Low temperature and <Emphasis Type="Italic">Daphnia</Emphasis>-associated infochemicals promote colony formation of <Emphasis Type="Italic">Scenedesmus obliquus</Emphasis> and its harvesting

Objective

To explore the combined effects of temperature and Daphnia-associated infochemicals on colony formation of Scenedesmus obliquus to faciliate harvesting the algal biomass.

Results

A three-parameter modified Gaussian model fitted the changes of the number of cells per particle in S. obliquus induced by Daphnia culture filtrate well under any temperature. Decreases in temperature enhanced the induced–colony formation of Scenedesmus. The maximum colony size at 15–25 °C was significantly larger than those at 30–35 °C. An additional 1 or 2 days at low temperature was needed to reach the maximum colony size, which indicates the best harvest time for algal biomass.

Conclusion

Induced-colony formation of Scenedesmus by Daphnia culture filtrate at 15–25 °C is recommended to settle algal cells. This condition facilitates harvesting the biomass.

     

The use of natural infochemicals for sustainable and efficient harvesting of the microalgae <Emphasis Type="Italic">Scenedesmus</Emphasis> spp. for biotechnology: insights from a meta-analysis

Open raceway ponds are regarded as the most economically viable option for large-scale cultivation of microalgae for low to mid-value bio-products, such as biodiesel. However, improvements are required including reducing the costs associated with harvesting biomass. There is now a growing interest in exploiting natural ecological processes within biotechnology. We review how chemical cues produced by algal grazers induce colony formation in algal cells, which subsequently leads to their sedimentation. A statistical meta-analysis of more than 80 studies reveals that Daphnia grazers can induce high levels of colony formation and sedimentation in Scenedesmus obliquus and that these natural, infochemical induced sedimentation rates are comparable to using commercial chemical equivalents. These data suggest that natural ecological interactions can be co-opted in biotechnology as part of a promising, low energy and clean harvesting method for use in large raceway systems.     

Grazer density-dependent response of induced colony formation of Scenedesmus obliquus to grazing-associated infochemicals

To investigate the effects of infochemicals from Daphnia magna reared at different densities on the growth and morphological development of Scenedesmus obliquus, we cultured S. obliquus with filtrates from D. magna with a density of 0, 6, 30, 150, 300, and 600 ind L⁻¹, respectively. Results showed no significant differences in S. obliquus growth rates among the treatments with different Daphnia densities. On days 3 and 5, the proportion of eight-celled colonies in the treatments constituted 30% of the S. obliquus populations. Importantly, the number of cells per particle increased significantly with increasing density of Daphnia, indicating a grazer density-dependent response. Specifically, the density-response relationship followed a rectangular hyperbolic model in the first five days of treatment, and then switched to a linear model, which implies that the relationship between the inducible colony size of S. obliquus and Daphnia density may also be dependent upon incubation time.     

Does microcystin disrupt the induced effect of Daphnia kairomone on colony formation in Scenedesmus?

In natural aquatic system, Scenedesmus and Microcystis species usually coexist. Microcystins are released into water after lysis of Microcystis cells during the collapse of heavy blooms. The released toxins can then come into contact with a wide range of aquatic organisms. In this study, we used filtered Daphnia test water containing kairomone from Daphnia magna to stimulate the inducible colony formation in Scenedesmus obliquus under microcystin-contaminated system, to examine how microcystin affects the induced effect of Daphnia kairomone on colony formation in S. obliquus. The results showed neither microcystin nor Daphnia kairomone affected the growth of S. obliquus. Microcystin neither promoted nor impaired the overall Daphnia-induced colony formation in S. obliquus, except reducing the proportion of eight-celled colonies on day 2, indicating that the effect of microcystin was just short-term and in general did not disrupt grazer-induced colony formation of S. obliquus.     

No difference in colony formation of Scenedesmus obliquus exposed to lakes with different nutrient levels

The algal genus Scenedesmus is famous for its highly phenotypic plasticity in response to various environmental factors. In laboratory, axenic cultures of Scenedesmus often fail to form colonies and remain in unicellular morph. To examine whether unicellular Scenedesmus can form colonies after exposure to natural lakes, dialysis bags and plastic bottles which contained the precultured Scenedesmus obliquus were exposed in two lakes with different nutrient levels for four weeks. Results showed S. obliquus grew well in dialysis bags but not in plastic bottles; S. obliquus can form colonies when incubated in dialysis bags which allow exchange with the aquatic environment of the substances, regardless of the nutrient levels of the two lakes. However, no colonies were observed in S. obliquus incubated in plastic bottles exposed in both lakes. This suggested that active growth and zooplankton infochemicals contributed together to the colony formation of S. obliquus exposed in situ environment.     

