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Sewer overflows are an environmental concern due to their potential to introduce contaminants that can adversely affect downstream aquatic ecosystems. As these overflows can occur during rainfall events, the influence of rainwater ingress from inflow and infiltration on raw untreated wastewater (influent) within the sewer is a critical factor influencing the dilution and toxicity of the contaminants. The Vineyard sewer carrier in the greater city of Sydney, Australia, was selected for an ecotoxicological investigation of a sanitary (separate from stormwater) sewerage system and a wet-weather overflow (WWO). Three influent samples were collected representing dry-weather (DW), intermediate wet-weather (IWW) and wet-weather (WW). In addition, a receiving water sample was also collected downstream in Vineyard Creek (WW-DS) coinciding with a WWO. We employed direct toxicity assessment (DTA) and toxicity identification evaluation (TIE) approaches to gain comprehensive insights into the nature and magnitude of the impact on influent from rainwater ingress into the sewer. Three standard ecotoxicological model species, a microalga, Chlorella vulgaris, the water flea, Ceriodaphnia dubia and the midge larva, Chironomus tepperi were used for both acute and chronic tests. The study revealed variable toxicity responses, with the sample of influent collected in wet-weather displaying lower toxicity compared to the dry-weather sample of influent. Ammonia, and metals, were identified in dry weather as contributors to the observed toxicity, however, this risk was alleviated through rainwater ingress in wet-weather with further dilution within the receiving water. Based on toxicity data, dilutions of influent to minimise effects on C. vulgaris and C. dubia ranged from 1 in 12 in DW to 1 in 2.8 in WW, and further diminished in the receiving water to 1 in 1.8. The successful application of ecotoxicological approaches enabled the assessment of cumulative effects of contaminants in influent, offering valuable insights into the sanitary sewer system under rainwater ingress.

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Irgarol 1051 is an herbicide extensively utilized in antifouling paint due to its ability to inhibit photosynthesis. Irgarol and its photodegradation products are highly persistent in waters and sediments, although they are present in low concentrations. However, our understanding of the harmful effects of Irgarol on non-target organisms remains limited. In this study, we assessed the effects of acute (24 h) and chronic (14 days across three generations) exposure to different concentrations (including the 1/10 NOEC, NOEC, and 1/10 LC50 calculated from the 24-h acute toxicity test) of Irgarol using the water flea Moina macrocopa. Acute exposure to 1/10 LC50 significantly decreased survival, feeding rate, thoracic limb activity, heart rate, and acetylcholinesterase activity. Elevated levels of intracellular reactive oxygen species and malondialdehyde, along with a significant increase in catalase and superoxide dismutase activity, suggested the induction of oxidative stress in response to 1/10 LC50. An initial boost in glutathione level and the enzymatic activities of glutathione peroxidase and glutathione reductase, followed by a plunge, implies some compromise in the antioxidant defense system. Upon chronic exposure to the NOEC value, both generations F1 and F2 displayed a significant decrease in survival rate, body length, number of neonates per brood, and delayed sexual maturation, suggesting maternal transfer of potential damage through generations. Taken together, Irgarol induced acute toxicity through physiological and cholinergic damage, accompanied by the induction of oxidative stress, in the water flea. Even its sub-lethal concentrations can induce detrimental effects across generations when consistently exposed.

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Aqueous calcium (Ca) decline is threatening freshwater ecosystems worldwide. There are great concerns about the possible ecological consequences of Ca limitation combined with biological pressures like predation. Here we investigated the interactions between Ca restriction and fish predation risk on the phenotypic plasticity in the keystone herbivore Daphnia, together with physiological responses underlying the plastic trait changes. Fish predation risk induced D. pulex to mature earlier and produce more but smaller offspring at adequate Ca. Declining Ca inhibited the expression of defensive traits, with the inhibitive degree showing a linear or threshold-limited dynamic. The presence of predation risk mitigated the negative effect of declining Ca on reducing body size but exacerbated the delay in maturity, indicating a life history trade-off for larger body size rather than the current reproduction in multi-stressed Daphnia. Actin 3-mediated cytoskeleton and AMPK ß-mediated energy metabolism were highly correlated with these plastic trait changes. Altered phenotypic plasticity in planktonic animals is expected to trigger many ecological impacts from individual fitness to community structure, thus providing new insights into the mechanisms underlying decreased Ca affecting lake ecosystems.

