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In recent years most studies of the benthic microbial food web have either been descriptive or were restricted to the measurement of within sediment process rates. Little is known about benthic-pelagic coupling processes such as recruitment. We, therefore, developed an ex situ core incubation procedure to quantify the potential for microbial recruitment from the benthos to the pelagic in an acidic mining lake, Mining Lake 111 (ML 111; pH 2.6), in eastern Germany. Our data suggest that considerable zooplankton recruitment from the benthos takes place. Heliozoan and rhizopod recruitment in both summer and winter sediment cores was highest when they were incubated at 20°C. Maximum heliozoan recruitment was 23 (± 9 s.e.) individuals cm-2 d-1 (40% initial standing stock daily) in the winter 20°C incubation. Maximum rhizopod recruitment was 6 (± 2 s.e.) individuals cm-2 d-1 in the summer 20°C incubation. Little or no recruitment was apparent for either taxa when winter cores were incubated at 5°C, implying a temperature cue. Conversely, the rotifer, Cephalodella hoodi, exhibited a maximum recruitment of 6 (± 2 s.e.) individuals cm-2 d-1 during the winter 5°C incubation, representing 30% of initial standing stock daily, but little recruitment when incubated at 20°C. Cephalodella may have responded to an increased winter benthic food supply; in situ winter Chl a concentrations in the benthos were 3.4 times higher than those in the summer. The importance of this was reinforced by the poor pelagic food supply available in ML 111. In situ, Heliozoa, rhizopods and Cephalodella were first observed in the epilimnion of ML 111 in spring or early summer, suggesting active or passive recruitment following lateral transport from littoral sediments. Benthic-pelagic coupling via recruitment is potentially important in understanding the pelagic food web in ML 111 and warrants further investigation in this and other aquatic environments.
1. The in situ abundance, biomass and mean cell volume of Actinophrys sol (Sarcodina: Heliozoa), the top predator in an extremely acidic German mining lake (Lake 111; pH 2.65), were determined over three consecutive years (spring to autumn, 2001-03). 2. Actinophrys sol exhibited pronounced temporal and vertical patterns in abundance, biomass and mean cell volume. Increasing from very low spring densities, maxima in abundance and biomass were observed in late June/early July and September. The highest mean abundance recorded during the study was 7 x 10(3) Heliozoa L-1. Heliozoan abundance and biomass were higher in the epilimnion than in the hypolimnion. Actinophrys sol cells from this acidic lake were smaller than individuals of the same species found in other aquatic systems. 3. We determined the growth rate of A. sol using all potential prey items available in, and isolated and cultured from, Lake 111. Prey items included: single-celled and filamentous bacteria of unknown taxonomic affinity, the mixotrophic flagellates Chlamydomonas acidophila and Ochromonas sp., the ciliate Oxytricha sp. and the rotifers Elosa worallii and Cephalodella hoodi. Actinophrys sol fed over a wide-size spectrum from bacteria to metazoans. Positive growth was not supported by all naturally available prey. Actinophrys sol neither increased in cell number (k) nor biomass (k(b)) when starved, with low concentrations of single-celled bacteria or with the alga Ochromonas sp. Positive growth was achieved with single- celled bacteria (k = 0.22 +/- 0.02 d(-1); k(b) = -0.06 +/- 0.02 d(-1)) and filamentous bacteria (k = 0.52 +/- < 0.01 d(- 1); k(b) = 0.66 d(-1)) at concentrations greater than observed in situ, and the alga C. acidophila (up to k = 0.43 +/- 0.03 d(-1); k(b) = 0.44 +/- 0.04 d(-1)), the ciliate Oxytricha sp. (k = 0.34 +/- 0.01 d(-1)) and in mixed cultures containing rotifers and C. acidophila (k = 0.23 +/- 0.02-0.32 +/- 0.02 d(-1); maximum k(b) = 0.42 +/- 0.05 d(-1)). The individual- and biomass-based growth of A. sol was highest when filamentous bacteria were provided. 4. Existing quantitative carbon flux models for the Lake 111 food web can be updated in light of our results. Actinophrys sol are omnivorous predators supported by a mixed diet of filamentous bacteria and C. acidophila in the epilimnion. Heliozoa are important components in the planktonic food webs of 'extreme' environments
Predator-prey cycles rank among the most fundamental concepts in ecology, are predicted by the simplest ecological models and enable, theoretically, the indefinite persistence of predator and prey(1-4). However, it remains an open question for how long cyclic dynamics can be self-sustained in real communities. Field observations have been restricted to a few cycle periods(5-8) and experimental studies indicate that oscillations may be short-lived without external stabilizing factors(9-19). Here we performed microcosm experiments with a planktonic predator-prey system and repeatedly observed oscillatory time series of unprecedented length that persisted for up to around 50 cycles or approximately 300 predator generations. The dominant type of dynamics was characterized by regular, coherent oscillations with a nearly constant predator-prey phase difference. Despite constant experimental conditions, we also observed shorter episodes of irregular, non-coherent oscillations without any significant phase relationship. However, the predator-prey system showed a strong tendency to return to the dominant dynamical regime with a defined phase relationship. A mathematical model suggests that stochasticity is probably responsible for the reversible shift from coherent to non-coherent oscillations, a notion that was supported by experiments with external forcing by pulsed nutrient supply. Our findings empirically demonstrate the potential for infinite persistence of predator and prey populations in a cyclic dynamic regime that shows resilience in the presence of stochastic events.
