Ingestion and digestion of epilithic algae by larval insects in a heavily grazed montane stream

1. Epilithic algae grown on elevated or non-elevated ceramic tiles were exposed (to produce assemblages with different grazing histories) in a heavily grazed, montane stream in New Mexico, U.S.A. to Ameletus nymphs (Ephemeroptera) and Ecclisomyia larvae (Trichoptera) and the algal composition in insect faeces was compared to that on the tiles. Differences in grazing and digestion efficiency between grazers were then assessed and also differences in susceptibility to ingestion and digestibility among common algae. 2. Ordination of tile and faecal samples, using the relative abundance of common algae, revealed that: (i) algal assemblages on elevated vs. non-elevated tiles differed only slightly; (ii) the taxonomic composition of algae in faeces of both caddis and mayflies differed substantially from that on the tiles, indicating low grazing efficiency for some algal taxa; and (iii) the algal composition of faeces produced by caddis larvae and mayflies was similar, indicating little difference in grazing efficiency between them. However, some algal taxa were more susceptible to ingestion by caddisfly larvae when occurring on elevated tiles than on non-elevated tiles, suggesting that previous exposure to caddis grazing influenced assemblage attributes. 3. Although Ameletus and Ecclisomyia differed little in grazing efficiency, the percentage of diatoms that were dead after passage through the gut was greatest in the mayfly treatment, suggesting that mayflies digested diatoms more efficiently than the caddis. Analyses of differences in the condition of chloroplasts within diatoms in tile and faecal samples showed that losses of ’live‘ diatom cells (i.e. those containing full chloroplasts) during gut passage through mayflies equalled the increase, in faeces, of ’dead‘ (empty frustules) cells of all common diatoms. In contrast, some diatoms were digested inefficiently by caddis larvae. 4. Algae on elevated tiles contained a higher proportion of dead diatoms than those on non-elevated tiles, possibly because mayflies visited raised tiles more often and, consequently, ingested and defaecated cells at a higher rate in the absence of caddis larvae. Moreover, diatom taxa differed in the percentage of cells that were dead within tile assemblages, with populations of typically grazer-resistant taxa (e.g. Achnanthidium minutissimum, Planothidium lanceolatum and Cocconeis placentula var. euglypta) containing significantly more dead cells than grazer-susceptible taxa [e.g. small, chain-forming Fragilaria (= Staurosirella)]. This result suggests that a trade-off exists between ingestion vs. digestion resistance of microalgae. Both the ingestion and digestion efficiency of algivorous macroinvertebrates could influence the structure and function of algal assemblages. In heavily grazed systems, where algal cells are probably processed through grazer guts repeatedly, differential resistance to digestion among algae may be particularly important.     

Algal pigment diversity in coastal sediments from Kerguelen (sub-Antarctic Islands) reflecting local dominance of green algae,euglenoids and diatoms

Summary High Performance Liquid Chromatography analysis of algal pigments from inter- and subtidal (deep and shallow) sediments from the Kerguelen Islands showed clear differences in the pigment composition at the different stations. High concentrations of chlorophyll c and fucoxanthin were present at all locations, indicating significant diatom densities, chlorophyll b was detected at all sites. At one station the other green algal pigments were also present; here green algae contributed more to chlorophyll a concentrations than diatoms, as estimated by using pigment ratios and microscopic observations. At another location chlorophyll b was associated with a high concentration of diadinoxanthin, indicating an abundance of euglenoids. This indicates that chemotaxonomy can be powerful tool in microphytobenthos studies since enumeration of living cells are difficult as many algae are attached to sediment particles (epipsammic algae). Ways of diagenesis, carotenoid degradation and the role of grazing are briefly mentioned. Phaeophorbide a-like pigments were the most significant chlorophyll a degradation products, with concentrations up to 110 g · g^–1 dry weight sediment, i.e. 10 times the chlorophyll a concentration. Some taxonomic estimations, based on pigments ratios, and their limits, are discussed.     