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Parabens, a group of alkyl esters of p-hydroxybenzoic acid, have been found in aquatic systems in particular, leading to concerns about their potential impact on ecosystems. This study investigated the effects of three commonly used parabens, methylparaben (MeP), ethylparaben (EtP), and propylparaben (PrP), on the brackish water flea Diaphanosoma celebensis. The results showed that PrP had the most adverse impact on survival rates, followed by EtP and MeP, while MeP and EtP induced significant adverse effects on reproductive performance. A transcriptome analysis revealed significant differential gene expression patterns in response to paraben exposure, with MeP associated with the most significant effects. MeP and EtP exposure produced greater disruption in the microbiota of D. celebensis than did PrP compared with control groups, and we identified eight key microbiota, including Ruegeria and Roseovarius. Correlation analysis between transcriptome and microbiome data revealed key interactions between specific microbiota and host gene expression. Certain microbial taxa were associated with specific genes (e.g. cuticle related genes) and toxicological pathways, shedding light on the complex molecular response and in vivo toxicity effects of parabens. These findings contribute to a deeper understanding of the molecular mechanisms underlying paraben toxicity and highlight the importance of considering the ecological impact of chemical contaminants in aquatic ecosystems.

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The toxicity of triclosan (TCS) on the freshwater cladoceran Moina macrocopa was investigated by acute and chronic toxicity assessments followed by genotoxicity and oxidative stress response analyses. The 48-h LC50 of TCS for ≤24-h-old M. macrocopa was determined as 539 µg L-1 . Chronic exposure to TCS at concentrations ranging from 5 to 100 µg L-1 showed a stimulatory effect at low concentrations (≤10 µg L-1 ) and an inhibitory effect at high concentrations (≥50 µg L-1 ) on growth, reproduction, and population-growth-related parameters of M. macrocopa. The genotoxicity test results indicated that TCS concentrations ranging from 50 to 100 µg L-1 can alter individuals' DNA. Analysis of the antioxidant enzymes catalase (CAT) and glutathione s-transferase (GST) demonstrated increased levels of these enzymes at high TCS concentrations. Our results indicated that TCS concentrations found in the natural environment have minimal acute toxicity to M. macrocopa. However, TCS at even low concentrations can significantly affect its growth, reproduction, and population-growth-related characteristics. The observed responses suggest a hormetic dose-response pattern and imply a potential endocrine-disrupting effect of TCS. Our molecular and biochemical findings indicated that high concentrations of TCS have the potential to induce oxidative stress that may lead to DNA alterations in M. macrocopa.

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Rising ocean temperatures are driving unprecedented changes in global marine ecosystems. Meanwhile, there is growing concern about microplastic and nanoplastic (MNP) contamination, which can endanger marine organisms. Increasing ocean warming (OW) and plastic pollution inevitably cause marine organisms to interact with MNPs, but relevant studies remain sparse. Here, we investigated the interplay between ocean warming and MNP in the marine water flea Diaphanosoma celebensis. We found that combined exposure to MNPs and OW induced reproductive failure in the F2 generation. In particular, the combined effects of OW and MNPs on the F2 generation were associated with key genes related to reproduction and stress response. Moreover, populations of predatory bacteria were significantly larger under OW and MNP conditions during F2 generations, suggesting a potential link between altered microbiota and host fitness. These results were supported by a host transcriptome and microbiota interaction analysis. This research sheds light on the complex interplay between environmental stressors, their multigenerational effects on marine organisms, and the function of the microbiome.

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Triclosan, a chlorinated biphenyl ether is widely used in industrial products and cosmetics due to its antibiotic activity. Although relatively levels of triclosan have been detected in aquatic ecosystems, limited information is available regarding the acute and chronic impacts of triclosan on aquatic invertebrates, especially planktonic crustaceans. In this study, we analyzed the acute (24 h) and chronic (14 days exposure across three generations) effects of different concentrations of triclosan [1/10 of the no observed effect concentration (NOEC), the NOEC, and 1/10 of the LC50] calculated from the 24 h acute toxicity value, on the water flea Moina macrocopa. In the acute exposure experiment, the 1/10 LC50 value of triclosan significantly reduced survival, feeding rate, thoracic limb activity, heart activity, and acetylcholinesterase activity. In response to the 1/10 LC50 value, intracellular reactive oxygen species increased along with elevated levels of malondialdehyde and glutathione. Enzymatic activities of catalase, superoxide dismutase, glutathione peroxidase, and glutathione reductase were significantly increased by the 1/10 LC50 value, suggesting active protection of the antioxidant defense system against oxidative stress. Chronic exposure to the 1/10 NOEC and NOEC values revealed multigenerational adverse impacts of triclosan. The second generation was found to be the most sensitive to triclosan, as the NOEC value significantly reduced the survival rate, body length, and the number of neonates per brood, along with a delayed hatching period. Taken together, these results indicate that even sublethal levels of triclosan can have detrimental effects on the water flea population's maintenance through intergenerational toxicity.