Effects of Daphnia ‐Associated Infochemicals on the Morphology,Polysaccharides Content and PSII‐Efficiency in Scenedesmus obliquus

We investigated the effects of infochemicals from Daphnia carinata on the morphology, polysaccharides yield and PSII‐efficiency in Scenedesmus obliquus. Infochemicals released from D. carinata induced colony formation in S. obliquus. The contents of soluble extracellular polysaccharides, bounded extracellular polysaccharides, and the total polysaccharides per cell in the induced colonies of S. obliquus increased significantly relative to those of the unicells, which indicated that Daphnia ‐associated infochemicals could also stimulate S. obliquus to increase the synthesis of extracellular polysaccharides. The increased extracellular polysaccharides may play an important role in cementing S. obliquus cells together to form colonies. In addition, no significant differences in growth, the maximal efficiency of PSII photochemistry (F_v/F_m), and the effective quantum yields of PSII (Φ_PSII) were detected between unicellular and induced colonial populations, which showed that the cost of induced colony formation was not reflected on these indices. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Influence of <Emphasis Type="Italic">Daphnia</Emphasis> infochemicals on functional traits of <Emphasis Type="Italic">Microcystis</Emphasis> strains (Cyanobacteria)

We conducted a laboratory experiment to investigate the influence of Daphnia infochemicals on growth rate, microcystin production, colony formation and cell size of eight Microcystis strains isolated from two lakes. The strains were characterized genetically by their 16S-23S rDNA ITS sequence. The experiment was composed of four treatments: (1) a control using filtered WC medium, (2) addition of Scenedesmus obliquus culture medium filtrate, (3) addition of Daphnia magna culture medium filtrate and (4) addition of sodium octyl sulphate, a commercially available Daphnia infochemical. Our results showed that sympatric strains differed strongly for the measured functional traits, while no correlations between traits were found. Between-strain differences in growth rate, microcystin production, colony formation and cell size were generally larger than the differences in phenotypes observed between treatments. Despite this, several strains reacted to the infochemicals by changing functional trait values. Daphnia culture medium filtrate and, to a lesser extent, sodium octyl sulphate had a negative influence on the growth rate of half of the strains and stimulated microcystin production in one strain, but the latter effect was not Daphnia-specific as Scenedesmus culture medium filtrate had the same effect. Daphnia culture medium filtrate also induced colony formation in one strain. Our data suggest that Daphnia infochemicals generally have a weak influence on growth rate, microcystin production and colony formation of Microcystis strains as compared to the inter-strain variability, while existing inducible effects are highly strain-specific.     

Possible toxic effects on Daphnia resulting from the green alga Scenedesmus obliquus

We cultured individuals of two Daphnia species and their hybrid on two different algae, Scenedesmus obliquus and Chlamydomonas globosa, in different concentrations. Our results suggest that culture conditions of S. obliquus can be such that the algal cells become toxic to Daphnia     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Combined effects of four Microcystis aeruginosa strains and Scenedesmus obliquus concentrations on population dynamics and resting egg formation of two Daphnia species

We examined the hypothesis that toxic effects of Microcystis aeruginosa to population dynamics, ephippial production, and resting egg formation of Daphnia were restricted by food quality levels. Four unicellular, toxic M. aeruginosa strains with contrasting concentration of microcystin and two Daphnia species were used in this study. The number of offspring at first reproduction, population densities, and maximum population growth rate of two Daphnia species were lower at the highest toxic M. aeruginosa 7820 treatment compared with other treatments under two concentrations of Scenedesmus obliquus. The maximum number of offspring at first reproduction of two Daphnia species appeared in the higher toxic M. aeruginosa 526 strain at the high concentration of S. obliquus. The effects of M. aeruginosa strains, S. obliquus, and their combination on the number of offspring at first reproduction and the maximum population growth rate of two Daphnia species were significant (P < 0.001). Two Daphnia species could not reproduce in the highest toxic M7820 treatment under two concentrations of S. obliquus. They had lower population size and maximum population growth rate in the higher toxic M526 treatment at the low concentration of S. obliquus, but they were higher at the high concentration of S. obliquus. This result suggests that high S. obliquus concentration could relieve the toxicity of M. aeruginosa to Daphnia, and Daphnia could utilize the lower toxic Microcystis as food. The cumulative ephippia numbers of two Daphnia species were more at the high concentration of S. obliquus than those at the low concentration. The percentage of ephippia containing no resting eggs of two Daphnia species was evidently higher at the low concentration of S. obliquus but was lower at the high concentration of S. obliquus. Our results indicated that the cumulative ephippia numbers of Daphnia were population density dependent at the high-level food, and the productions of ephippia in Daphnia were significantly controlled by microcystin concentration at the low-level food.     