Despite the increasing number of species invasions, the factors driving invasiveness are still under debate. This is particularly the case for “invisible” invasions by aquatic microbial species. Since in many cases only a few individuals or propagules enter a new habitat, their genetic variation is low and might limit their invasion success, known as the genetic bottleneck. Thus, a key question is, how genetic identity and diversity of invading species influences their invasion success and, subsequently, affect the resident community. We conducted invader-addition experiments using genetically different strains of the globally invasive, aquatic cyanobacterium Raphidiopsis raciborskii (formerly: Cylindrospermopsis raciborskii) to determine the role of invader identity and genetic diversity (strain richness) at four levels of herbivory. We tested the invasion success of solitary single strain invasions against the invader genetic diversity, which was experimentally increased up to ten strains (multi-strain populations). By using amplicon sequencing we determined the strain-specific invasion success in the multi-strain treatments and compared those with the success of these strains in the single-strain treatments. Furthermore, we tested for the invasion success under different herbivore pressures. We showed that high grazing pressure by a generalist herbivore prevented invasion, whereas a specialist herbivore enabled coexistence of consumer and invader. We found a weak effect of diversity on invasion success only under highly competitive conditions. When invasions were successful, the magnitude of this success was strain-specific and consistent among invasions performed with single-strain or multi-strain populations. A strain-specific effect was also observed on the resident phytoplankton community composition, highlighting the strong role of invader genetic identity. Our results point to a strong effect of the genetic identity on the invasion success under low predation pressure. The genetic diversity of the invader population, however, had little effect on invasion success in our study, in contrast to most previous findings. Instead, it is the interaction between the consumer abundance and type together with the strain identity of the invader that defined invasion success. This study underlines the importance of strain choice in invasion research and in ecological studies in general.
Despite the increasing number of species invasions, the factors driving invasiveness are still under debate. This is particularly the case for “invisible” invasions by aquatic microbial species. Since in many cases only a few individuals or propagules enter a new habitat, their genetic variation is low and might limit their invasion success, known as the genetic bottleneck. Thus, a key question is, how genetic identity and diversity of invading species influences their invasion success and, subsequently, affect the resident community. We conducted invader-addition experiments using genetically different strains of the globally invasive, aquatic cyanobacterium Raphidiopsis raciborskii (formerly: Cylindrospermopsis raciborskii) to determine the role of invader identity and genetic diversity (strain richness) at four levels of herbivory. We tested the invasion success of solitary single strain invasions against the invader genetic diversity, which was experimentally increased up to ten strains (multi-strain populations). By using amplicon sequencing we determined the strain-specific invasion success in the multi-strain treatments and compared those with the success of these strains in the single-strain treatments. Furthermore, we tested for the invasion success under different herbivore pressures. We showed that high grazing pressure by a generalist herbivore prevented invasion, whereas a specialist herbivore enabled coexistence of consumer and invader. We found a weak effect of diversity on invasion success only under highly competitive conditions. When invasions were successful, the magnitude of this success was strain-specific and consistent among invasions performed with single-strain or multi-strain populations. A strain-specific effect was also observed on the resident phytoplankton community composition, highlighting the strong role of invader genetic identity. Our results point to a strong effect of the genetic identity on the invasion success under low predation pressure. The genetic diversity of the invader population, however, had little effect on invasion success in our study, in contrast to most previous findings. Instead, it is the interaction between the consumer abundance and type together with the strain identity of the invader that defined invasion success. This study underlines the importance of strain choice in invasion research and in ecological studies in general.
