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Cytostatic drugs are one of the most important therapeutic options for cancer, a disease that is expected to affect 29 million individuals by 2040. After being excreted, cytostatics reach wastewater treatment plants (WWTPs), which are unable to efficiently remove them, and consequently, they will be released into the aquatic environment. Due to the highly toxic properties of cytostatics, it is particularly relevant to evaluate their potential ecological risk. Yet, cytostatics toxicity data is still not available for various species. In this work, the ecotoxicity of two widely consumed cytostatics, cyclophosphamide (CYP-as a model cytostatic) and mycophenolic acid (MPA-as a priority cytostatic), was evaluated on three freshwater species-Raphidocelis subcapitata, Brachionus calyciflorus, and Danio rerio, and the risk quotient (RQ) was assessed. Both drugs significantly affected the yield and growth inhibition of the microalgae, while for rotifers, the least sensitive species, only significant effects were registered for CYP. These drugs also caused significant effects on the mortality and morphological abnormalities on zebrafish. The estimation of the RQ discloses that CYP seems to pose a low risk to aquatic biota while MPA poses a very high risk. Altogether, these results emphasize the need for more complete environmental risk assessments, to properly prioritize and rank cytostatics according to their potentially toxic effects on the environment and aquatic biota.

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The growth rate hypothesis (GRH) posits that the relative body phosphorus content of an organism is positively related to somatic growth rate, as protein synthesis, which is necessary for growth, requires P-rich rRNA. This hypothesis has strong support at the interspecific level. Here, we explore the use of the GRH to predict microevolutionary responses in consumer body stoichiometry. For this, we subjected populations of the rotifer Brachionus calyciflorus to selection for fast population growth rate (PGR) in P-rich (HPF) and P-poor (LPF) food environments. With common garden transplant experiments, we demonstrate that in HP populations evolution toward increased PGR was concomitant with an increase in relative phosphorus content. In contrast, LP populations evolved higher PGR without an increase in relative phosphorus content. We conclude that the GRH has the potential to predict microevolutionary change, but that its application is contingent on the environmental context. Our results highlight the potential of cryptic evolution in determining the performance response of populations to elemental limitation of their food resources.

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Zooplankton bodies are organic-rich micro-environments that support fast bacterial growth. Therefore, the abundance of zooplankton-associated bacteria is much higher than that of free-living bacteria, which has profound effects on the nutrient cycling of freshwater ecosystems. However, a detailed analysis of associated bacteria is still less known, especially the relationship between those bacteria and bacterioplankton. In this study, we analyzed the relationships between Brachionus calyciflorus-associated bacterial and bacterioplankton communities in freshwater using high-throughput sequencing. The results indicated that there were significant differences between the two bacterial communities, with only 29.47% sharing OTUs. The alpha diversity of the bacterioplankton community was significantly higher than that of B. calyciflorus-associated bacteria. PCoA analysis showed that the bacterioplankton community gathered deeply, while the B. calyciflorus-associated bacterial community was far away from the whole bacterioplankton community, and the distribution was relatively discrete. CCA analysis suggested that many environmental factors (T, DO, pH, TP, PO43-, NH4+, and NO3-) regulated the community composition of B. calyciflorus-associated bacteria, but the explanatory degree of variability was only 37.80%. High-throughput sequencing revealed that Raoultella and Delftia in Proteobacteria were the dominant genus in the B. calyciflorus-associated bacterial community, and closely related to the biodegradation function. Moreover, several abundant bacterial members participating in carbon and nitrogen cycles were found in the associated bacterial community by network analysis. Predictive results from FAPROTAX showed that the predominant biogeochemical cycle functions of the B. calyciflorus-associated bacterial community were plastic degradation, chemoheterotrophy, and aerobic chemoheterotrophy. Overall, our study expands the current understanding of zooplankton-bacteria interaction and promotes the combination of two different research fields.

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Predation is a strong species interaction causing severe harm or death to prey. Thus, prey species have evolved various defence strategies to minimize predation risk, which may be immediate (e.g., a change in behaviour) or transgenerational (morphological defence structures). We studied the behaviour of two strains of a rotiferan prey (Brachionus calyciflorus) that differ in their ability to develop morphological defences in response to their predator Asplanchna brightwellii. Using video analysis, we tested: (a) if two strains differ in their response to predator presence and predator cues when both are undefended; (b) whether defended individuals respond to live predators or their cues; and (c) if the morphological defence (large spines) per se has an effect on the swimming behaviour. We found a clear increase in swimming speed for both undefended strains in predator presence. However, the defended specimens responded neither to the predator presence nor to their cues, showing that they behave indifferently to their predator when they are defended. We did not detect an effect of the spines on the swimming behaviour. Our study demonstrates a complex plastic behaviour of the prey, not only in the presence of their predator, but also with respect to their defence status.