HPLC analysis of chlorophyll a breakdown products to interpret microalgae dynamics in a shallow bay

Phytoplankton production is determined by growth, senescence, sinking and zooplankton grazing. In an attempt to follow algal senescence and grazing, some authors have used HPLC fluorescence detection of chlorophyll a breakdown products. Laboratory grazing experiments have shown that copepods reduce chlorophyll a from diatoms leading to an increase in pheophytin a rather than pheophorbide a. However, field measurements only indicated a slight increase of pheopigment concentrations in summer. During this period, high heterotrophic activities (zooplankton and bacteria) seemed to be responsible for rapid pheopigment disappearance. On the other hand, highest chlorophyllide a levels appeared to be related to spring accumulation of nutrient-limited senescent algae. While increases in pheophytin a accounted for chlorophyll a consumption, changes in pheophorbide a concentrations could be linked to chlorophyllide a abundance. These results suggest that laboratory studies cannot be uncritically extrapolated to the field.     

Carotenoid Distribution Patterns in Bacillariophyceae (Diatoms)

The distribution of carotenoids in 10 species of diatoms was determined qualitatively and quantitatively. Monoacetylenic carotenoids (diatoxanthin and diadinoxanthin, 12–43% of total), allenic carotenoids (fucoxanthin and fucoxanthinol, 49–81% of total), monoepoxides (diadinoxanthin, fucoxanthin and fucoxanthinol, 79–91% of total) and β,β-carotene (3–8% of total) were identified and analysed by methods including TLC, HPLC, MS, VIS and cochromatography with authentic standards. Trace amounts of echinenone and canthaxanthin were obtained from Navicula pelliculosa. Chemosystematic considerations are made.     

Feeding of biofilm-dwelling nematodes examined using HPLC-analysis of gut pigment contents

The natural feeding behaviour of the nematodes Chromadorina bioculata (Schultze in Carus 1857) and Chromadorina viridis (Linstow 1876) was studied in situ, within epilithic biofilms of the Garonne River (France). Based on their feeding-type characteristics and population dynamics, it was hypothesised that these species feed selectively on microphytobenthos (MPB) within the biofilm, and that among MPB groups, diatoms are preferred. High-performance liquid chromatography (HPLC) was used for separation, identification and quantification of pigments both in nematode guts and in the biofilm. This is the first time that nematode gut pigment contents were examined under natural conditions. Diatoms dominated the MPB which also comprised cyanobacteria and green microalgae. The comparison between chlorophyll a content in nematode guts versus in the biofilm showed that C. bioculata and C. viridis fed opportunistically (non-selectively) on MPB within the biofilm. Only diatom biomarker pigments were found in nematode guts suggesting that they could preferentially fed on diatoms among MPB groups. However, the non-detection of biomarker pigments for other microphyte groups could be also linked to HPLC detection limits. It was estimated that Chromadorina nematodes daily ingested on average 0.03–0.67% of the MPB standing stock. This grazing covered only a small part of their energetic requirements, suggesting that besides MPB they probably also fed on other biofilm food sources. Some considerations on the applicability of the HPLC gut pigment analysis technique for the examination of nematode feeding are also presented.     

Comparison of Plankton-Water Chemistry Relationships in Three Acid Stressed Lakes in Nova Scotia,Canada

A field program was conducted in 1980 and 1981 on three lakes in Kejimkujik National Park, Nova Scotia, to study plankton-water chemistry relationships in a region of potential acid precipitation stress. Among lake and between year comparisons were made. The three study sites represent two different types of lakes. Kejimkujik Lake (pH 4.8) and Pebbleloggitch Lake (pH 4.5) are both high-colored, dystrophic acid lakes. Beaverskin Lake is a clear, oligotrophic lake which is not as acidic (pH 5.4). The phytoplankton of Kejimkujik Lake was dominated by diatoms, while Pebbleloggitch Lake was dominated by chlorophytes and chrysophytes, and Beaverskin Lake was dominated by cyanophytes. Kejimkujik Lake had the highest total algal cell volume per liter, and Pebbleloggitch Lake the lowest. Rotifer populations composed the majority of the zooplankton communities, while the crustacean zooplankton were dominated by the acid-tolerant copepod Diaptomus minutus. Kejimkujik Lake had the lowest zooplankton biomass and Pebbleloggitch Lake the highest zooplankton biomass per m³. A pattern of highest algal density in Beaverskin Lake, but highest zooplankton density in Pebbleloggitch Lake emerged in both years. This is likely an effect of abundant, but unsuitable food (blue-green algae) in Beaverskin Lake and important detrital food resources in Pebbleloggitch Lake in both years. Multiple regression analysis of water chemistry variables with plankton species produced many significant effects, but failed to show clear patterns. Cause and effect relationships in aquatic ecosystems are poorly delineated by such techniques.     