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Global warming and nanoplastics (NPs) are critical global issues. Among NPs, one of the most hazardous types of plastics, polystyrene (PS), poses ecotoxicological threats to several freshwater organisms. The degree of toxicity of PS-NPs is strongly influenced by various environmental factors. This study illustrates the combined effects of temperature and PS-NPs on the water flea Daphnia magna. The sensitivity of D. magna to PS-NPs was tested under control (23 °C) and elevated temperatures (28 °C). As a result, increased temperatures influenced the uptake and accumulation of PS-NPs. Co-exposure to both higher temperatures and PS-NPs resulted in a drastic decrease in reproductive performance. The level of oxidative stress was found to have increased in a temperature-dependent manner. Oxidative stress was stimulated by both stressors, leading to increased levels of reactive oxygen species and antioxidant enzyme activity supported by upregulation of antioxidant enzyme-related genes under combined PS-NPs exposure and elevated temperature. In the imbalanced status of intracellular redox, activation of the p38 mitogen-activated protein kinase signaling pathway was induced by exposure to PS-NPs at high temperatures, which supported the decline of the reproductive capacity of D. magna. Therefore, our results suggest that PS-NPs exposure along with an increase in temperature significantly affects physiological processes triggered by damage from oxidative stress, leading to severely inhibited reproduction of D. magna.

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In aquatic environments, nanoplastics (NPls) can adsorb pharmaceuticals. However, throughout the scientific community, there is scarce knowledge about the interactive effects of the mixture nanoplastics (NPls) with pharmaceuticals to aquatic organisms. Therefore, this study aimed to investigate if the pharmaceutical diphenhydramine (DPH) toxicological effects alters when in presence of polystyrene NPls (PSNPls). To achieve this, Daphnia magna immobilization and different biochemical biomarkers (48-hours exposure) were assessed. Synergistic interactions occurred at environmentally relevant concentrations, PSNPls+DPH induced oxidative damage, whereas no effect was observed at single exposures. With the increase of PSNPls concentration, the DPH concentration causing 50% of effect (EC50) for organisms' immobilization decreased to 0.001 mg/L. In silico analysis suggested that the DPH toxicity to D. magna occurs via the sodium-dependent serotonin transporter. The results showed interactive effects between PSNPls and DPH (implying harmful effects on D. magna), allowing more thoughtful decisions by society and policymakers regarding plastics and pharmaceuticals.

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Daphnia magna is one species of water flea that has been used for a long time for ecotoxicity studies. In addition, Daphnia has a myogenic heart that is very useful for cardiotoxicity studies. Previous attempts to calculate the cardiac parameter endpoints in Daphnia suffer from the drawback of tedious operation and high variation due to manual counting errors. Even the previous method that utilized deep learning to help the process suffer from either overestimation of parameters or the need for specialized equipment to perform the analysis. In this study, we utilized DeepLabCut software previously used for animal pose tracking and demonstrated that ResNet_152 was the best fit for training the network. The trained network also showed comparable results with ImageJ and Kymograph, which was mostly done manually. In addition to that, several macro scripts in either Excel or Python format were developed to help summarize the data for faster analysis. The trained network was then challenged to analyze the potential cardiotoxicity of imidacloprid and pendimethalin in D. magna, and it showed that both pesticides cause alteration in their cardiac performance. Overall, this method provides a simple and automatic method to analyze the cardiac performance of Daphnia by utilizing DeepLabCut. The method proposed in this paper can contribute greatly to scientists conducting fast and accurate cardiotoxicity measurements when using Daphnia as a model.

