Marine macroalgae as foods for fishes: an evaluation of potential food quality

Synopsis A revitalized view of feeding by herbivorous marine fishes is sought through two questions. First, What characteristics of major taxa of algae identify them as predictably high or low quality foods? Second, are marine algae valuable foods for fishes which do not mechanically disrupt cell walls and do not harbor specialized enzymes or microbes capable of lysing cell walls? Energy, ash and nutrient content of 16 species of marine algae were employed to assess food quality of fleshy red, green, brown and calcareous red algae. On the basis of ash, calories, total protein and total lipid content, fleshy algae should be superior to calcareous algae as foods for fishes; in addition, green algae should be superior to brown algae and brown algae superior to red algae. When the probable digestibility of storage and extracellular carbohydrates is considered, green and red algae are predicted superior to brown algae as food. Two species of damselfishes (Pomacentridae) from the Gulf of California,Eupomacentrus rectifraenum andMicrospathodon dorsalis, eat red and green algae and ignore brown and calcareous algae. They feed, therefore, in a fashion consistent with predictions based only on algal chemistry. These fishes absorb at least 20–24% of the biomass, 57–67% of the protein, 46–56% of the lipid and 37–44% of the carbohydrate contained in algae eaten in the wild. Since these damselfishes do not masticate their food, it appears that herbivorous fishes can digest major fractions of algal nutrients without mechanical destruction of algal cells.     

Positive feedbacks between bottom-up and top-down controls promote the formation and toxicity of ecosystem disruptive algal blooms: A modeling study

Harmful algal blooms that disrupt and degrade ecosystems (ecosystem disruptive algal blooms, EDABs) are occurring with greater frequency and severity with eutrophication and other adverse anthropogenic alterations of coastal systems. EDAB events have been hypothesized to be caused by positive feedback interactions involving differential growth of competing algal species, low grazing mortality rates on EDAB species, and resulting decreases in nutrient inputs from grazer-mediated nutrient cycling as the EDAB event progresses. Here we develop a stoichiometric nutrient–phytoplankton–zooplankton (NPZ) model to test a conceptual positive feedback mechanism linked to increased cell toxicity and resultant decreases in grazing mortality rates in EDAB species under nutrient limitation of growth rate. As our model EDAB alga, we chose the slow-growing, toxic dinoflagellate Karenia brevis, whose toxin levels have been shown to increase with nutrient (nitrogen) limitation of specific growth rate. This species was competed with two high-nutrient adapted, faster-growing diatoms (Thalassiosira pseudonana and Thalassiosira weissflogii) using recently published data for relationships among nutrient (ammonium) concentration, carbon normalized ammonium uptake rates, cellular nitrogen:carbon (N:C) ratios, and specific growth rate. The model results support the proposed positive feedback mechanism for EDAB formation and toxicity. In all cases the toxic bloom was preceded by one or more pre-blooms of fast-growing diatoms, which drew dissolved nutrients to low growth rate-limiting levels, and stimulated the population growth of zooplankton grazers. Low specific grazing rates on the toxic, nutrient-limited EDAB species then promoted the population growth of this species, which further decreased grazing rates, grazing-linked nutrient recycling, nutrient concentrations, and algal specific growth rates. The nutrient limitation of growth rate further increased toxin concentrations in the EDAB algae, which further decreased grazing-linked nutrient recycling rates and nutrient concentrations, and caused an even greater nutrient limitation of growth rate and even higher toxin levels in the EDAB algae. This chain of interactions represented a positive feedback that resulted in the formation of a high-biomass toxic bloom, with low, nutrient-limited specific growth rates and associated high cellular C:N and toxin:C ratios. Together the elevated C:N and toxin:C ratios in the EDAB algae resulted in very high bloom toxicity. The positive feedbacks and resulting bloom formation and toxicity were increased by long water residence times, which increased the relative importance of grazing-linked nutrient recycling to the overall supply of limiting nutrient (N).     