Grazing resistance in nutrient-stressed phytoplankton

Grazing experiments were performed with the zooplankters Daphnia pulex and Daphnia magna, feeding on phosphorus-saturated and phosphorus-limited cells of two green algae (Scenedesmus subspicatus and Selenastrum capricornutum). P-limited algal cells passed largely intact through the gut and were thus spared from heavy grazing pressure. P-saturated algal cells, in contrast, were efficiently assimilated. Structural and morphological changes in the P-limited cells most probably reduced their digestibility. This phenomenon may be an important factor in zooplankton production and competition, and may serve as an example of a highly efficient strategy of P-limited algae to resist heavy grazing pressure.     

LIPOPHILIC PIGMENTS FROM CYANOBACTERIAL (BLUE-GREEN ALGAL) AND DIATOM MATS IN HAMELIN POOL,SHARK BAY,WESTERN AUSTRALIA1

Lipophilic pigments were examined in microbial mat communities dominated by cyanobacteria in the intertidal zone and by diatoms in the subtidal and sublittoral zones of Hamelin Pool, Shark Bay, Western Australia. These microbial mats have evolutionary significance because of their similarity to lithified stromatolites from the Proterozoic and Early Paleozoic eras. Fucoxanthin, diatoxanthin, diadinoxanthin, β-carotene, and chlorophylls a and c characterized the diatom mats, whereas cyanobacterial mats contained myxoxanthophyll zeaxanthin, echinenone, β-carotene, chlorophyll a and, in some cases, sheath pigment. The presence of bacteriochlorophyll a with in the mats suggest a close association of photosynthetic bacteria with diatoms and cyanobacteria. The high carotenoids: chlorophyll a ratios (0.84–2.44 wt/wt) in the diatom mats suggest that carotenoids served a photoprotective function in this high light environment. By contrast, cyanobacterial sheath pigment may have largely supplanted the photoprotective role of carotenoids in the intertidal mats.     

Grazing experiments with two freshwater zooplankters:fate of chlorophyll and carotenoid pigments

In order to evaluate the validity of the gut pigment methodto assess grazing and diet in two freshwater zooplankters, experimentswere carried out to check chlorophyll a and xanthophyll conservationduring feeding. For both animals, two sets of experiments wereconducted by incubating animals in the laboratory, either isolatedfrom a reservoir (the calanoid copepod, Eudiaptomus gracilis)or cultured under high-food conditions (the cladoceran, Daphniagaleata). For both animals, gut pigments and clearance rateson different types of algae were determined from the same incubations.Chlorophyll a and derivatives, as well as major algal carotenoids,were analysed by High Performance Liquid Chromatography (HPLC).In copepods, the pigment profiles from the gut extracts reflectedthe diet of the animals poorly. The animal extracts containedalmost exclusively alloxanthin (or an alloxanthin-like pigment)in large amounts, whereas the other pigments were lost in highproportions (>70% for lutein and fucoxanthin; 57 and 78%for a-phorbins). The cladocerans fed on the main types of algaeabundant in the suspensions, with a preference, however, forsmall cells. Although the main xanthophylls from these algaewere detected in the Daphnia extracts, some destruction of luteinand fucoxanthin may have occurred (18.7 and 30%). The loss ratefor alloxanthin seemed more variable (0 and 68%), possibly dependingon food concentration. As for the transformation of a-phorbins,E.gracilis and D.galeata behaved quite differently. The HPLCprofiles of copepod extracts always showed a very small chlorophylla peak, along with phaeophytin a and pyrophaeophytin a. Thosefrom the cladoceran exhibited a large phaeophorbide a peak,along with some chlorophyll a and phaeophytin a. In fact, D.galeatadid not destroy a-phorbins under our experimental conditionsbut converted chlorophyll a mainly into phaeophorbide. Froma comparison of our results with data from other studies, itseems that in these two zooplankters, use of gut pigment datafor quantitative grazing assessment should be considered withcaution.     

