Coping with antagonistic predation risks: Predator-dependent unique responses are dominant in Ceriodaphnia cornuta.

Inducible defences of prey are evolved under diverse and variable predation risks. However, during the co-evolution of prey and multiple predators, the responses of prey to antagonistic predation risks, which may put the prey into a dilemma of responding to predators, remain unclear. Based on antagonistic predation pressure from an invertebrate (Chaoborus larvae) and a vertebrate (Rhodeus ocellatus) predator, we studied the responses of multiple traits and transcriptomes of the freshwater crustacean Ceriodaphnia cornuta under multiple predation risks. Chaoborus predation risk altered the expression of genes encoding cuticle proteins and modulated the biosynthesis of steroid hormones, cutin, suberine, and wax, leading to the development of horns and increase in size at the late developmental stage. Meanwhile, fish predation risk primarily triggered genes encoding ribosomes and those involved in unsaturated fatty acid biosynthesis and cysteine and methionine metabolism, resulting in smaller individual size and earlier reproduction. Inducible responses of both transcriptome and individual traits revealed that predator-dependent unique responses were dominant and the dilemma of antagonistic responses was relatively limited. However, the unique individual traits in response to invertebrate predation could be significantly impaired by vertebrate predation risk, even though the unique responses to different predators were extremely weakly correlated and could be elicited simultaneously. These results indicate that diverse predator-dependent unique responses are favoured by Ceriodaphnia during its co-evolution with multiple predators. Nonetheless, Ceriodaphnia is not a generalist that can fully adopt all predator-dependent unique responses simultaneously under multiple predation risks.

Understanding host-microbiome-environment interactions: Insights from Daphnia as a model organism.

Microbes perform a variety of vital functions that are essential for healthy ecosystems, ranging from nutrient recycling, antibiotic production and waste decomposition. In many animals, microbes become an integral part by establishing diverse communities collectively termed as "microbiome/s". Microbiomes defend their hosts against pathogens and provide essential nutrients necessary for their growth and reproduction. The microbiome is a polygenic trait that is dependent on host genotype and environmental variables. However, the alteration of microbiomes by stressful condition and their recovery is still poorly understood. Despite rapid growth in host-associated microbiome studies, very little is known about how they can shape ecological processes. Here, we review current knowledge on the microbiome of Daphnia, its role in fitness, alteration by different stressors, and the ecological and evolutionary aspects of host microbiome interactions. We further discuss how variation in Daphnia physiology, life history traits, and microbiome interactive responses to biotic and abiotic factors could impact patterns of microbial diversity in the total environment, which drives ecosystem function in many freshwater environments. Our literature review provides evidence that microbiome is essential for Daphnia growth, reproduction and tolerance against stressors. Though the core and flexible microbiome of Daphnia is still debatable, it is clear that the Daphnia microbiome is highly dependent on interactions among host genotype, diet and the environment. Different environmental factors alter the microbiome composition and diversity of Daphnia and reduce their fitness. These interactions could have important implications in shaping microbial patterns and their recycling as Daphnia are keystone species in freshwater ecosystem. This review provides a framework for studying these complex relationships to gain a better understanding of the ecological and evolutionary roles of the microbiome.

Disentangling Diet- and Medium-Associated Microbes in Shaping Daphnia Gut Microbiome.

Host genotype and environment are considered crucial factors in shaping Daphnia gut microbiome composition. Among the environmental factors, diet is an important factor that regulates Daphnia microbiome. Most of the studies only focused on the use of axenic diet and non-sterile medium to investigate their effects on Daphnia microbiome. However, in natural environment, Daphnia diets such as phytoplankton are associated with microbes and could affect Daphnia microbiome composition and fitness, but remain relatively poorly understood compared to that of axenic diet. To test this, we cultured two Daphnia magna genotypes (genotype-1 and genotype-2) in sterile medium and fed with axenic diet. To check the effects of algal diet-associated microbes versus free water-related microbes, Daphnia were respectively inoculated with three different inoculums: medium microbial inoculum, diet-associated microbial inoculum, and medium and diet-mixed microbial inoculum. Daphnia were cultured for 3 weeks and their gut microbiome and life history traits were recorded. Results showed that Daphnia inoculated with medium microbial inoculum were dominated by Comamonadaceae in both genotypes. In Daphnia inoculated with mixed inoculum, genotype-1 microbiome was highly changed, whereas genotype-2 microbiome was slightly altered. Daphnia inoculated with diet microbial inoculum has almost the same microbiome in both genotypes. The total number of neonates and body size were significantly reduced in Daphnia inoculated with diet microbial inoculum regardless of genotype compared to all other treatments. Overall, this study shows that the microbiome of Daphnia is flexible and varies with genotype and diet- and medium-associated microbes, but not every bacteria is beneficial to Daphnia, and only symbionts can increase Daphnia performance.