Stoichiometry, herbivory and competition for nutrients: simple models based on planktonic ecosystems

Models are examined in which two prey species compete for two nutrient resources, and are preyed upon by a predator that recycles both nutrients. Two factors determine the effective relative supply of the nutrients, hence competitive outcomes: the external nutrient supply ratio, and the relative recycling of the two nutrients within the system. This second factor is governed by predator stoichiometry--its relative requirements for nutrients in its own biomass. A model with nutrient resources that are essential for the competing prey is detailed. Criteria are given to identify the limiting nutrient for a food chain of one competitor with the predator. Increased supply of this limiting nutrient increases predator density and concentration of this nutrient at equilibrium, while decreasing the concentration of a non-limiting nutrient. Changes in supply or recycling of a non-limiting nutrient affect only the concentration of that nutrient. Criteria for the invasion of a second prey competitor are presented. When different nutrients limit growth of the resident prey and the invader, increased supply or recycling of the invader's limiting nutrient assists invasion, while increased supply or recycling of the resident's limiting nutrient hinders invasion. If the same nutrient limits both resident and invader, then changes in supply and recycling have complex effects on invasion, depending on species properties. In a parameterized model of a planktonic ecosystem, green algae and cyanobacteria coexist over a wide range of nitrogen:phosphorus supply ratios, without predators. When the herbivore Daphnia is added, coexistence is eliminated or greatly restricted, and green algae dominate over a wide range of supply conditions, because the effective supply of P is greatly reduced as Daphnia rapidly recycles N.     

Genotype × environment interactions,stoichiometric food quality effects,and clonal coexistence in <Emphasis Type="Italic">Daphnia pulex</Emphasis>

The role of stoichiometric food quality in influencing genotype coexistence and competitive interactions between clones of the freshwater microcrustacean, Daphnia pulex, was examined in controlled laboratory microcosm experiments. Two genetically distinct clones of D. pulex, which show variation in their ribosomal rDNA structure, as well as differences in a number of previously characterized growth-rate-related features (i.e., life-history features), were allowed to compete in two different arenas: (1) batch cultures differing in algal food quality (i.e., high vs. low carbon:phosphorus (C:P ratio) in the green alga, Scenedesmus acutus); (2) continuous flow microcosms receiving different light levels (i.e., photosynthetically active radiation) that affected algal C:P ratios. In experiment 1, a clear genotype x environment interaction was determined with clone 1 out-competing clone 2 under high nutrient (i.e., low food C:P) conditions, while the exact opposite pattern was observed under low nutrient (i.e., high C:P) conditions. In experiment 2, clone 1 dominated over clone 2 under high light (higher C:P) conditions, but clonal coexistence was observed under low light (low C:P) conditions. These results indicate that food (nutrient) quality effects (hitherto an often overlooked factor) may play a role in microevolutionary (genotypic) responses to changing stoichiometric conditions in natural populations.     

Effects of solar ultraviolet radiation on the periphyton community in lotic systems: comparison of attached algae and bacteria during their development

The effects of solar ultraviolet radiation (UVR) on the development of a periphyton community were studied in an outdoor artificial stream apparatus. Algal biomass, species composition, and bacterial cell density were measured under full sunlight and non-UVR (photosynthetically active radiation [PAR]-only) conditions. Attachment of algae was detected on days 6–9. Although the chlorophyll-a concentration under non-UVR conditions was 2–4 times that under full sunlight (PAR + UVR) throughout the experiment, neither net algal growth rate nor species composition differed significantly between the two light conditions. The relative carotenoid pigment contents of attached algae in the PAR + UVR condition were 1.1–1.3 times those in the non-UVR condition. Rates of increase of bacterial cell densities under the PAR + UVR condition were depressed by solar UVR for the first few days, although there were no apparent differences in the rates of increase between the light conditions later in the experiment. The small effect of UVR on the development of this periphyton community may be attributable to low UV flux at this study site and to the experimental conditions under which the algae were kept: a high physiological state with high nutrient conditions. Attached bacteria and algae that colonize substrata first are likely to be sensitive to solar UVR, and the negative effects of UVR are mitigated by the development of a periphyton community.     