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Multigenerational toxicity testing is a valuable tool for understanding the long-term effects of contaminants on aquatic organisms. This review focuses on the use of multigenerational tests with Daphnia, a widely used model organism in aquatic toxicological studies. The review highlights the importance of studying multiple generations to assess Daphnia spp. reproductive, growth, and physiological responses to various contaminants. We discuss the outcomes of multigenerational tests involving different contaminants, including nanoparticles, pesticides, and pharmaceuticals. The results reveal that multigenerational exposure can lead to transgenerational effects, where the impacts of contaminants are observed in subsequent generations even after the initial exposure has ceased. These transgenerational effects often manifest as reproduction, growth, and development alterations. Furthermore, we emphasize the need for standardized protocols in multigenerational testing to ensure comparability and reproducibility of results across studies. We also discuss the implications of multigenerational testing for ecological risk assessment, as it provides a more realistic representation of the long-term effects of contaminants on populations and ecosystems. Overall, this review highlights the significance of multigenerational tests with Daphnia in advancing our understanding of the ecological impacts of contaminants. Such tests provide valuable insights into the potential risks associated with long-term exposure to pollutants and contribute to the development of effective mitigation strategies for aquatic ecosystems.

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Heavy metals such as lead (Pb), cadmium (Cd), and arsenic (As) are of great concern in aquatic ecosystems because of their global distribution, persistence, and biomagnification via the food web. They can induce the expression of cellular protective systems (e.g., detoxification enzymes and antioxidant enzymes) to protect organisms from oxidative stress, which is a high-energy-consuming process. Thus, energy reserves (e.g., glycogen, lipids, and proteins) are utilized to maintain metabolic homeostasis. Although a few studies have suggested that heavy metal stress can modulate the metabolic cycle in crustaceans, information on changes in energy metabolism under metal pollution remains lacking in planktonic crustaceans. In the present study, the activity of digestive enzymes (amylase, trypsin, and lipase) and the contents of energy storage molecules (glycogen, lipid, and protein) were examined in the brackish water flea Diaphanosoma celebensis exposed to Cd, Pb, and As for 48 h. Transcriptional modulation of the three AMP-activated protein kinase (AMPK) and metabolic pathway-related genes was further investigated. Amylase activity was highly increased in all heavy metal-exposed groups, whereas trypsin activity was reduced in Cd- and As-exposed groups. While glycogen content was increased in all exposed groups in a concentration-dependent manner, lipid content was reduced at higher concentrations of heavy metals. The expression of AMPKs and metabolic pathway-related genes was distinct among heavy metals. In particular, Cd activated the transcription of AMPK-, glucose/lipid metabolism-, and protein synthesis-related genes. Our findings indicate that Cd can disrupt energy metabolism, and may be a potent metabolic toxicant in D. celebensis. This study provides insights into the molecular mode of action of heavy metal pollution on the energy metabolism in planktonic crustaceans.

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Since a mixed state of environmental contaminants, including microplastics (MPs), heavy metals, pharmaceuticals, and personal care products (PPCPs), exists in aquatic ecosystems, it is necessary to evaluate not only the adverse effects of exposure to a single stressor but to combined stressors. In this study, we exposed the freshwater water flea Daphnia magna to 2 µm MPs and triclosan (TCS), one of PPCPs, for 48 h to investigate the synergistic toxic consequences of simultaneous exposure to both pollutants. We measured in vivo endpoints, antioxidant responses, multixenobiotic resistance (MXR) activity, and autophagy-related protein expression via the PI3K/Akt/mTOR and MAPK signaling pathways. While MPs single exposure did not show toxic effects in water fleas, simultaneous exposure to TCS and MPs was associated with significantly greater deleterious effects in the form of increased mortality and alterations in antioxidant enzymatic activities compared with water fleas exposed to TCS alone. In addition, MXR inhibition was confirmed by measurement of the expression of P-glycoproteins and multidrug-resistance proteins in MPs-exposed groups, which led to the accumulation of TCS. Overall, these results suggest that simultaneous exposure to MPs and TCS resulted in higher TCS accumulation via MXR inhibition, leading to synergistic toxic effects such as autophagy in D. magna.