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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).

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Classic population regulation theories usually concern the influence of immediate factors on current populations, but studies investigating the effect of parental environment factors on their offspring populations are scarce. The maternal environments can affect offspring life-history traits across generations, which may affect population dynamics and be a mechanism of population regulation. In cyclical parthenogens, sexual reproduction is typically linked with dormancy, thereby providing a negative feedback to population growth. In this study, we manipulated population sex ratios in the mother's environment to investigate whether this factor affected future population dynamics by regulating offspring sexual reproduction in the rotifer Brachionus calyciflorus. Compared with females in male-biased environments, those in female-biased environments produced fewer mictic (sexual) offspring, and their amictic (asexual) offspring also produced a lower proportion of mictic females at a gradient of population densities. Moreover, populations that were manipulated under male-biased conditions showed significantly smaller population sizes than those under female-biased conditions. Our results indicated that in cyclical parthenogens, mothers could adjust the sexual reproduction of their offspring in response to the current population sex ratio, thus providing fine-scale regulation of population dynamics in addition to population density.

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Global change encompasses many co-occurring anthropogenic stressors. Understanding the interactions between these multiple stressors, whether they be additive, antagonistic or synergistic, is critical for ecosystem managers when prioritizing which stressors to mitigate in the face of global change. While such interactions between stressors appear prevalent, it remains unclear if and how these interactions change over time, as the majority of multiple-stressor studies rarely span multiple generations of study organisms. Although meta-analyses have reported some intriguing temporal trends in stressor interactions, for example that synergism may take time to emerge, the mechanistic basis for such observations is unknown. In this study, by analysing data from an evolution experiment with the rotifer Brachionus calyciflorus (~35 generations and 31,320 observations), we show that adaptation to multiple stressors shifts stressor interactions towards synergism. We show that trade-offs, where populations cannot optimally perform multiple tasks (i.e. adapting to multiple stressors), generate this bias towards synergism. We also show that removal of stressors from evolved populations does not necessarily increase fitness and that there is variation in the evolutionary trajectories of populations that experienced the same stressor regimes. Our results highlight outstanding questions at the interface between evolution and global change biology, and illustrate the importance of considering rapid adaptation when managing or restoring ecosystems subjected to multiple stressors under global change.

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Zooplankton provides bacteria with a complex microhabitat richen in organic and inorganic nutrients, and the bacteria community also changes the physiochemical conditions for zooplankton, where the symbiotic relationship between them plays an important role in the nutrient cycle. However, there are few studies on the effect of associated bacteria on the population dynamics of rotifers. In order to make clear their relationships, we reconstructed the associated bacterial community in Brachionus calyciflorus culture, and examined the life history and population growth parameters, and analyzed the diversity and community composition of the associated bacteria at different growth stages of B. calyciflorus. The results showed that the addition of bacteria from natural water can promote the population growth and asexual reproduction of B. calyciflorus, but has no significant effect on sexual reproduction, exhibited by the improvement of its life expectancy at hatching, net reproduction rates and intrinsic growth rate, no significant effects on the generation time and mixis ratio of offspring. It was found that the B. calyciflorus-associated bacterial community was mainly composed of Proteobacteria, Bacteroidota, Actinobacteriota, Cyanobacteria and Firmicutes. Through correlation network analysis, the members of Burkholderiales, Pseudomonadales, Micrococcales, Caulobacterales and Bifidobacteriales were the keystone taxa of B. calyciflorus-associated bacteria. In addition, the relative abundance of some specific bacteria strains increased as the population density of B. calyciflorus increased, such as Hydrogenophaga, Acidovorax, Flavobacterium, Rheinheimera, Novosphingobium and Limnobacter, and their relative abundance increased obviously during the slow and exponential phases of population growth. Meanwhile, the relative abundance of adverse taxa (such as Elizabethkingia and Rickettsiales) decreased significantly with the increase in rotifer population density. In conclusion, the closely associated bacteria are not sufficient for the best growth of B. calyciflorus, and external bacterioplankton is necessary. Furthermore, the function of keystone and rare taxa is necessary for further exploration. The investigation of the symbiotic relationship between zooplankton-associated bacterial and bacterioplankton communities will contribute to monitoring their roles in freshwater ecosystems, and regulate the population dynamics of the micro-food web.