Photophysiology of Surface Phytoplankton Communities in a Transect from the Mouth of the Peene‐Strom to the Arkona Sea (Baltic)

The potential of surface phytoplankton to withstand photostress was investigated in August 1998 along a transect from the mouth of the Peene‐Strom (Pomeranian Bight) to the open Arkona Sea (Baltic). Photosynthetic efficiency, algal class composition and pigment pattern were determined. Algae were photoinhibited by artificial illumination and the kinetics of recovery were recorded. Under photoinhibitory treatment, algae from the estuary showed a low effective quantum yield but a high potential to recover their maximum photosynthetic efficiency. Contrary to this, the relatively high effective quantum yield of open sea algae under photoinhibitory treatment is accompanied by a low final recovery of maximum photosynthetic efficiency. These phenomena are discussed with respect to nutrient supply, algal class composition and to different strategies of algae to react to light stress. Literature data of summer primary productivity of open sea and coastal algae are compared with our data on electron transport rates. This revealed a low influence of photoinhibitory effects on productivity.     

The dynamics of the restoration of mechanical damage to colonies of the scleractinian coral Porites lutea under conditions of competition with algal settlers for substratum

Natural and anthropogenic catastrophes in the last decades of the 20th century and at the beginning of the new century have evoked significant changes in the biological diversity of coral reefs, viz., the mortality of hermatypic scleractinian corals that are replaced by communities of marine algae and seaweeds. In this connection, the study of the dynamics of algal colonization of a newly formed substratum and competition for the substratum between hermatypic corals and algae is of topical interest. This paper describes the results of investigations conducted in 2003 at the Sesoko Marine Biological Station of Ryukyu University (Okinawa, Japan). Colonies of the massive scleractinian coral Porites lutea were sampled from a fringing reef of Sesoko Island. Damages were inflicted on the upper portions of the colonies, the damaged coral fragments were placed in aquaria and maintained for six months under different light intensities in the absence of herbivorous and corallivorous animals. The experiments showed that under bright (70–90% of incident photosynthetically active radiation, PAR₀) and under moderate light (20–30% of PAR₀), the damaged parts of colonies of about 25 cm² were overgrown by newly formed polyps within six months. Under dim light (2–5% of PAR₀), this period was not sufficient for recovery from lesions. During the first month, the recovery rate of colonies was the highest and depended on the physiological state of the corals and light intensity: in bright and moderate light it was two times and more higher than under low light. During the subsequent months, the recovery rate was significantly lower and depended mainly on the degree of overgrowth of dead portions with algae and on their species composition. Under bright and moderate light, algal colonization of damaged colonies started with the settling of colonial and filamentous diatoms, cyanobacteria and green algae, followed by the formation of the algal turf community. Communities of red calcareous crusts and fleshy algae were mainly formed on damaged corals under low light. A comparison of experimental results showed that damaged colonies maintained in aquaria in the absence of herbivorous and corallivorous animals recovered 1.5 times faster than in the sea. We hypothesize that herbivorous fish have a negative rather than positive influence on the recovery of damaged colonies.     

Mitigation of adverse effects of rising CO2 on a planktonic herbivore by mixed algal diets

Putative future increase in atmospheric CO₂ is expected to adversely affect herbivore growth due to decrease in contents of key nutrients such as nitrogen and phosphorus (P) relative to carbon in primary producers including plant and algal species. However, as many herbivores are polyphagous and as the response of primary producers to elevated CO₂ is highly species-specific, effects of elevated CO₂ on herbivore growth may differ between feeding conditions with monospecific and multiproducer diets. To examine this possibility, we performed CO₂ manipulation experiments under a P-limited condition with a planktonic herbivore, Daphnia , and three algal species, Scenedesmus obliquus (green algae), Cyclotella sp. (diatoms) and Synechococcus sp. (cyanobacteria). Semibatch cultures with single algal species (monocultures) and multiple algal species (mixed cultures) were grown at ambient (360 ppm) and high CO₂ levels (2000 ppm) that were within the natural range in lakes. Both in the mono- and mixed cultures, algal steady state abundance increased but algal P : C and N : C ratios decreased when they were grown at high CO₂. As expected, Daphnia fed monospecific algae cultured at high CO₂ had decreased growth rates despite increased algal abundance. However, when fed mixed algae cultured at high CO₂, especially consisting of diatoms and cyanobacteria or the three algal species, Daphnia maintained high growth rates despite lowered P and N contents relative to C in the algal diets. These results imply that algal diets composed of multiple species can mitigate the adverse effects of elevated CO₂ on herbivore performance, although the magnitude of this mitigation depends on the composition of algal species involved in the diets.     