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Along with bisphenol-A (BPA), conventional phthalate esters (PAEs) have been reported as environmental hormones, despite their functional usefulness as plasticizers. Nevertheless, they are frequently found in various products, including children's utensils and toys made of poly (vinyl chloride). This is tremendously important because PAEs are harmful to infants. In addition, gel/slime-type toys made of poly (vinyl alcohol) are currently popular for developing infant' tactile senses. In this study, we developed a method to qualitatively and quantitatively detect PAEs in gel/slime-type toys mimicking, infants playing with them in a bathtub. As a result, 1,2-cyclohexanedicarboxylic acid diisononyl ester (DINCH), one of the PAE alternatives, transferred into the water from the toys and was detected most commonly (108-719 µg g-1; 0.01-0.07 wt%) among PAEs. The detected DINCH levels were below the universally accepted levels for PAEs (0.1 wt%). However, the amount of DINCH detected could still be toxic, in accordance with toxicity tests using water fleas. Furthermore, unpleasant odors were emitted when the toys containing toxic volatile organic compounds were unpacked. This is the first study to develop a method to analyze PAE in gel/slime-type toys and determine that alternatives to conventional PAEs cannot be unconditionally regarded as safe chemicals. Therefore, the revised standards for regulating PAEs and their alternatives must be reconsidered.

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The maternal effect of microplastics (MPs) toxicity is likely influenced by age and brood of test species. This study investigated the maternal effect of polyethylene MP fragments (18.23 ± 8.02 µm) with benzophenone-3 (BP-3; 2.89 ± 0.20% w/w) on chronic toxicity to Daphnia magna over two generations. Neonate (< 24 h old) and adult (5 d old) daphnids in the F0 generation were exposed until 21 d old, then first and third brood neonates in the F1 generation were recovered in clean M4 medium for 21 d. Higher chronic toxicity and maternal effect of MP/BP-3 fragments were observed in the adult group compared with the neonate group, reducing growth and reproduction in both F0 and F1 generations. First brood neonates in the F1 generation showed a higher maternal effect of MP/BP-3 fragments than third brood ones, resulting in enhanced growth and reproduction relative to the control. This study provided insights into the ecological risk of MPs containing plastic additives in the natural environment.

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The water flea Daphnia O.F. Müller 1776 (Crustacea: Cladocera) is an important model of recent evolutionary biology. Here, we report a complete genome of Daphnia (Ctenodaphnia) arabica (Crustacea: Cladocera), recently described species endemic to deserts of the United Arab Emirates. In this study, genome analysis of D. arabica was carried out to investigate its genomic differences, complexity as well as its historical origins within the subgenus Daphnia (Ctenodaphnia). Hybrid genome assembly of D. arabica resulted in ~116 Mb of the assembled genome, with an N50 of ~1.13 Mb (BUSCO score of 99.2%). From the assembled genome, in total protein coding, 5374 tRNA and 643 rRNA genes were annotated. We found that the D. arabica complete genome differed from those of other Daphnia species deposited in the NCBI database but was close to that of D. cf. similoides. However, its divergence time estimate sets D. arabica in the Mesozoic, and our demographic analysis showed a great reduction in its genetic diversity compared to other Daphnia species. Interestingly, the population expansion in its diversity occurred during the megadrought climate around 100 Ka ago, reflecting the adaptive feature of the species to arid and drought-affected environments. Moreover, the PFAM comparative analysis highlights the presence of the important domain SOSS complex subunit C in D. arabica, which is missing in all other studied species of Daphnia. This complex consists of a few subunits (A, B, C) working together to maintain the genome stability (i.e., promoting the reparation of DNA under stress). We propose that this domain could play a role in maintaining the fitness and survival of this species in the desert environment. The present study will pave the way for future research to identify the genes that were gained or lost in this species and identify which of these were key factors to its adaptation to the harsh desert environment.

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Owing to their widespread distribution and high bioaccumulation, microplastics (MPs) and mercury (Hg) are considered major threats to the ocean. MP interacts with Hg because of its high adsorption properties. However, their toxicological interactions with marine organisms, especially combined effects at the molecular level, are poorly understood. This study investigated the single and combined effects of MP and Hg on the metabolic profile of the brackish water flea Diaphanosoma celebensis. A total of 238 metabolites were significantly affected by MP, Hg, or MP + Hg. Metabolite perturbation patterns showed that toxicity of Hg and MP + Hg was similar and that of MP was not significant. Among the 223 metabolites affected by Hg, profiles of 32 unannotated metabolites were significantly different from those of MP + Hg, and combined effects of MP + Hg decreased the effect of Hg on 25 of these metabolites. Only 11 annotated metabolites were significantly affected by Hg or MP + Hg and were related to carbohydrate, lipid, vitamin, and ecdysteroid metabolism. Ten metabolites were decreased by Hg and MP + Hg and were not significantly different between the exposure groups. Enrichment analysis showed that galactose, starch, and sucrose metabolism were the most affected pathways. These findings suggest that MP has negligible toxic effect, and Hg can induce energy depletion, membrane damage, and disruption of growth, development, and reproduction. Although the impact of MP was negligible, the combined effects of MP + Hg could be metabolite specific. This study provides better understanding of the combined effects of MP and Hg on marine organisms.