Changes in pigment content of the diatom Navicula pelliculosa (Bréb.) Hilse in silicon-starvation synchrony

Summary Exponentially grown cells of the freshwater diatom Navicula pelliculosa (Bréb) Hilse, contained chlorophyll a, chlorophyll c, fucoxanthin, diadinoxanthin, diatoxanthin, neofucoxanthin, -carotene, and an unknown pigment, the absorption spectrum of which is reported. Changes in amounts of chlorophyll a, fucoxanthin and diadinoxanthin were determined during the course of silicon-starvation synchrony carried out in the light or dark. Changes in the rate of chlorophyll a and fucoxanthin syntheses were similar. Synthesis ceased after 5–7 hr of silicon starvation, but recommenced in cultures kept in the light, once silicon was re-introduced. In cultures kept in the dark no significant synthesis was observed after re-introduction of silicon. Diadinoxanthin synthesis continued in the light at all times, although at a lower rate during the silicon-starvation period. In the dark, synthesis of this pigment ceased when cell division stopped, and the amount per unit volume of culture decreased. These results are discussed in relation both to the effect of silicon on the metabolism of the diatom and to the possible function of the carotenoids.Dedicated to Prof. C. B. van Niel on the occasion of his 70th birthday.     

Factors influencing the variability of pigments in the surface sediments of mountain lakes

1. Surface sediment biofilm samples from 82 Pyrenean lakes were analysed for marker pigment composition using high performance liquid chromatography (HPLC). 2. Variability in the pigment composition among lakes was investigated by multivariate statistical analyses using a large data set of factors describing lake chemical, physical, morphological and catchment characteristics. 3. Due to the widely varying light penetration in the lakes, the most significant gradient of pigment composition extended from a benthic to a planktonic signal. The most important pigments in the gradient were alloxanthin (cryptophytes marker pigment, planktonic signal) and diatoxanthin (diatoms marker pigment, benthic signal). The molar ratio between these two marker pigments was positively correlated with lake depth. 4. Chlorophyll‐a preservation was found to be positively related to light penetration and the development of an autothrophic biofilm on the surface sediment and negatively related to decreasing pH and the percentage of alpine meadows in the lake catchments. 5. Zooplankton marker pigments in the surface sediment, including grazing by‐products (e.g. phaeophorbides) and carotenoids (astaxanthin, canthaxanthin, echinenone) incorporated into their tissues, were correlated with the areal abundance of zooplankton. 6. Marker pigments for photosynthetic bacteria, BChl‐e and okenone, were found mainly in relatively shallow lakes with large catchments that are forested, probably because of their higher loading of allochthonous organic matter. 7. The evaluation of a preservation index (Chl‐a expressed as a percentage of a‐phorbins) and the alloxanthin/diatoxanthin ratios throughout the sediment record of mountain lakes can provide evidence of historical changes in the relative importance of planktonic versus benthic primary production and might ultimately be interpreted in terms of climatic or environmental changes.     

Particle size selection in individuals from epifaunal versus infaunal populations of the nereidid polychaete Neanthes succinea(Polychaeta: Nereididae)

Neanthes succinea (Frey & Leuckart, 1847) is a common nereidid polychaete of both epifaunal and infaunal estuarine habitats. The gut contents of individuals collected from two epifaunal and two infaunal habitats are compared. Our a priori expectation was that individuals from epifaunal habitats would be classified as macrophagous with guts indicating carnivory and/or macroalgal herbivory, while individuals from infaunal habitats would be classified as microphagous with guts indicating deposit feeding. At all four locations gut contents indicated deposit feeding with little indication of macrophagous feeding. Average particle sizes for mineral grains did not differ between the four collection sites. For the two infaunal locations mean size of the mineral grains in gut contents was significantly smaller than ambient sediments. In addition to mineral grains, guts contained diatoms, dinoflagellates, macrophytic detritus, protozoan tests, and a variety of metazoans. Our study demonstrates that caution is necessary when inferring feeding type from morphology and that population and habitat specific differences in diet can occur within the same species.     