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Various types of pollutants derived from rapid industrialization and urbanization have largely threaten biodiversity and functioning of freshwater ecosystems globally. Morphological plasticity, especially body size-associated traits, is considered a functional response to water pollution in species, as such changes are often directly related to functioning of freshwater ecosystems through dynamics of food webs. However, detailed dynamics of pollution impacts on morphological plasticity remain largely unknown, particularly in the wild. Here, we used the model planktonic rotifer Brachionus calyciflorus to assess morphological response to chemical pollution in a river reach disturbed by sewage discharges. Multiple analyses showed dynamic morphological response to water pollution in wild B. calyciflorus populations. The distance between anterior lateral spines, lorica length, and egg short diameter were the most sensitive morphological indicators to water pollution, while spine length was stable in varied pollution conditions. Interestingly, body size and egg size were increased with accentuated water pollution, suggesting that wild populations maintain fitness by increasing feeding efficiency and reducing vulnerability to predation and ensure survival by producing large newborns in polluted environments. Total ammonia nitrogen was the leading nitrogen pollutant affecting body size, while total phosphorus and elements of Mn and As were the key factors relating to egg size. The results obtained here provide new sights into biological consequences of environmental pollution in the wild, thus advancing our understanding of pollution impacts on structure and functioning of freshwater ecosystems.

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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.

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Much of the recent literature concerning the threat posed by anthropogenic microscopic pollution has focussed on marine organisms although freshwater environments face the same degree of pollution and therefore risk. Although several studies have documented the ingestion of nanoparticles (NPs) in species of the pelagic freshwater rotifer genus Brachionus, unambiguous evidence for its cellular uptake in this group remains lacking. We therefore used transmission electron microscopy (TEM) of ultrathin sections through the digestive tract of individuals of Brachionus calyciflorus exposed in vitro to citrate stabilized gold nanoparticles (AuCit NPs) in their culture medium to provide the first concrete evidence for the cellular uptake of NPs in rotifers, a group of organisms that comprise an important part of the zooplankton community. Using this method, AuCit NPs with average diameters of 8.5 ± 1.4 nm and 12.5 ± 1.5 nm could be localized clearly within large vacuoles within the stomach cells. Moreover, the occasional presence of pits containing AuCit NPs in the outer membranes of these cells hints that the particles are taken up by some form of endocytosis. In all cases, the ingestion of AuCit NPs showed lethal effects after only one day with virtually no individuals surviving more than two days of exposure. Combined with the TEM evidence above, we hypothesize that death might derive from some form of lysosomal overload. In total, our results document the potential threat that microscopic pollution also poses for freshwater organisms. Through this, we hope that additional emphasis in this context will be directed toward freshwater environments and the potential for such pollution both to enter as well as to move up the food chain via trophic transfer events.

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The baking industries and disinfection of tap water released a considerable amount of bromate into surface water, which has been reported as a carcinogenic compound to mammals. Rotifers play an important role in freshwater ecosystems and are model organisms to assess environmental contamination. In the present study, the effects of different concentrations (0.001, 0.01, 0.1, 1, 10, 100 and 200 mg/L) of bromate on the life-table and population growth parameters were investigated in the rotifer Brachionus calyciflorus. The results showed that the 24-h LC50 of bromate to B. calyciflorus was 365.29 mg/L (95%Cl: 290.37-480.24). Treatments with 0.01, 10 and 200 mg/L bromate shorten the reproductive period. High levels of bromate (100 and 200 mg/L) significantly decreased net reproductive rate, intrinsic rate of population increase, life span, mictic rate of B. calyciflorus. To investigate the underlying mechanisms, swimming speed and antioxidative biomarkers were compared between bromate treatments and the control. The results showed that glutathione (GSH) and malondialdehyde (MDA) contents, total superoxide dismutase (T-SOD) and peroxidase (POD) activities decreased significantly in response to bromate exposure and the reasons required further investigation. Treatments with 0.001-200 mg/L bromate all significantly reduced swimming linear speed to rotifer larvae and treatments with 100-200 mg/L bromate significantly suppressed the swimming linear speed of adult rotifer. These changes would reduce filtration of algal food and could explain the decreased survival and reproduction. Overall, bromate may not show acute toxicity to rotifers, but still have potential adverse effects on rotifer behavior, which may then influence the community structure in aquatic ecosystems.