1. During the last couple of decades, invasive species have become a worldwide problem in many freshwater systems. Besides higher plants and animals, microbes, in particular the potentially toxic cyanobacterium Cylindrospermopsis raciborskii, has attracted increasing attention, due to its spread towards temperate zones of the northern and southern hemisphere. A number of advantageous functional traits and a high intraspecific plasticity have been suggested to explain its invasion success. 2. The aim of this study was to examine intraspecific functional trait variability in 12 different isolates of C.raciborskii originating from different lakes in an invaded region in Northeast Germany. We measured growth rate, C:N:P ratios, chlorophyll-a content and the abundance of heterocysts under nutrient-replete and phosphorus-limited conditions. Moreover, the isolate-specific morphology and grazing losses by an herbivorous rotifer, as a top-down force, were studied. 3. DNA fingerprinting revealed that all isolates were genetically different. C.raciborskii exhibited a large variability in all measured traits among isolates. The C:P, N:P and Chl-a:C ratios differed by a factor of two or more. The trait variability among isolates was higher under nutrient-replete conditions, except for the C:P ratio, which varied most during phosphorus limitation. The susceptibility to grazing, calculated as maximum ingestion rates of the rotifer Brachionus calyciflorus on C.raciborskii, varied most among isolates, but was not related to any of the measured physiological or morphological traits, i.e. no trade-off was found. 4. Ecological and genetic clustering did not match, indicating that the genetic relationship based on DNA fingerprinting did not cover ecological differences. 5. Our results show a high trait variability within locally occurring and partly co-occurring C.raciborskii isolates. No overall trade-offs between the measured functional traits were found. This demonstrates the ecological relevance of linking multiple traits, e.g. competitive and consumptive. Furthermore, this study emphasises the importance of analysing more than one strain of a species, as different strains show different trait values potentially relevant for their invasibility and the field of general trait-based ecology.
We report the influence of different nutritional modes-autotrophy, mixotrophy, and heterotrophy-on the fatty acid and sterol composition of the freshwater flagellate Ochromonas sp. and discuss the ecological significance of our results with respect to the resource competition theory (rct). Polyunsaturated fatty acids (PUFAs) are the most efficient biochemical variable distinguishing between nutritional modes of Ochromonas sp. Decreasing concentrations of PUFAs were observed in the order autotrophs, mixotrophs, heterotrophs. In mixotrophs and heterotrophs, concentrations of saturated fatty acids were higher than those of monounsaturated fatty acids and PUFAs as a result of bacterivory. Stigmasterol was the main sterol in Ochromonas sp., regardless of nutritional mode. Mixotrophs showed higher growth rates than heterotrophs, which could not be explained by rct. Heterotrophs, in turn, exhibited higher growth rates than autotrophs, which were cultured under the same light conditions as mixotrophs. Mixotrophs can synthesize PUFAs, which are important for many physiological functions such as membrane permeability and growth. Thus, mixotrophy facilitated efficient growth as well as the ability to synthesize complex and essential biomolecules. These strong synergetic effects are due to the combination of biochemical benefits of heterotrophic and autotrophic metabolic pathways and cannot be predicted by rct.
Negative phototactic response to UVR in three cosmopolitan rotifers: a video analysis approach
(2019)
In recent years, increasing concerns have been raised about the environmental risk of microplastics in freshwater ecosystems. Small microplastics enter the water either directly or accumulate through disintegration of larger plastic particles. These particles might then be ingested by filter-feeding zooplankton, such as rotifers. Particles released into the water may also interact with the biota through the formation of aggregates, which might alter the uptake by zooplankton. In this study, we tested for size-specific aggregation of polystyrene microspheres and their ingestion by a common freshwater rotifer Brachionus calyciflorus. The ingestion of three sizes of polystyrene microspheres (MS) 1-, 3-, and 6-mu m was investigated. Each MS size was tested in combination with three different treatments: MS as the sole food intake, MS in association with food algae and MS aggregated with biogenic matter. After 72 h incubation in pre-filtered natural river water, the majority of the 1-mu m spheres occurred as aggregates. The larger the particles, the higher the relative number of single particles and the larger the aggregates. All particles were ingested by the rotifer following a Type-II functional response. The presence of algae did not influence the ingestion of the MS for all three sizes. The biogenic aggregation of microspheres led to a significant size-dependent alteration in their ingestion. Rotifers ingested more microspheres (MS) when exposed to aggregated 1- and 3-mu m MS as compared to single spheres, whereas fewer aggregated 6-mu m spheres were ingested. This indicates that the small particles when aggregated were in an effective size range for Brachionus, while the aggregated larger spheres became too large to be efficiently ingested. These observations provide the first evidence of a size- and aggregation-dependent feeding interaction between microplastics and rotifers. Microplastics when aggregated with biogenic particles in a natural environment can rapidly change their size-dependent availability. The aggregation properties of microplastics should be taken into account when performing experiments mimicking the natural environment.