Algal Diet of Small-Bodied Crustacean Zooplankton in a Cyanobacteria-Dominated Eutrophic Lake

Small-bodied cladocerans and cyclopoid copepods are becoming increasingly dominant over large crustacean zooplankton in eutrophic waters where they often coexist with cyanobacterial blooms. However, relatively little is known about their algal diet preferences. We studied grazing selectivity of small crustaceans (the cyclopoid copepods Mesocyclops leuckarti, Thermocyclops oithonoides, Cyclops kolensis, and the cladocerans Daphnia cucullata, Chydorus sphaericus, Bosmina spp.) by liquid chromatographic analyses of phytoplankton marker pigments in the shallow, highly eutrophic Lake Võrtsjärv (Estonia) during a seasonal cycle. Copepods (mainly C. kolensis) preferably consumed cryptophytes (identified by the marker pigment alloxanthin in gut contents) during colder periods, while they preferred small non-filamentous diatoms and green algae (identified mainly by diatoxanthin and lutein, respectively) from May to September. All studied cladoceran species showed highest selectivity towards colonial cyanobacteria (identified by canthaxanthin). For small C. sphaericus, commonly occuring in the pelagic zone of eutrophic lakes, colonial cyanobacteria can be their major food source, supporting their coexistence with cyanobacterial blooms. Pigments characteristic of filamentous cyanobacteria and diatoms (zeaxanthin and fucoxanthin, respectively), algae dominating in Võrtsjärv, were also found in the grazers’ diet but were generally avoided by the crustaceans commonly dominating the zooplankton assemblage. Together these results suggest that the co-occurring small-bodied cyclopoid and cladoceran species have markedly different algal diets and that the cladocera represent the main trophic link transferring cyanobacterial carbon to the food web in a highly eutrophic lake.     

Degradation of diatom chlorophyll to colourless,non-fluorescing compounds during copepoo grazing

During grazing of the copepodTemora longicornis onThalassiosira eccentrica phaeophorbide-a and several phaeophytins-a appeared in the culture vessels. However, a variable portion of the diatom's chlorophyll-a was degraded to substances that could not be detected either spectrophotometrically or fluorometrically. Chlorophyll-c and fucoxanthin were also lost during diatom cell transit throughTemora's gut. The variability between experiments in the destruction of chl.a to colourless residues is ascribed to differences in coprophagy and in transit time through the copepod gut, processes related to filtering rate and the ambient food concentration. Because the pigment loss is variable, grazing pressure cannot be estimated by simply recording bulk phaeopigment concentrations. The net result of the appearance of a fucoxanthinol-like pigment while all other pigments disappeared is a gradual carotenoid enrichment of the samples during grazing.     

Gut fluorescence in herbivorous copepods: an attempt to justify the method

Recently the gut fluorescence technique has been critisized because of the possible degradation of chlorophyll into nonfluorescent derivatives during passage through copepod guts and changes of the gut passage time with food concentration. Here pigment budgets have been calculated in 6 experiments with Calanus finmarchicus CIV caught 2 km offshore of the Murmansk Marine Biological Institute (the Barents Sea, Dalnije Zelentsi) in September 1992. Copepods were fed with culture of Platymonas viridis at different concentrations. Gut pigment and ingestion rate increased with food concentration in a similar way. On average between 78% and 89% of ingested chlorophyll was recovered in the guts and faecal pellets. No trend for a greater loss of fluorescence at low food concentration than at high was observed. Pigment content of faecal pellets incubated in filtered seawater decreased by 20–30% in the first 7–12 h and by up to 60% in 48 h. The decline of pigment content was accompanied by a rapid bacterial growth (by a factor of 3 in 48 h). Gut passage time increased with decreasing food concentration (from 40 min at 9 µg pigm l^–1 to 64 min at 0.9 µg pigm l^–1). These results together with some data by other authors suggest that the gut fluorescence method can be used to estimate in situ grazing rate providing gut passage time is measured properly and there are no losses of faecal material. However, careful consideration should be given to the previous feeding history of copepods.     