Cyanobacterial dominance and succession: Factors, mechanisms, predictions, and managements.

Eutrophication of natural water bodies worldwide has led to cyanobacteria becoming the dominant species in phytoplankton communities, causing serious harm environmentally and economically. Cyanobacterial succession makes effective treatment of cyanobacterial blooms a challenge. Although there are many studies about cyanobacterial dominance and succession, it is still lack of relevant review summarizing the advances on this topic. To control cyanobacterial blooms and manage water quality effectively, we conducted a critical review and drew the following conclusions: (1) cyanobacterial dominance and succession occur from spring to summer, with changes of multiple environmental factors dominated by temperature and nutrients conditions; (2) the cyanobacterial dominance and succession are inherently attributed to the distinctive traits of cyanobacteria including colony formation, gas vesicles, toxin release, and nitrogen fixation; (3) given the current meta-omics explorations on mechanisms of cyanobacterial succession, how to combine the extensive data to draw general conclusions is a challenge in the future; (4) the dominant niche of high temperature-adapted cyanobacteria genera will be further reinforced with global warming and elevated carbon dioxide in the future; (5) considering the causes and future developments of cyanobacterial blooms, the management strategies for controlling cyanobacterial blooms include reducing external nutrient input and removing internal nutrient in sediment, artificial mixing waters to decrease buoyancy of cyanobacteria, and biological control using allelopathy of aquatic plants and/or enhancing zooplankton feeding.

The thermal regime modifies the response of aquatic keystone species Daphnia to microplastics: Evidence from population fitness, accumulation, histopathological analysis and candidate gene expression.

The water bodies are greatly influenced by chemical contamination and global increasing temperature. As an emerging pollutant, microplastics are widely distributed in the freshwater environment, raising concerns regarding their potential toxicity to organisms. Especially for zooplankton filter feeders, many of microplastics are in similar size as their food. Individually, both microplastics and temperature have profound effects on zooplankton populations and their function in ecosystems. However, the strength and direction of their interactive effects are still not clear. Here, we performed a comprehensive biotoxicity assessment providing empirical evidence that the temperature played a key role in shaping the sensitivity of the zooplankter, Daphnia magna, against microplastic toxicity. We found that exposure to microplastics generally caused negative effects on Daphnia individual fitness, such as increased lethality, declined fecundity and reduced population growth rate. This microplastic toxicity was more prominent at 30 °C than at 20 °C, and was rather minor at 15 °C. Moreover, the warming accelerated the ingestion of microplastics, and triggered abnormal ultrastructure of intestinal epithelial cells. In addition, the expression profiling of candidate genes revealed oxidative damage, fecundity impairment and energy retardation by microplastics were amplified with increasing temperature, which may contribute to the enhancement of microplastic toxicity under warming. Given that high temperature fluctuations are becoming more common and difficult to predict, the interactive effects of microplastics and climate warming on Daphnia population dynamics and biomass production may become increasingly aggravated in nature. Collectively, extrapolation for environmental risk assessment studies conducted under different temperature contexts may broaden our knowledge microplastic toxicity on aquatic organism fitness.

Elevated temperature and toxic Microcystis reduce Daphnia fitness and modulate gut microbiota.

The gut microbiota has been increasingly recognized to regulate host fitness, which in turn is dependent on stability of community structure and composition. Many biotic and abiotic factors have been demonstrated to shape gut microbiota of cladocerans. However, the interactive effects of these variables on cladocerans fitness due to alteration of gut microbiota and their linkage with life history parameters are poorly understood. Here, we investigated the responses of Daphnia magna gut microbiota to the combined effects of toxic Microcystis aeruginosa and high temperature and its associations with fitness. We found that under good food regime, the temperature has no effect on the composition of the gut microbiota, whereas under high proportion of toxic M. aeruginosa and high temperature conditions, D. magna lost their symbionts. High proportion of toxic M. aeruginosa and high temperature had synergistically negative effects on D. magna performance due to altered gut microbiota. The high abundance of symbiotic Comamonadaceae and good food increased D. magna fitness. The present study illustrates that understanding life history strategies in response to multiple stressors related to changes in the gut microbiota diversity and composition requires integrated approaches that incorporate multiple linked traits and tether them to one another.

Changes in the life history traits of Daphnia magna are associated with the gut microbiota composition shaped by diet and antibiotics.