Paradoxes of enrichment: effects of increased light versus nutrient supply on pelagic producer-grazer systems

Energy-based plant-herbivore models produce the "paradox of enrichment," a destabilizing influence of enrichment on population dynamics. Because many plants change their carbon : nutrient stoichiometry in response to the light : nutrient supply ratio, enrichment with light can cause a mismatch between the elemental compositions of plants and their herbivores. Herbivore growth rates may then decrease with increased light supply, which is termed the "paradox of energy enrichment." I present a stoichiometric phytoplankton-grazer model that accounts for the dynamical vertical light gradient and explore how algal and grazer densities, mineral nutrient concentration, algal nutrient stoichiometry, and system stability respond to enrichment with light (through changes in irradiance, background turbidity, and water column depth) versus enrichment with nutrients. Parameterized for Daphnia, the model produces several "unusual" phenomena: multiple equilibria (with grazers extinct in spite of high algal biomass at one equilibrium), inconsistent light enrichment effects on stability (light enrichment first destabilizes and then stabilizes), and the paradox of energy enrichment. These phenomena are restricted to the low end of realistic nutrient supplies except in very shallow systems, where high sedimentation rates effectively deplete the water column of nutrients. At higher nutrient supplies, light enrichment produces the classical paradox of enrichment, leading first to an increase in grazers at a stable equilibrium and then to algae-grazer oscillations.     

Succession in Stream Biofilms is an Environmentally Driven Gradient of Stress Tolerance

The century-long research on succession has bestowed us with a number of theories, but little agreement on what causes species replacements through time. The majority of studies has explored the temporal trends of individual species in plant and much less so in microbial communities, arguing that interspecific interactions, especially competition, play a key role in community organization throughout succession. In this experimental investigation of periphytic succession in re-circulating laboratory streams, we examined the density and the relative abundance of diatoms and soft algae for 35 days across gradients of low to high nutrient supply (nitrogen + phosphorus) and low to intermediate current velocity (10 vs. 30 cm·s⁻¹). All algal species were classified into trophic groups and morphological guilds, both of which responded more strongly to nutrient than current velocity manipulations, as shown by regression analyses. We concluded that within the manipulated environmental ranges: (1) Succession was a gradient of stress tolerance, driven primarily by nutrient supply and secondarily, by current velocity. Nutrient supply had a qualitative effect in determining whether the contribution of species tolerant vs. sensitive to nutrient limitation would increase through time, while current velocity had a quantitative influence and affected only the rate of this increase. (2) The mechanism of algal succession at a functional level was a neutral coexistence, whereby the tolerant low profile guild maintained high density when overgrown by sensitive species, while sensitive species, constituting mostly the motile and high profile guilds, were neither facilitated nor inhibited by tolerant species but controlled by the environment. It is suggested that the mechanism of succession may depend on the level of biological organization with interspecific interactions giving way to neutral coexistence along the hierarchy from species to functional groups. Considering that the functional makeup is strictly environmentally defined, while species composition reflects local and regional species pools that may exhibit substantial geographic variability, succession is deterministic at a functional level but stochastic at a species level.     

Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

Influence of two Daphnia species on summer phytoplankton assemblages from eutrophic lakes

We conducted grazing experiments to test whether larger-bodiedDaphnia pulicaria have a different effect from smaller-bodiedDaphnia galeata mendotae on the composition of summer algalassemblages in eutrophic lakes. Three separate cubitainer experimentswere run for 5 days in a replicated factorial design utilizingtwo algal community types and the two Daphnia species. Inorganicphosphorus and nitrogen were added to prevent nutrient limitationof the algae. Both edible and inedible size fractions of chlorophylla increased in cubitainers without Daphnia spp. Grazer additionusually resulted in a reduction in edible chlorophyll; reductionswere greater in D.pulicaria cubitainers. Grazing by Daphniaspp. on presumed inedible chlorophyll was variable. Algal sizewas not always a good predictor of grazeability. The resultsof this study indicate that D.pulicaria, because of its greaterfiltration potential and ability to ingest larger particles,provides a stronger control on inedible-sized algae when comparedto equal numerical densities of D.g.mendotae. However, Aphanizomenonincreased as a response to heavy grazing pressure by D.pulicariaon other algal species. This suggests that biomanipulation effortsthat promote large-bodied Daphnia may not produce desirableresults if nutrient inputs remain high.     

Effects of avian grazing on the algal community and small invertebrates in the rocky intertidal zone

In this study of a rocky intertidal habitat in northern Japan, feeding by avian consumers had significant effects on algal assemblages and small herbivorous invertebrates. The effects of the birds on algae were different from those of invertebrate grazers such as urchins and gastropods. The abundance of the dominant algal species decreased during the grazing period, increased again after the grazing period, and indirectly affected algal species richness and evenness. Avian grazing also decreased the density of tube-dwelling amphipods on the dominant alga, but did not change the density of mobile and free-living isopods. These results suggest that avian grazers may act as habitat modifiers rather than exploitative competitors for the small herbivorous crustaceans. Avian herbivores consumed only the upper parts of large algal fronds, apparently reducing the amount of suitable microhabitat for the small herbivorous crustaceans, which are subject to a variety of physical or biological stress. Thus, avian herbivores function as ecosystem engineers, regulating community structure in a manner different to invertebrate herbivores in rocky intertidal habitats.     

Harmful algal blooms of allelopathic microalgal species: The role of eutrophication

The ability of certain harmful algal species to produce and release chemicals that inhibit the growth of co-occurring phytoplankton species, here considered as allelopathy, is closely associated with competition for limiting nutrient resources. Many phytoplankton cells are known to release elevated amounts of organic compounds under nutrient limitation. Eutrophication alters the nitrogen-to-phosphorus balance and, when nutrient availability is unbalanced, nutrient limitation may result. Algal species that can compete successfully for available growth-limiting nutrient(s) have the potential to become dominant and form blooms. The stress conditions imposed by the shifted nutrient supply ratios can, in some algae, stimulate production of allelochemicals that inhibit potential competitors. Thus, under cultural eutrophication, altered nutrient (N, P) ratios and limiting nutrient supplies can stimulate increased production of allelochemicals, including toxins, by some algal species and accentuate the adverse effects of these substances on other algae. Future investigation on the characterization of the chemical compounds involved in the allelopathic process are needed to advance the study of the mode of action of phytoplankton allelochemicals.     

Modeling the bacterial contribution to planktonic community respiration in the regulation of solar energy and nutrient availability

In planktonic ecosystems, algae and bacteria exhibit complex interrelationships, as algae provide an important organic matter source for microbial growth while microbial metabolism recycles limiting nutrients for algae in a loose commensalism. However, algae and bacteria can also compete for available nutrients if supplies of organic matter are sufficient to satisfy bacterial demand. We developed a stoichiometrically explicit model of bacteria–algae interactions that incorporated realistic assumptions about algal light and nutrient utilization, algal exudation of organic matter, and bacterial processing of organic matter and nutrients. The model makes specific predictions about how the relative balance of algae and bacteria should change in response to varied nutrient and light availability seen in lakes and oceans. The model successfully reproduces published empirical data and indicates that, under moderate nutrient supply, the bacterial percentage of total respiration should be maximal at intermediate light intensity.     