Consumer-dependent responses of lake ecosystems to nutrient loading

The nutrient loading concept proposes that algal biomass, waterclarity and the processes of lake eutrophication are a functionof nutrient loading. We hypothesized that grazers play an importantrole in determining the impacts of nutrient loading on algalbiomass and water clarity, and the overall eutrophication process.To test how the contrasting grazer communities modify the fateof nutrients, we added nutrients (nitrate and phosphate) ata known loading rate to four large enclosures, but in two ofthe four enclosures large cladoceran grazers (Daphnia >1mm mean length) were allowed to develop by removing the planktivorousfish. In the remaining two enclosures, the development of largeDaphnia was prevented by adding planktivorous fish. The concentrationsof epilimnetic total phosphorus (TP) increased at a similarrate in all four enclosures. However, the daily accumulationof added phosphate into the participate or planktonic forms,especially into plankton <20 µm, was three times fasterwhen large Daphnia were absent than when large Daphnia wereabundant. In the enclosures with large Daphnia, added phosphatewas accumulated in the dissolved pool instead. At a constantnutrient loading, algal biomass (chlorophyll a) increased fourtimes faster in the enclosures without large Daphnia than inthose with large Daphnia. Similarly, Secchi depth declined from3.5 to <1 m when Daphnia were absent, but did not declinewhen Daphnia were common. Our results demonstrate that the samenutrient loading and the resultant increase in epilimnetic TPdo not produce the same trophic conditions, as indicated byalgal biomass and water clarity, if the grazers of the majorassimilators of nutrients (the fraction of plankton edible toDaphnia) are different. We suggest that stratified lake ecosystemshaving functionally dominant large grazer communities may beless prone to eutrophication than those lacking large grazers.Consistent with the nutrient loading concept, epilimnetic concentrationsof phosphorus increase proportionately with increased loadingof phosphorus, but the trophic conditions of ecosystems indicatedby algal biomass and water clarity do not follow the same patternsunder contrasting conditions of grazer communities. We suggestthat models predicting algal biomass from loading rates shouldaccount for the role of grazers.     

PSII-efficiency,polysaccharide production,and phenotypic plasticity of Scenedesmus obliquus in response to changes in metabolic carbon flux

To investigate the response of Scenedesmus obliquus to changes in metabolic carbon flux, S. obliquus was cultured in medium with different concentrations of glyoxylate over 9 days. Results showed that growth rates were not affected in the lower concentration glyoxylate (0.25 and 0.5 mM). However, growth rate of S. obliquus was inhibited in the higher concentration glyoxylate (0.85 and 1.25 mM) during the early phase before recovering at higher densities. Changes in growth rates in different glyoxylate concentrations were in line with changes in F_v/F_m and Φ_PSII. Colony formation was observed in S. obliquus in the four glyoxylate treatments. As a consequence, the mean number of cells per particle of S. obliquus in the glyoxylate treatments were significantly higher than those in the control. The total polysaccharide content of S. obliquus cells increased with increased glyoxylate concentrations. The increased glyoxylate-stimulated polysaccharide levels were directly correlated with colony size of S. obliquus.     

Influence of photoperiods on the growth rate and biomass productivity of green microalgae