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The effects of potassium bromate (KBrO3), sodium bromate (NaBrO3), and potassium bromide (KBr) on the sexual reproduction of the rotifer Brachionus calyciflorus were studied by 2-d population growth, 4-d sexual reproduction, and 7-d resting egg production tests. The results showed that low concentrations of bromate promote 2-d and 4-d rotifer population growth, while high concentrations limit it. Bromate stress significantly affected parameters of rotifer sexual reproduction, including the ratio of mictic to amictic females, the mictic rate of rotifers, and the fertilization rate of mictic females. KBrO3 at 0.001, 0.01, 1, and 10 mg/L, NaBrO3 at 1 and 10 mg/L, and KBr at 100 and 200 mg/L significantly increased resting egg production, while KBrO3 at 100 and 200 mg/L, and NaBrO3 at 200 mg/L significantly decreased it. Resting egg production appears to provide a sensitive endpoint in evaluating the effect of bromate on rotifer sexual reproduction.

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Titanium dioxide nanoparticles (nTiO2 ) have raised environmental concerns and display potential hazards to aquatic organisms and even humans. However, only a few reports tested the toxicity of nTiO2 to rotifers. In the present study, acute and chronic toxicity of nTiO2 to the freshwater rotifer Brachionus calyciflorus was investigated at different temperatures. At 25°C, the 24 and 48-hr LC50 was 117.14 and 60.11 mg/L, respectively. At 15°C, 20°C, 25°C, and 30°C, exposure to nTiO2 significantly decreased life expectancy at birth, net reproductive rate, generation time, average lifespan, and/or intrinsic rate of population increase of B. calyciflorus (p < .05). High temperature enhanced the toxicity of nTiO2 to rotifers. The swimming linear speed of rotifers significantly increased (p < .05) in treatments with 200 µg/L nTiO2 , compared with the control. In addition, treatments with 8 µg/L to 5 mg/L nTiO2 significantly increased superoxide dismutase activity (p < .05). Glutathione content and catalase activity increased significantly after exposure to 8 µg/L nTiO2  but decreased significantly in treatments with nTiO2 concentrations ranging from 40 µg/L to 5 mg/L (p < .05). There were no significant changes in malondialdehyde contents among nTiO2 treatments and control. Overall, the present study indicated that nTiO2 revealed high toxicity to rotifers, displaying high environmental risks to aquatic ecosystems.

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Anthropogenic activities have reshaped the relative supply rates of essential elements to organisms. Recent studies suggested that consumer performance is strongly reduced by food that is either very high or very low in relative phosphorus content. However, the generality of such 'stoichiometric knife-edge' and its underlying mechanisms are poorly understood. We studied the response of a planktonic rotifer to a 10-fold food carbon : phosphorus (C : P) gradient and confirmed the existence of the stoichiometric knife-edge. Interestingly, we observed a complete homeostatic breakdown associated with strong growth reductions at high food C : P. In contrast, at low food C : P, animals maintained homeostasis despite pronounced performance reductions. Our results suggest that the mechanisms underlying adverse effects of stoichiometric imbalance are determined by both the identity of elements that are limiting and those that are present in excess. Negative effects of excess P reveal an additional way of how eutrophication may affect consumers.

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Many organisms display oscillations in population size. Theory predicts that these fluctuations can be generated by predator-prey interactions, and empirical studies using life model systems, such as a rotifer-algae community consisting of Brachionus calyciflorus as predator and Chlorella vulgaris as prey, have been successfully used for studying such dynamics. B. calyciflorus is a cyclical parthenogen (CP) and clones often differ in their sexual propensity, that is, the degree to which they engage into sexual or asexual (clonal) reproduction. Since sexual propensities can affect growth rates and population sizes, we hypothesized that this might also affect population oscillations. Here, we studied the dynamical behaviour of B. calyciflorus clones representing either CPs (regularly inducing sex) or obligate parthenogens (OPs). We found that the amplitudes of population cycles to be increased in OPs at low nutrient levels. Several other population dynamic parameters seemed unaffected. This suggests that reproductive mode might be an important additional variable to be considered in future studies of population oscillations.