The gut microbiota has a crucial role in host physiology and fitness. Host-microbiota relationships can be disrupted by environmental stressors, which further affect host growth and survival. However, the link between host performance and the gut microbiota composition shaped by increasing antibiotic pollution under different food conditions is not clearly understood. In the present study, we used Daphnia magna as a model organism to investigate the interactive effects of diets (Chlorella with or without Microcystis) and antibiotics on its life history traits, gut microbiota alterations, and their relationship. The results showed that poor diet consumption by D. magna at low and high antibiotic concentrations reduced reproduction and survival. Under good diet conditions, the fitness was reduced only at a high antibiotic concentration. Under good diet conditions, high concentration of antibiotics reduced the abundance of Comamonadaceae and increased the abundance of Pseudomonadaceae, whereas under poor diet conditions, both low and high concentrations of antibiotics increased the abundance of Pseudomonadaceae. Performances of life history traits were positively correlated with an increased abundance of Comamonadaceae but were negatively correlated with increased Pseudomonadaceae abundance. The results of this study revealed the interactive effects of diet and antibiotics on D. magna fitness and correlations between bacterial abundance and life history traits, which has important implications for understanding the effects of pollutants on host-microbiota interactions through changes in phenotypes.

Insight into chronic exposure effects of nanosized titanium dioxide on Typha angustifolia leaf litter decomposition.

Advancement in nanotechnology has increased production of nanoparticles which initiates concerns for freshwater ecosystems. Nanosized TiO2 is one of the most used materials and its ecotoxicity has been extensively studied. Here, a freshwater microcosm experiment was performed to investigate the effects of nanosized (10, 60, and 100 nm) and bulk TiO2 at 1 g L-1 on the alteration in community structure of fungal decomposers and the consequences on litter decomposition of Typha angustifolia leaves. After 209 days of exposure, the decomposition rate was significantly higher in 100 nm TiO2 treatment compared to the control, which was caused by its promotion on fungal biomass and metabolic activity. Therefore, the study provides the multifaceted evidences for different effects of TiO2 with varied sizes on T. angustifolia leaf decomposition and highlights the importance of understanding the potential effects of varying sizes and long-term exposure in nanoparticle risk assessments.

Molecular characterization of thioredoxin reductase in waterflea Daphnia magna and its expression regulation by polystyrene microplastics.

Global scale concerns regarding rise in microplastics pollution in the environment have recently aroused. Ingestion of microplastics by biota, including freshwater zooplankton has been well studied, however, despite keystone species in freshwater food webs, the molecular response (e.g. oxidative defense) of zooplankton in response to microplastics is still in its infancy. The thioredoxin (TRx) system has a vital function in cellular antioxidative defense via eliminating the excessive generation of reactive oxygen species (ROS). Therefore, it is necessary to investigate the effects of thioredoxin reductase (TRxR), due to its triggering the TRx catalysis cascade. The present study identified TRxR in Daphnia magna (Dm-TRxR) for the first time, and found that the full-length cDNA was 1862 bp long, containing an 1821-bp open reading frame. Homologous alignments showed the presence of conserved catalytic domain CVNVGC and the seleocysteine (SeCys) residue (U) located in the N- and C- terminal portions. Subsequently, the expression of Dm-TRxR, together with permease, arginine kinase (AK), was investigated by approach of quantitative real-time PCR after exposure to four (1.25-µm) polystyrene (PS) microbeads concentrations: 0 (control), 2, 4 and 8 mg L-1 for 10 days. Dm-TRxR, permease and AK mRNA were significantly upregulated after exposure to 2, 4 mg L-1 of PS, but then declined in the presence of 8 mg L-1 PS. The gene expression results suggested that oxidative defense, energy production and substance extra cellular transportation were significantly regulated by microplastic exposure. Collectively, the present study will advance our knowledge regarding the biological effects of microplastic pollution on zooplankton, and builds a foundation for freshwater environmental studies on mechanistic and biochemical responses to microplastics.

Temperature-dependent elimination efficiency on Phaeocystis globosa by different initial population sizes of rotifer Brachionus plicatilis.

Due to sea water eutrophication and global warming, the harmful Phaeocystis blooms outbreak frequently in coastal waters, which cause a serious threat to marine ecosystem. The application of rotifer to control the harmful alga is a promising way. To investigate the influence of initial rotifer density and temperature on the ability of rotifer Brachionus plicatilis to eliminate Phaeocystis globosa population, we cultured P. globosa with different initial rotifer densities (1, 3, 5 inds mL-1) at 19, 22, 25, 28, and 31 °C for 9-16 d. Results showed that the population of rotifer feeding on Phaeocystis increased rapidly and higher temperatures favored the growth of P. globosa and B. plicatilis. With increased initial rotifer density and temperature, both the clearance rate of rotifer and the reduction rate of P. globosa increased, and thus P. globosa were eliminated earlier. Both temperature and initial rotifer density had significant effects on clearance rate of rotifer and the time to Phaeocystis extinction, and there was a significant interaction between the two factors on the two parameters, i.e., the effect of initial rotifer density on eliminating Phaeocystis decreased with increasing temperature. The rotifer in 5 inds mL-1 at 28 °C eliminated P. globosa in 4 d, whereas the rotifer in 1 ind mL-1 at 19 °C spent about 16 d on eliminating P. globosa. In conclusion, higher temperature and bigger initial rotifer density promote rotifer to eliminate the harmful P. globosa, and the optimal temperature for rotifer to clear P. globosa is 28 °C.