The effect of different photoperiods: 24 h illumination and a 12:12-h light/dark (12L:12D) cycle on the growth rate and biomass productivity was studied in five algal species: Neochloris conjuncta, Neochloris terrestris, Neochloris texensis, Botryococcus braunii and Scenedesmus obliquus. The green microalgae examined differ in the reproduction mode. Continuous illumination stimulated the growth of B. braunii and S. obliquus more effectively than the growth of the microalgal species from the genus Neochloris. However, under shorter duration of light of the same intensity (12L:12D cycle), the growth of all the three species of Neochloris was stimulated. Under continuous illumination, the specific growth rate in the first phase of B. braunii and S. obliquus cultures was higher than the growth rate of Neochloris, whereas under the 12L:12D cycle, the specific growth rate of all the three Neochloris species was generally higher than that in B. braunii and S. obliquus. As a result, the light regime influenced algal biomass productivity differently. The maximum biomass productivity was obtained in B. braunii and S. obliquus cultures carried out at continuous illumination. All the Neochloris species produced biomass more efficiently at the 12L:12D cycle, which was two–threefold higher than that of B. braunii and S. obliquus. The unicellular species of the green microalgae from the genus Neochloris, examined for the first time in this study, are promising prospective objects for algal biotechnology.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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Is bigger better? A possibility for adaptation of <Emphasis Type="Italic">Daphnia</Emphasis> to filamentous cyanobacteria in the face of global warming

Body size is a key determinant of fitness in Daphnia. Bigger size means higher feeding efficiency and reproduction. However, filamentous cyanobacteria have a more detrimental effect on larger daphnids. Predicted global warming is expected to reduce size of Daphnia and facilitate frequent occurrence of cyanobacteria. Therefore, we asked two questions: (i) does elevated temperature cause reduced size of daphnids, and (ii) is temperature-dependent size reduction adaptive in an environment dominated by filamentous cyanobacteria? To address these issues, we obtained 8 clones of the Daphnia longispina complex from artificially heated lakes and 8 clones from control lakes, and exposed them to a favorable food Scenedesmus obliquus and a mixture of S. obliquus and the filamentous cyanobacterium Cylindrospermopsis raciborskii in life table experiments at 16, 20, and 24°C. Individuals from heated lakes attained larger body size than those from control lakes. Moreover, exposure to the filamentous cyanobacterium led to reduced fecundity of Daphnia from the non-heated lakes and did not affect reproduction of Daphnia from the heated lakes. We conclude that Daphnia display some evolutionary adaptations to cope with filamentous cyanobacteria, linked to long-term exposition to elevated temperature. Our results violate broadly accepted assumptions that ectotherms are smaller in warmer environments.     

Predaceous insects may limit algal grazers in high-nutrient environments: implications for wastewater remediation in open bioreactors

Algal biofuel has potential as a source of renewable fuel and a tool for wastewater remediation. Open algal bioreactors fertilized with wastewater can have net energy gain but are vulnerable to colonization by algal grazers. However, colonizing predaceous insects may limit grazer impacts on algae. Here, we investigate the effects of grazers, predators, and invading algae species on algal production and community structure in high-nutrient environments. First, we grew diverse algal assemblages in treated municipal wastewater in a greenhouse with Daphnia grazers and different insect predators that were added experimentally. When Daphnia were present without predators, they eliminated suspended algae. But, dragonfly larvae [Odonata: Libellulidae] and backswimmers [Hemiptera: Notonectidae], but not larval diving beetles [Coloeoptera: Dytiscidae], suppressed Daphnia allowing suspended algae to persist. Second, we grew Chlorella algae in field tanks that were open or protected from natural invertebrate colonization and half the tanks received wild-collected plankton in a factorial design. Mosquito larvae [Culex sp.] readily colonized open tanks and reduced algal mass and dissolved phosphorus concentrations. Colonist addition to open tanks shifted algal functional and taxonomic composition but did not impact suspended algal production. Our study indicates that large numbers of grazer individuals can rapidly colonize open bioreactors. Experimentally added and naturally colonizing grazers altered algal community structure and reduced algal standing crops but may also aid in nutrient removal from wastewater-fed bioreactors. Effective operation of open algal bioreactors must consider cultivated algae species’ vulnerability to competition and local grazers as well as the ability of potential predators to both naturally disperse into bioreactors and to control grazers.     

Changes in food quality of the green alga Scenedesmus induced by Daphnia infochemicals: biochemical composition and morphology