Effect of food preservation on the grazing behavior and on the gut flora of the harpacticoid copepod Paramphiascella fulvofasciata

Harpacticoids owe their ‘reputation’ as primary consumers in aquatic food webs to their substantial grazing on diatoms, thus assuring an efficient energy flow to higher trophic levels. Due to the complex feeding behavior of harpacticoids, the nature and dynamics of diatom-harpacticoid trophic interactions remain poorly understood. In addition, there is a growing interest from aquaculture industry in mass-culturing harpacticoids with algal foods but the labor costs of maintaining algal stock cultures are high. This study focuses on the palatability of preserved diatoms for copepods and considers the possible role of bacterially mediated effects on diatom food uptake.The grazing of Paramphiascella fulvofasciata on a preserved freeze-dried diatom diet was tested and compared to the grazing on fresh cells. P. fulvofasciata assimilated the preserved diet, but assimilation of fresh cells was higher. When both cell types were mixed, no selective feeding was observed. Community fingerprinting of the bacteria associated with diatoms and fecal pellets suggests that the copepod gut flora was modified depending on the food source. Furthermore, the results suggest that the egestion of gut bacteria enriches the microenvironment and this can have an additional influence on the feeding behavior of the copepod.Experimental research using preserved foods must take into account that copepod grazing assimilations of fresh foods are likely to be significantly higher. Yet, the stated high assimilation of the mixed diet, encourages further exploration of the application of preserved ‘balanced’ foods for harpacticoid mass-culturing.     

Pigment destruction by Calanus pacificus: impact on the estimation of water column fluxes

Measurements of gut pigment made prior to fecal pellet productionin Calanus pacificus females and CV copepodites suggest that(i) chlorophyll a and/or its pheopigment derivatives are degradedinto molecules that are not detectable by the standard fluorometrictechnique; and (ii) the percentage of ingested chlorophyll awhich degrades into fluorometrically undetected molecules isnot constant. Thus, measurements of chlorophyll and pheopigmenta in the guts of zooplankton can only yield minimum estimatesof in situ grazing rates. Estimates of the vertical flux ofprimary particulates based on chlorophyll and pheopigment abudgets may also be underestimated.     

Seasonal variation in the biochemical composition of particulate material collected by sediment traps at Signy Island, Antarctica

Particulate material recovered over an 18-month period from sediment traps deployed at a shallow-water nearshore Antarctic site was analysed for photosynthetic pigments, aliphatic hydrocarbons and fatty acids. All components showed a distinct seasonal variation, with high recovery rates during the summer open-water phytoplankton bloom and low rates under winter fast ice. The amount of trapped material differed between the two summers, indicating inter-annual variability of vertical flux associated with differences in the intensity of the summer phytoplankton bloom. Particulate material trapped in summer was dominated by that which originated in diatoms. High recoveries of chlorophyll a, fucoxanthin, n-C_21:6 hydrocarbon, 20:5(n-3) fatty acid and shorter chain (C₁₅–C₂₄) aliphatic hydrocarbons all pointed to a significant summer flux of ungrazed diatoms. There were, however, also signals of zooplankton grazing activity (notably pyrophaeophorbide a), and the presence of C18:4(n-3) and C22:6(n-3) fatty acids suggested a small flux of material from flagellates and other sources. Longer chain n-alkanes (C₂₅–C₃₄) indicative of nanoplankton were detected all year, but there was no significant deposition of zooplankton material in any sample. The major recovery rate of photosynthetic pigments was in late summer (February to April), and the major grazing signal occurred after the peak of the summer diatom bloom. Most of the diatom bloom appeared to settle out from the water column without being grazed. The major seasonal contrast in the biochemistry of the trapped material was the dominance of the diatom signature in summer, and in winter the predominance (but at much lower recovery rates) of material from nanoplankton. Received: 2 March 1998 / Accepted: 12 June 1998     

Microphytobenthic species composition,pigment concentration,and primary production in sublittoral sediments of the Trondheimsfjord (Norway)1

