algal protection in rock crevices

1. Refuges can be functionally important if they harbour sufficient organisms during disturbance to augment population recovery. I quantified the protection of stream algae in crevice refuges using the applied, severe disturbance of scrubbing. Scrubbing effectively removed visible surface algae, and algae remaining on stones were considered protected. 2. In a field experiment, substrata with different quantities of crevice (glass bottles; greywacke; schist; pumice) were incubated in a channelised stream. The possible influence of growth conditions was investigated concurrently; half of the substrata were suspended in the water column, the rest were placed on the bottom (providing differential access to grazers and exposing them to different flow conditions). 3. Rougher substrata had greater total algal biomass than smoother substrata; this pattern resulted from more algal biomass in crevices of the rougher substrata. Protection from scrubbing ranged from about 5% of total algal biomass on glass and greywacke to 80% on pumice. In contrast, algal biomass removed by scrubbing was similar among the experimental substrata. Suspended substrata had more chironomid grazers than those on the bottom, and also greater algal biomass, possibly because of high algal concentrations in the chironomid retreats. 4. A survey of stones from three rivers supported the experimental results; namely, rough pumice protected more algae from scrubbing than did smoother greywacke. 5. In a separate experimental assay, there was no difference in algal growth on agar plates with and without added powdered rock substrata, suggesting that crevice characteristics and not substratum chemistry produced the differences in algal assemblages between rough and smooth surfaces. 6. Results indicate that rough stream stones may protect sufficient algae to augment their recovery in streams following disturbance.     

STREAM PERIPHYTON PHOTOACCLIMATION RESPONSE IN FIELD CONDITIONS: EFFECT OF COMMUNITY DEVELOPMENT AND SEASONAL CHANGES1

The photochemical behavior of intact stream periphyton communities in France was evaluated in response to the time course of natural light. Intact biofilms grown on glass substrata were collected at three development stages in July and November, and structural parameters of the biofilms were investigated (diatom density and taxonomy). At each season, physiological parameters based on pigment analysis (HPLC) and pulse‐amplitude‐modulated (PAM) chl fluorescence technique were estimated periodically during a day from dawn to zenith. Regardless of the community studied, the optimal quantum yield of PSII (F_v/F_m), the effective PSII efficiency (Φ_PSII), the nonphotochemical quenching (NPQ), and the relative electron transport rate (rETR) exhibited clear dynamic patterns over the morning. Moreover, microalgae responded to the light increase by developing the photoprotective xanthophyll cycle. The analysis of P‐I parameters and pigment profiles suggests that July communities were adapted to higher light environments in comparison with November ones, which could be partly explained by a shift in the taxonomic composition. Finally, differences between development stages were significant only in July. In particular, photoinhibition was less pronounced in mature assemblages, indicating that self‐shading (in relation to algal biomass) could have influenced photosynthesis in older communities.     

The response of stream periphyton to Pacific salmon: using a model to understand the role of environmental context

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Top‐down and bottom‐up control of periphyton by benthivorous fish and light supply in two streams

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Contribution of biofilm‐dwelling consumers to pelagic–benthic coupling in a large river

1. Over the course of this 17‐month study, we assessed the potential loss of plankton (bacteria, algae, heterotrophic flagellates) to consumers (ciliates and rotifers) within mature biofilms established on natural substrata exposed to the main current of the River Rhine (Germany). Once a month, in flow cells in a bypass system to the River Rhine, we measured the clearance rates of the biofilm‐associated consumers on the different groups within the natural plankton. 2. Ciliates were the most dominant consumers, among which planktivorous groups, particularly peritrichs and (in spring and summer) heterotrichs dominated. Consumer biomass varied with season, with the highest density occurring directly after the appearance of the phytoplankton spring peak. 3. Clearance rates on plankton ranged from 96 to 565 L m^?2 d^?1 for bacteria and 66–749 L m^?2 d^?1 for algae, with a preference for algae in summer and for bacteria in winter. This pattern coincided with seasonal changes in the structures of the grazer communities. The consumers (both ciliates and rotifers with total standing stocks ranging between 19 and 572 mg C m^?2) imported a substantial amount of organic matter (between 15 and 137 mg C m^?2 d^?1) into the biofilm. 4. These results highlight the potential importance of consumers in the biofilm as a trophic link between the plankton and the benthos, a function that has hitherto mostly been attributed to filter‐feeding bivalves. In contrast to bivalves, the biofilm‐dwelling consumers show a more dynamic response towards the plankton density and composition. Such dynamic components need to be considered when estimating total plankton consumption by the benthos.     

In situ interactions between the effects of season,current velocity and pollution on a river biofilm

1. The development of periphytic algae and bacteria is controlled by a combination of interacting biotic processes and abiotic factors. Distinguishing between the selection pressure resulting from pollution and that of natural environmental factors is therefore one of the most critical aspects of assessing the impact of pollutants on the diversity and function of benthic microbial communities in natural ecosystems. 2. We studied how current velocity and season affect the ability of river biofilms to cope with complex chemical pollution. We compared the diversity, structure and production of periphytic algae and bacteria from four sampling zones with differing chemical water quality levels and different flow velocities over the course of two seasons (summer and winter). 3. The three factors tested all influenced biofilm development, but this depended on the biological variable being measured. Bacterial and algal densities were highly dependent on season and chemical water quality. Algal density was lower in summer than in winter, but bacterial density and production increased from upstream (reference) to downstream (polluted), and this increase was more marked in winter. The impact of chemical water quality was also dependent on the season. 4. An interaction between current velocity and pollution was also detected. During the summer, there was no difference in bacterial density or production between the upstream and downstream segments in the fast current zones, whereas both variables were higher downstream in the slow current zones. Such interactions between environmental factors and the impact of water quality on biofilms must be taken into account in assessments of the effects of chemicals on biofilm community structure and functioning in rivers.     

Qualitative examination of diatom diversity in black fly (Diptera: Simuliidae) gut contents across a mortality gradient after exposure to Bacillus thuringiensis subsp. israelensis de Barjac, 1978

While Bacillus thuringiensis subsp. israelensis de Barjac, 1978 de Barjac, H. (1978), ‘Une Nouvelle Variété de Bacillus thuringiensis Très Toxique Pour les Moustiques: B. thuringiensis var. israelensis Sérotype H14’, Comptes Rendus de l'Académie des Sciences de Paris 286D, 797–800. [Google Scholar] (Bti) is generally effective in reducing Simulium Latreille, 1802 Latreille, P.A. (1802–1803), Histoire Naturelle, Générale et Particulière des Crustacés et des insectes (Vol. 3), Paris: F. Dufart. [Google Scholar] pest levels, impaired toxicity has been identified. Diatoms are typically abundant in Simulium larval gut contents and thought to affect feeding behaviour and/or Bti ingestion. We sought to determine diatom composition in larval Simulium gut contents and periphyton in relation to mortality subsequent in situ Bti application across three stream reaches. Generally, lower larval mortality after Bti exposure was associated with comparatively greater diatom diversity inside gut contents, corresponding with an inverse relationship between periphyton diatom diversity and larval mortality across our study reaches. High profile diatoms inside gut contents were positively correlated with mortality, suggesting microbial morphology may affect Bti efficacy. Diatoms inside live larval guts were most similar to periphyton which indicates that these larvae may have been feeding less on lower quality detrital material and/or exhibiting comparatively greater benthic feeding.