1. The effects of Daphnia infochemicals on the morphology and biochemical composition of Scenedesmus were studied and subsequently their influence on Daphnia life history. Three species of Scenedesmus ( S. acutus, S. obliquus and S. subspicatus ) were tested for Daphnia -induced colony formation. Life history experiments were performed with Daphnia cucullata and D. magna feeding on unicellular or colonial S. acutus.
2. Colony formation was promoted when S. acutus and S. obliquus were exposed to filtered water from a D. magna culture. S. subspicatus did not form colonies when exposed to culture water, but showed some colony formation when exposed to live D. magna.
3. No clear differences were found in total lipid, protein or carbohydrate content between the unicellular and colonial ecomorphs of the three Scenedesmus species. However, fatty acid (FA) concentration and composition were changed. Total FA as a percentage of frozen dry weight (DW) were increased in colonies. The ratio of monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA) and the ratio of ω3 : ω6 fatty acids decreased in colonies.
4. A short-term grazing experiment with three size classes of Scenedesmus revealed lower clearance rates for small Daphnia when feeding on large colonial Scenedesmus.
5. Life history parameters of both Daphnia species were influenced by food type. The intrinsic rate of population increase ( r) was significantly lower for daphnids feeding on colonies.
6. The negative influence of colonial Scenedesmus on population growth of Daphnia seems to be determined by algal morphology rather than biochemical composition.     

Linking Cascading Effects of Fish Predation and Zooplankton Grazing to Reduced Cyanobacterial Biomass and Toxin Levels Following Biomanipulation

Eutrophication has been one of the largest environmental problems in aquatic ecosystems during the past decades, leading to dense, and often toxic, cyanobacterial blooms. In a way to counteract these problems many lakes have been subject to restoration through biomanipulation. Here we combine 13 years of monitoring data with experimental assessment of grazing efficiency of a naturally occurring zooplankton community and a, from a human perspective, desired community of large Daphnia to assess the effects of an altered trophic cascade associated with biomanipulation. Lake monitoring data show that the relative proportion of Daphnia spp. grazers in June has increased following years of biomanipulation and that this increase coincides with a drop in cyanobacterial biomass and lowered microcystin concentrations compared to before the biomanipulation. In June, the proportion of Daphnia spp. (on a biomass basis) went from around 3% in 2005 (the first year of biomanipulation) up to around 58% in 2012. During months when the proportion of Daphnia spp. remained unchanged (July and August) no effect on lower trophic levels was observed. Our field grazing experiment revealed that Daphnia were more efficient in controlling the standing biomass of cyanobacteria, as grazing by the natural zooplankton community never even compensated for the algal growth during the experiment and sometimes even promoted cyanobacterial growth. Furthermore, although the total cyanobacterial toxin levels remained unaffected by both grazer communities in the experimental study, the Daphnia dominated community promoted the transfer of toxins to the extracellular, dissolved phase, likely through feeding on cyanobacteria. Our results show that biomanipulation by fish removal is a useful tool for lake management, leading to a top-down mediated trophic cascade, through alterations in the grazer community, to reduced cyanobacterial biomass and lowered cyanobacterial toxin levels. This improved water quality enhances both the ecological and societal value of lakes as units for ecosystem services.     

Temporal patterns of grazer-periphyton interactions in laboratory streams

SUMMARY. 1. The snail Juga silicuta (500 m^?2) and the caddisfly Dicosmoecus gilvipes (50 m^?2) were introduced into separate laboratory streams on days 1, 9, 16 and 28 of algal development. The mayfly Baetis spp. (500 m^?2) was introduced on days 1 and 16, and two streams did not receive grazers. We assessed the interaction between succession in the pcriphyton, herbivore type and time of encounter in a 40-day experiment. 2. In ungrazed streams, the chlorophyte Scenedesmus obliquus was the most abundant early colonizer. The relative abundance of diatoms increased after day 9, and at day 40 the algal assemblage consisted of a thick mat of diatoms and S. obliquus with an overstorey of filaments of the chlorophyte Stigeoclonium tenue. In general, introductions of grazers at any stage altered this pattern by removing biomass, accelerating the replacement of S. obliquus by diatoms, and suppressing the growth of filaments. Grazing also reduced the relative abundance of the larger diatom Nitzschia oregona but increased the relative abundance of the smaller adnate diatoms Nitzschia frustulum var. perpusilla and Navicula minima. 3. Dicosmoecus decreased algal biomass and altered successional trajectories to a greater degree than either Juga or Baetis. Dicosmoecus rapidly grazed the entire substrate, whereas Juga and Baetis only cleared patches in the assemblages. Little alteration in algal development was observed in the Baetis streams after day 16, probably because (he periphyton assemblages attained a size and structure that prevented effective grazing by Baetis. 4. The patchy grazing patterns of Juga and Baetis resulted in more diverse algal assemblages than either the Dicosmoecus grazed or ungrazed streams. In natural streams, the temporal and spatial pattern of grazing relative to the developmental stage of the periphyton may contribute to maintaining a mosaic of algal patches in different serai stages.