Plastic pollution is an increasing environmental problem, but a comprehensive understanding of its effect in the environment is still missing. The wide variety of size, shape, and polymer composition of plastics impedes an adequate risk assessment. We investigated the effect of differently sized polystyrene beads (1-, 3-, 6-µm; PS) and polyamide fragments (5–25 µm, PA) and non-plastics items such as silica beads (3-µm, SiO2) on the population growth, reproduction (egg ratio), and survival of two common aquatic micro invertebrates: the rotifer species Brachionus calyciflorus and Brachionus fernandoi. The MPs were combined with food quantity, limiting and saturating food concentration, and with food of different quality. We found variable fitness responses with a significant effect of 3-µm PS on the population growth rate in both rotifer species with respect to food quantity. An interaction between the food quality and the MPs treatments was found in the reproduction of B. calyciflorus. PA and SiO2 beads had no effect on fitness response. This study provides further evidence of the indirect effect of MPs in planktonic rotifers and the importance of testing different environmental conditions that could influence the effect of MPs.
Plastic pollution is an increasing environmental problem, but a comprehensive understanding of its effect in the environment is still missing. The wide variety of size, shape, and polymer composition of plastics impedes an adequate risk assessment. We investigated the effect of differently sized polystyrene beads (1-, 3-, 6-µm; PS) and polyamide fragments (5–25 µm, PA) and non-plastics items such as silica beads (3-µm, SiO2) on the population growth, reproduction (egg ratio), and survival of two common aquatic micro invertebrates: the rotifer species Brachionus calyciflorus and Brachionus fernandoi. The MPs were combined with food quantity, limiting and saturating food concentration, and with food of different quality. We found variable fitness responses with a significant effect of 3-µm PS on the population growth rate in both rotifer species with respect to food quantity. An interaction between the food quality and the MPs treatments was found in the reproduction of B. calyciflorus. PA and SiO2 beads had no effect on fitness response. This study provides further evidence of the indirect effect of MPs in planktonic rotifers and the importance of testing different environmental conditions that could influence the effect of MPs.
The uptake of resources from the environment is a vital process for all organisms. Many experimental studies have revealed that the rate at which this process occurs depends critically on the resource concentration, a relationship called "functional response." However, whether the concentration of the consumer normally affects the functional response has been the subject of a long-standing, predominantly theoretical, debate in ecology. Here we present an experimental test between the alternative hypotheses that food uptake depends either only on the resource concentration or on both the resource and the consumer concentrations. In short-term laboratory experiments, we measured the uptake of radioactively labeled, unicellular green algae (Monoraphidium minutum, resource) by the rotifer Brachionus calyciflorus (a consumer) for varying combinations of resource and consumer concentrations. We found that the food uptake by Brachionus depended on the algal concentration with the relationship best described by a Holling type 3 functional response. We detected significant consumer effects on the functional response only at an extraordinarily high Brachionus density (similar to 125 rotifers/mL), which by far exceeds concentrations normally encountered in the field. We conclude that con sumer-dependent food uptake by planktonic rotifers is a phenomenon that can occur under extreme conditions, but probably plays a minor role in natural environments
A characteristic vertical sequence of phytoplankton populations was observed below the metalimnetic oxycline of a stratified, mesotrophic lake. Ceratium spp., Closterium acutum and Aphanizomenon flos- aquae were present in the epilimnion but had distinct population maxima in the microaerobic chemocline. Below these populations, Cryptomonas phaseolus, Planktothrix clathrata, Pseudanabaena catenata and Limnothrix sp. followed each other in the transition zone between the chemocline and the sulphide-containing hypolimnion. The dominating populations of P. clathrata and P. catenata caused a deep chlorophyll maximum. Phytoplankton structure was determined by the vertical gradients of sulphide and light. Compared with the epilimnion, nutrient availability was not fundamentally better below the oxycline but the algae might have benefited from reduced grazing pressure in their habitat.