In spring 2005, monthly sampling was carried out at a sublittoral site near Tautra Island. Microphytobenthic identification, abundance (ABU), and biomass (BIOM), were performed by microscopic analyses. Bacillariophyceae accounted for 67% of the total ABU, and phytoflagellates constituted 30%. The diatom floristic list consisted of 38 genera and 94 species. Intact light‐harvesting pigments chl a, chl c, and fucoxanthin and their derivatives were identified and quantified by HPLC. Photoprotective carotenoids were also observed (only as diadinoxanthin; no diatoxanthin was detected). Average fucoxanthin content was 4.57 ± 0.45 μg fucoxanthin · g sediment dry mass^?1, while the mean chl a concentration was 2.48 ± 0.15 μg · g^?1 dry mass. Both the high fucoxanthin:chl a ratio (considering nondegraded forms) and low amounts of photoprotective carotenoids indicated that the benthic microalgal community was adapted to low light. Microphytobenthic primary production was estimated in situ (MPPs, from 0.15 to 1.28 mg C · m^?2 · h^?1) and in the laboratory (MPPp, from 6.79 to 34.70 mg C · m^?2 · h^?1 under light saturation) as ¹⁴C assimilation; in April it was additionally estimated from O₂‐microelectrode studies (MPP_O2) along with the community respiration. MPP_O2 and the community respiration equaled 22.9 ± 7.0 and 7.4 ± 1.8 mg C · m^?2 · h^?1, respectively. A doubling of BIOM from April to June in parallel with a decreasing photosynthetic activity per unit chl a led us to suggest that the microphytobenthic community was sustained by heterotrophic metabolism during this period.     

Does light intensity modify the effect mayfly grazers have on periphyton?

1. A factorial experiment was conducted in artificial outdoor streams to quantify the effects of irradiance (two levels) and two mayfly grazers (four densities of each) on periphytic community structure. The mayflies were Ecdyonurus venosus (Heptageniidae), a grazer using brushing mouthparts, and Baetis spp. (Baetidae) a grazer which uses mandibles and maxilla to scrape and gather periphyton. The experiment ran for 16 days. 2. Grazer densities in channels approximated those existing in a shoreline habitat in the River Sihl, Switzerland. Light treatments were natural (daily mean = 810 μmol m^–2 s^–1) and shaded (daily mean = 286 μmol m^–2 s^–1). 3. Higher irradiance increased total algal abundance by a factor of 4. Algae most affected were prostrate/motile and erect diatoms, filamentous chlorophytes and Hydrurus foetidus. 4. Both species of mayfly reduced periphytic and algal biomass. Mayfly–mayfly interactions, however, were associated with statistical increases in algal biovolume and chlorophyll-a content, indicating that the two grazers may have interfered with one another as their densities increased. The mayfly–mayfly interaction did not influence periphytic ash-free dry mass (AFDM). Light modified the influence of Ecdyonurus such that this mayfly produced greater reductions in algal biovolume under high irradiance. 5. Despite efforts to exclude other grazers, chironomids colonized experimental channels. Chironomid biomass was approximately eight times less than mayflies across treatments and was positively correlated with all measures of periphytic abundance, suggesting that these grazers were responding to periphyton rather than controlling it. Chironomids were also associated with an increase in the abundance of diatoms having a prostrate/motile physiognomy. The only physiognomy to show a negative relationship with chironomid biomass was the thallus type, a form which comprised less than 1% of the algal biovolume across channels. 6. Ecdyonurus and Baetis had distinct influences on algal physiognomy. Ecdyonurus, for example, reduced adnate, stalked and Achnanthes-type physiognomies, but was associated with a significant increase in the abundance of filamentous chlorophytes (primarily Ulothrix sp.). Baetis reduced erect, Achnanthes-type and thallus physiognomies. Neither mayfly influenced the abundance of prostrate/motile diatoms; a physiognomy that comprised 21% of the algae in channels. 7. Light and mayfly interactions affected algal community structure. The interaction of Ecdyonurus with light had a negative effect on erect diatoms, filamentous chlorophytes and the thallus physiognomy, but a positive effect on stalked and Achnanthes-type physiognomies. Baetis interacting with light had a positive effect on adnate diatoms. 8. Although both mayfly taxa influenced periphytic community structure, physiognomy was not a good predictor of algal susceptibility to grazing. The type of substratum to which an alga is attached (detritus or algal filaments vs hard surfaces) and location within the periphytic matrix may be better indicators of vulnerability to grazing than physiognomy.