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The two complete mitochondrial genomes were sequenced from the Netherlands strain of the freshwater monogonont rotifer Brachionus calyciflorus. The mitochondrial genome sequences were 27,698 bp and 9,906 bp in size, respectively. The gene order and contents of the two B. calyciflorus strains were mostly identical to one another, except for the additional identification and translocation of several tRNAs in mitochondrial DNA I and II. Of 13 protein-coding genes (PCGs), three genes (ND1, ND5, and ND3) had incomplete stop codons. Furthermore, the start codon of ND2, CO2, and CO3 and ND4 genes was ATT, GTG, and ATA, respectively, while the start codon of other PCGs was ATG. The base composition of 13 PCGs of B. calyciflorus (the Netherlands strain) mitogenome showed 31.1% for A, 37.6% for T, 16.5% for C, and 14.8% for G, respectively.

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Under global warming scenarios, rising temperatures can constitute heat stress to which species may respond differentially. Within a described species, knowledge on cryptic diversity is of further relevance, as different lineages/cryptic species may respond differentially to environmental change. The Brachionus calyciflorus species complex (Rotifera), which was recently described using integrative taxonomy, is an essential component of aquatic ecosystems. Here, we tested the hypothesis that these (formerly cryptic) species differ in their heat tolerance. We assigned 47 clones with nuclear ITS1 (nuITS1) and mitochondrial COI (mtCOI) markers to evolutionary lineages, now named B. calyciflorus sensu stricto (s.s.) and B. fernandoi We selected 15 representative clones and assessed their heat tolerance as a bi-dimensional phenotypic trait affected by both the intensity and duration of heat stress. We found two distinct groups, with B. calyciflorus s.s. clones having higher heat tolerance than the novel species B. fernandoi This apparent temperature specialization among former cryptic species underscores the necessity of a sound species delimitation and assignment, when organismal responses to environmental changes are investigated.

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Theory predicts that fitness decline via mutation accumulation will depend on population size, but there are only a few direct tests of this key idea. To gain a qualitative understanding of the fitness effect of new mutations, we performed a mutation accumulation experiment with the facultative sexual rotifer Brachionus calyciflorus at six different population sizes under UV-C radiation. Lifetime reproduction assays conducted after ten and sixteen UV-C radiations showed that while small populations lost fitness, fitness losses diminished rapidly with increasing population size. Populations kept as low as 10 individuals were able to maintain fitness close to the nonmutagenized populations throughout the experiment indicating that selection was able to remove the majority of large effect mutations in small populations. Although our results also seem to imply that small populations are effectively immune to mutational decay, we caution against this interpretation. Given sufficient time, populations of moderate to large size can experience declines in fitness from accumulating weakly deleterious mutations as demonstrated by fitness estimates from simulations and, tentatively, from a long-term experiment with populations of moderate size. There is mounting evidence to suggest that mutational distributions contain a heavier tail of large effects. Our results suggest that this is also true when the mutational spectrum is altered by UV radiation.

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Maternal effects play important evolutionary and ecological roles. Amictic female mothers of monogonont rotifer Brachionus calyciflorus can transmit predatory information of Asplanchna brightwellii in their environment to their offspring by changing the offspring's defensive morphology to increase their fitness. However, it remains unclear whether such maternal effects also exist during sexual reproduction of a mictic mother. This study explored the maternal effect in mictic mothers using the B. calyciflorus and A. brightwellii as a prey-predator model. We collected resting eggs from two groups of mictic mothers that previously experienced environments with (P environment) or without (NP environment) Asplanchna kairomones. Stem females from the resting eggs of each maternal group were also hatched and reared in P and NP environments. The population growth rate of offspring who experienced the same environment as their mictic mothers was significantly higher than those that experienced a different environment. When exposed to a gradient of predator kairomone levels, the posterolateral spine of the offspring elongated with increasing kairomone concentration. Offspring from the P mictic mother showed significantly shorter posterolateral spines than those from the NP mictic mother at each predator kairomone level. Offspring originating from NP mictic mothers clearly elongated their posterolateral spines at low concentrations of predator kairomones, while those from P mothers elongated their posterolateral spines only at the highest levels of predator kairomone. Our findings highlight the existence of anticipatory maternal effects during the sexual process via resting eggs of B. calyciflorus in response to predator kairomone.

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