Light and nutrients regulate energy transfer through benthic and pelagic food chains

The amount of energy flowing to top trophic levels depends on primary production and the efficiency at which it is converted to production at each trophic level. In aquatic systems, algal production is often limited by light and nutrients, and the nutritional quality of algae depends on the relative balance of these two resources. In this study, we used a mesocosm experiment to examine how light and nutrient variation affected food chain efficiency (FCE, defined as the proportion of primary production converted to top trophic level production), using a food web with benthic and pelagic food chains. We also related variation in benthic and pelagic efficiencies to the nutritional quality of primary producers, i.e. carbon:nitrogen:phosphorus stoichiometry. As predicted, pelagic and benthic FCEs were highest under low light/high nutrient conditions, the treatment with the best algal food quality, i.e. the lowest C:nutrient ratios. Pelagic FCE and pelagic herbivore efficiency (HE_P) were more responsive than benthic FCE to variation in light and nutrients. Furthermore, pelagic FCE and HE_P were highly correlated with algal C:P, suggesting ‘carryover effects’ of algal food quality on carnivores (larval fish) via effects on herbivore (zooplankton) quality. Benthic (tadpole) production was primarily explained by primary production rate, suggesting food quantity rather than quality drives their production. However, benthic FCE was also highest at low light/high nutrients and was significantly correlated with food quality. The stronger effect of food quality in mediating pelagic compared to benthic efficiencies, is consistent with differences in the stoichiometric mismatches between algae and consumers. Pelagic FCE and HE_P were more likely to be P‐limited, whereas benthic FCE was more likely N‐limited. This study is the first to examine both pelagic and benthic FCE within the same system, and highlights the importance of differential consumer needs in determining how food quality affects energy transfer efficiency.     

Promotion of harmful algal blooms by zooplankton predatory activity

The relationship between algae and their zooplanktonic predators typically involves consumption of nutrients by algae, grazing of the algae by zooplankton which in turn enhances predator biomass, controls algal growth and regenerates nutrients. Eutrophication raises nutrient levels, but does not simply increase normal predator-prey activity; rather, harmful algal bloom (HAB) events develop often with serious ecological and aesthetic implications. Generally, HAB species are outwardly poor competitors for nutrients, while their development of grazing deterrents during nutrient stress ostensibly occurs too late, after the nutrients have largely been consumed already by fast-growing non-HAB species. A new mechanism is presented to explain HAB dynamics under these circumstances. Using a multi-nutrient predator-prey model, it is demonstrated that these blooms can develop through the self-propagating failure of normal predator-prey activity, resulting in the transfer of nutrients into HAB growth at the expense of competing algal species. Rate limitation of this transfer provides a continual level of nutrient stress that results in HAB species exhibiting grazing deterrents protecting them from top-down control. This process is self-stabilizing as long as nutrient demand exceeds supply, maintaining the unpalatable status of HABs; such events are most likely under eutrophic conditions with skewed nutrient ratios.     

A conceptual ecological model to aid restoration of Cootes Paradise Marsh,a degraded coastal wetland of Lake Ontario,Canada