Understanding interactive inducible defenses of Daphnia and its phytoplankton prey.

Cyanobacterial and zooplankton inducible defenses are important but understudied process that regulate the trophic interactions of freshwater ecosystem. Daphnia due to its large size is considered an important zooplankton with the high potential to control cyanobacterial blooms. It has been shown that Daphnia through maternal induction transfer tolerance to their next generation against Microcystis toxicity. Maternal induction has been investigated in different Daphnia species without considering phenotypic plasticity of prey. Laboratory experiments were performed to explore cyanobacteria-Daphnia inducible defenses in order to better understand their interactions. Two Daphnia species were fed either with Microcystis aeruginosa PCC7806 (Ma) or Microcystis flos-aquae (Mf) mixed with Chlorella vulgaris (Cv) (exposed Daphnia), and or pure Cv (unexposed Daphnia). Exposed prey cultures were produced by prior exposure to Daphnia infochemicals. Neonates produced by exposed and unexposed Daphnia were fed with mixed diet (Microcystis+Cv) of either exposed and or unexposed prey. Growth parameters and toxin production of exposed prey cultures were significantly different than that of control. Exposed Daphnia fecundity and survival was higher as compared to unexposed Daphnia. Growth and reproduction was reduced in exposed Daphnia when fed with exposed prey as compared to those fed with unexposed prey. This study provides information on the interactive inducible defenses between cyanobacteria and its grazer under laboratory conditions and may increase our understanding of cyanobacteria and Daphnia interactions in the freshwater ecosystem.

The importance of calcium in improving resistance of Daphnia to Microcystis.

Changing environmental calcium (Ca) and rising cyanobacterial blooms in lake habitats could strongly reduce Daphnia growth and survival. Here, we assessed the effects of maternal Ca in Daphnia on transfer of resistance to their offspring against Microcystis aeruginosa PCC7806 (M. aeruginosa). Laboratory microcosm experiments were performed to examine effects in Daphnia carinata (D. carinata) and Daphnia pulex (D. pulex), and that how Ca induce responses in their offspring. The results showed that growth and survival were increased in offspring from exposed Daphnia as compared to unexposed, when raised in high Ca and increasing M. aeruginosa concentration. Among exposed Daphnia, offspring from high Ca mothers, produced more neonates with large size and higher survival as compared to offspring from low maternal Ca. Exposed D. carinata and D. pulex offspring, when reared in Ca deficient medium and increasing M. aeruginosa concentration, time to first brood increased, size become large and total offspring decreased subsequently in three alternative broods in offspring from low maternal Ca. In contrast, growth and reproduction in offspring from high Ca exposed mothers were consistent in three alternative broods. Despite species specific responses in growth, survival and variant life history traits in two Daphnia species, our results not only show maternal induction in Daphnia but also highlight that offspring response to M. aeruginosa varies with maternal Ca. This study demonstrates that Ca have role in Daphnia maternal induction against Microcystis, and recent Ca decline and increasing Microcystis concentration in lakes may decrease Daphnia growth and survival. Our data provide insights into the interactive effect of maternal Ca and Microcystis exposure on Daphnia and their outcome on offspring life history traits and survival.

Effect of heavy metals combined stress on growth and metals accumulation of three Salix species with different cutting position.

This study aimed to compare growth performance and heavy metal (HM) accumulation at different cutting positions of Salix species grown in multi-metal culture. Three Salix species stems cut at different positions (apical to basal) were grown hydroponically for four weeks. The plants were then treated for three weeks with 0, 5, 10, and 20 µM Cd, Cu, Pb, and Zn, resulting in total metal concentrations of 0, 20, 40, and 80 µM. The growth parameters and HM content in shoots and initial cutting were measured. Results showed that, compared with S. fragilis, S. matsudana grew more poorly in uncontaminated condition but grew better and accumulated lower metal in shoots under mixed HM treatment. In addition, cuttings from apical parent stem position exhibited poorer growth performance before and after treatment, as well as greater metal content in shoots than base parts under the HM treatment. These results suggest that S. matsudana may undergo a special mechanism to hinder metals in the initial cutting, thus mitigating growth damage. The apical portion also showed poor resistance against the invasion of mixed HMs because of the immature structure. Therefore, in the selection of phytoremediation plants, metal accumulation ability is not proportional to growth performance.