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Pharmaceuticals used in human medicine contaminate freshwater ecosystems. Chemotherapeutics applied in cancer treatment are found in freshwaters at low concentrations (in the range of ng L-1) which, however, can be toxic or mutagenic to aquatic organisms. The aim of this study was to determine the impact of the alkylating/crosslinking anticancer agents, cyclophosphamide (CP) and cisplatin (CDDP), at the concentration detected in water, on Daphnia magna life history, transcriptome, and proteome. This filter feeding cladoceran is an important member of the aquatic food webs controlling algal biomass and forming basic food for planktivorous fish. Here, observations of the D. magna growth rate, age at first reproduction, and the number of eggs produced were performed in the presence of CP or CDDP. The D. magna proteins and RNA were isolated and analysed by mass spectrometry and the mRNA-seq method, respectively. Five generations of contact with the pharmaceuticals in question significantly influenced the D. magna life history parameters with the growth rate and number of laid eggs decreased, whereas age at first reproduction was increased. A decrease in survivorship was observed when daphnids were exposed to CP. These changes are the result of modifications in the gene/transcript expression followed by differences in the proteome profile in comparison to the untreated control. The proteome changes were generally in accordance with the modified transcriptome. The ecotoxicogenomics approach makes it possible to get closer to a complete picture of the influence of CP and CDDP on Daphnia. We have gathered evidence that animals in the presence of anticancer pharmaceuticals attempt to cope with permanent stress by changing their proteome and transcriptome profile. Additionally, our analyses indicate that CDDP showed a stronger effect on tested organisms than CP.

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Mercury ion (Hg2+) is a toxic heavy metal ion and Hg2+ is convertible to methylmercury (MeHg) by many aquatic microorganisms, leading to bioaccumulation and biomagnification in aquatic organisms, which can interfere with brain development and function in humans. This study employs a newly developed AIEgen (Aggregation-induced emission fluorogen) to quantify and visualise the process of MeHg bioaccumulation in vivo on the species of water flea Daphnia carinata. Two approaches to MeHg absorption were taken, either by direct incubation in a MeHg solution or by indirect consumption of algae contaminated with MeHg. We analysed the relationship between the ratio of photoluminescence (PL) intensities (I585/I480) and MeHg concentration (CMeHg) and generated a master curve for determining MeHg concentration based on the measurement of PL intensities. Fluorescent image analysis showed the occurrence of MeHg in D. carinata to be mainly in the compound eyes, optic nerve and carapace. This study indicates that MeHg absorption can be quantified and visualised in the body of zooplankton, and the MeHg transfer to zooplankton is more likely through direct exposure than via indirect food intake. The accumulation of MeHg in the eye and the nervous system could be the cause of the high mortality of D. carinata exposed to MeHg in water.

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It is well-documented that perturbation of the gut bacterial community can influence the reproductive rates of the host. Less is known about how natural ecological processes can change the bacterial composition in the gut and how such changes influence the reproductive rate of the host. Here, we provide novel experimental insights into such processes using the clonally reproducing water flea, Daphnia magna. A total of 20 replicate cultures were reared for 5 weeks (Phase 1) to allow for divergence of bacterial communities through stochastic processes (i.e. drift, founder effects, and/or colonization). Duplicate cultures created from each of these were reared for 21 days (Phase 2) while recording reproductive rates. There was a significant repeatability in reproductive rates between these duplicates, suggesting that divergence of the bacterial communities during Phase 1 translated into reproductive rate effects during Phase 2. This was further supported by significant differences in the relative abundance of gut bacteria (investigated by amplicon sequencing of a part of the 16S rRNA gene) between cultures with high and low reproductive rate in Phase 2. These results are consistent with the hypothesis that stochastic processes can cause natural variation in the bacterial composition in the gut, which in turn affect host reproductive rates.

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