An ecological model is derived from recent studies, based on 60 years of empirical observations and experimental data, that conceptualizes how Cootes Paradise Marsh was transformed from a lush emergent marsh with considerable ecological diversity in all trophic levels, to one that is currently turbid, devoid of vegetation, and dominated by a few exotic plant and fish species. This conceptual model contains 17 key components that interact and contribute to the overall unhealthy state of the marsh. The most influential component is high water level which caused the initial loss of emergent vegetation in the 1940s and 1950s. In the absence of plants to attenuate sediment and assimilate nutrients, the marsh became turbid and windswept, and this led to the disappearance of submergent vegetation over the next two decades. Currently, high water turbidity is being maintained by wind re-suspension, high sediment loading from the watershed during the summer, high algal biomass resulting from excessive nutrient loads from sewage effluent and surface runoff, and the feeding and spawning activities of a very large population of common carp ( Cyprinus carpio). Due to vegetation loss, the substrate has become mostly loose sediment that is no longer suitable for the diverse assemblage of aquatic insect larvae that lived on the plants and detrital material in the 1940s. Benthic grazers have been kept in low abundances due to predation by benthivorous carp; consequently, epiphytic algae have proliferated and further contribute to light limitation of macrophytes. High nutrient loadings contribute to high diurnal fluxes in dissolved oxygen levels that tend to select against less tolerant organisms such as insect larvae (other than chironomids) and piscivores (northern pike and largemouth bass). Without piscivores in the marsh, the planktivores have become dominant and have virtually eliminated all of the large herbivorous zooplankton (e.g., Daphnia), except for a few pockets in the marsh inlets close to residual macrophyte beds. Because of the dominance of small-bodied inefficient grazers (rotifers and small cladocerans), algal biomass is high, and the community has a large proportion of heterotrophic forms that tolerate low light environments. This ecological model suggests that the current turbid un-vegetated state of Cootes Paradise may be very stable. It will persist as long as water levels remain unfavorable for natural re-colonization by the emergent flora, and/or water turbidities remain sufficiently high to suppress the growth of submergent vegetation. Using this conceptual model, I developed a model of how Cootes Paradise Marsh may have functioned as a healthy marsh prior to the 1940s, and use these models as a basis to explore a number of restoration and management options and discuss their implications on the aquatic foodweb.     

Temperature indirectly affects benthic microalgal diversity by altering effects of top–down but not bottom–up control

Understanding the ecological mechanisms that underlie species diversity decline in response to environmental change has become an urgent objective in current ecological research. Not only direct (lethal) effects on single species but also indirect effects altering biotic interactions between species within and across trophic levels comprise the driving force of ecosystem change. In an experimental marine benthic microalgae–grazer system we tested for indirect effects of moderate temperature change on algal diversity by manipulation of temperature, nutrient supply and grazer density. In our model system warming did not exert indirect effects on microalgal diversity via effects on resource competition. However, moderate warming strengthened consumer control and thereby indirectly affected algal community structure which ultimately resulted in decreased diversity. Only in low temperature and low nutrient regimes did the antagonizing mechanisms of bottom–up and top–down regulation establish a balancing effect on algal diversity within 29 days (corresponding to 15–29 algae generations). Effects of thermal habitat change did not appear before 9–18 algae generations, which points to the relevance of longer‐term experiments and ecological monitoring in order to separate transient biotic responses and subtle changes of community dynamics in consequence to global change.     

Use of dried algae as a food source for zooplankton growth and nutrient release experiments

Zooplankton growth and nutrient recycling are key processes in the operation of pelagic food webs. Most studies investigating these processes rely on complex methods and often require extensive laboratory facilities. Here we introduce a technique for preserving algae by rapid drying for later use in laboratory- or field-based growth and nutrient recycling experiments. Chemostat-grown Scenedesmus acutus was rapidly dried for later experiments evaluating its nutritional composition, suitability for animal growth and potential for use in nutrient release experiments. Reconstituted dried algae had slightly lower nitrogen (N), Phosphorus (P) and protein content (% dry weight) than fresh algae, but lipid content did not differ and elemental ratios were in the range considered to indicate favorable food quality. These elemental and biochemical differences did not appear functionally important, as Daphnia magna grew identically on fresh and dried food. Freeze-dried S.acutus did not work as an alternative to oven drying as it resulted in 100% mortality. NH4 and PO4 concentrations did not change over 24 h when dried algae were resuspended in normal media or boiled lake water. However, concentrations of PO4 decreased over 24 h, suggesting chemical adsorption of PO4 to the dried algae and reinforcing the need for animal-free controls in nutrient release experiments using this approach. N and P release rates for D.magna and natural zooplankton communities were estimated using dried algae, and values were comparable to published ones. Thus, dried algae may be a useful, simple technique for studying food quality and nutrient release in environments where maintaining active algal cultures may not be practical and a constant supply of consistent quality food is needed.