Under natural conditions, aboveground herbivory and plant-soil feedbacks (PSFs) are omnipresent interactions strongly affecting individual plant performance. While recent research revealed that aboveground insect herbivory generally impacts the outcome of PSFs, no study tested to what extent the intensity of herbivory affects the outcome. This, however, is essential to estimate the contribution of PSFs to plant performance under natural conditions in the field. Here, we tested PSF effects both with and without exposure to aboveground herbivory for four common grass species in nine grasslands that formed a gradient of aboveground invertebrate herbivory. Without aboveground herbivores, PSFs for each of the four grass species were similar in each of the nine grasslands-both in direction and in magnitude. In the presence of herbivores, however, the PSFs differed from those measured under herbivory exclusion, and depended on the intensity of herbivory. At low levels of herbivory, PSFs were similar in the presence and absence of herbivores, but differed at high herbivory levels. While PSFs without herbivores remained similar along the gradient of herbivory intensity, increasing herbivory intensity mostly resulted in neutral PSFs in the presence of herbivores. This suggests that the relative importance of PSFs for plant-species performance in grassland communities decreases with increasing intensity of herbivory. Hence, PSFs might be more important for plant performance in ecosystems with low herbivore pressure than in ecosystems with large impacts of insect herbivores.
We describe a new species of Cephalodella, C. acidophila n. sp., from the plankton of two extremely acidic mining lakes (pH <3) in Austria and Germany. The species is morphologically closely related to Cephalodella delicata Wulfert. It shares with the latter an almost identical trophi morphology and anatomical organization, but differs clearly by form and length of its toes, larger body size, and ecology. Laboratory experiments revealed that the species is acidophilic, i.e. it thrives at low pH (<4) and does not survive at circumneutral conditions. The species occurs in man-made habitats at low to moderate abundance (usually 5-22 individuals l(-1)) and in stock cultures thrives on the green alga Chlamydomonas acidophila. The easily cultured species has previously been used in various experimental studies, but has only now been recognized as an undescribed species.
Vertical differences in food web structure were examined in an extremely acidic, iron-rich mining lake in Germany (Lake 111; pH 2.6, total Fe 150mg L-1) during the period of stratification. We tested whether or not the seasonal variation of the plankton composition is less pronounced than the differences observed over depth. The lake was strongly stratified in summer, and concentrations of dissolved organic carbon and inorganic carbon were consistently low in the epilimnion but high in the hypolimnion. Oxygen concentrations declined in the hypolimnion but were always above 2mg L-1. Light attenuation did not change over depth and time and was governed by dissolved ferric iron. The plankton consisted mainly of single-celled and filamentous bacteria, the two mixotrophic flagellates Chlamydomonas sp. and Ochromonas sp., the two rotifer species Elosa worallii and Cephalodella hoodi, and Heliozoa as top predators. We observed very few ciliates and rhizopods, and no heterotrophic flagellates, crustaceans or fish. Ochromonas sp., bacterial filaments, Elosa and Heliozoa dominated in the epilimnion whereas Chlamydomonas sp., single-celled bacteria and Cephalodella dominated in the hypolimnion. Single-celled bacteria were controlled by Ochromonas sp. whereas the lack of large consumers favoured a high proportion of bacterial filaments. The primarily phototrophic Chlamydomas sp. was limited by light and CO2 and may have been reduced due to grazing by Ochromonas sp. in the epilimnion. The distribution of the primarily phagotrophic Ochromonas sp. and of the animals seemed to be controlled by prey availability. Differences in the plankton composition were much higher between the epilimnion and hypolimnion than within a particular stratum over time. The food web in Lake 111 was extremely species-poor enabling no functional redundancy. This was attributed to the direct exclusion of species by the harsh environmental conditions and presumably enforced by competitive exclusion. The latter was promoted by the low diversity at the first trophic level which, in turn, was attributed to relatively stable growth conditions and the independence of resource availability (inorganic carbon and light) from algal density. Ecological theory suggests that low functional redundancy promotes low stability in ecosystem processes which was not supported by our data.
The Brachionus calyciflorus species complex was recently subdivided into four species, but genetic resources to resolve phylogenetic relationships within this complex are still lacking. We provide two complete mitochondrial (mt) genomes from B. calyciflorus sensu stricto (Germany, USA) and the mt coding sequences (cds) from a German B. fernandoi. Phylogenetic analysis placed our B. calyciflorus sensu stricto strains close to the published genomes of B. calyciflorus, forming the putative sister species to B. fernandoi. Global representatives of B. calyciflorus sensu stricto (i.e. Europe, USA, and China) are genetically closer related to each other than to B. fernandoi (average pairwise nucleotide diversity 0.079 intraspecific vs. 0.254 interspecific).