Diapause of Cyclops vicinus (Uljanin) in Lake Søbygård: Indication of a Risk-spreading Strategy

Cyclops vicinus is reported to enter summer diapause triggered by day length in order to survive food scarcity and fish predation. Development ceases and the fourth or fifth copepodid stages persist in the sediment for several weeks. In Lake Søbygård, however, a small eutrophic lake in Denmark, C. vicinus is found in the sediment as well as in the open water throughout the year. We performed laboratory experiments to elucidate the life cycle strategy of this population. In order to find the factors that induce diapause we tested the influence of food composition and light regime on the proportion of copepods entering diapause. Further we examined the diapause pattern of the offspring from diapausing and non-diapausing parents to see whether the co-occurrence of diapausing and non-diapausing copepods results from genetic differentiation within the population. In all experiments some of the copepods developed directly into adults, while others remained at the fourth or fifth copepodid stage and displayed diapause features. The proportion of copepods that developed directly into adults was influenced by food supply. Fewer individuals entered diapause when they received a mixed diet of algae and ciliates or algae, ciliates and seston compared with a pure algal diet. The response to light was different than described in literature: about 30% of the copepods entered diapause under dark conditions, and the diapause frequency was not higher when copepods had been exposed to continuous light. The offspring from diapausing parents showed no higher tendency to enter diapause than the offspring of non-diapausing parents, indicating no genetic differentiation. We conclude that diapause in C. vicinus represents a risk-spreading strategy, modified by food. The simultaneous production of diapausing and non-diapausing offspring ensures survival under harsh conditions; the influence of the food supply on the proportion of individuals entering diapause may adjust the population to the actual environmental condition. This flexible life cycle strategy might contribute to the exclusive dominance of C. vicinus in Lake Søbygård.     

Marine microalgae attack and feed on metazoans

Free-living microalgae from the dinoflagellate genus Karlodinium are known to form massive blooms in eutrophic coastal waters worldwide and are often associated with fish kills. Natural bloom populations, recently shown to consist of the two mixotrophic and toxic species Karlodinium armiger and Karlodinium veneficum have caused fast paralysis and mortality of finfish and copepods in the laboratory, and have been associated with reduced metazooplankton biomass in-situ. Here we show that a strain of K. armiger (K-0688) immobilises the common marine copepod Acartia tonsa in a density-dependent manner and collectively ingests the grazer to promote its own growth rate. In contrast, four strains of K. veneficum did not attack or affect the motility and survival of the copepods. Copepod immobilisation by the K. armiger strain was fast (within 15 min) and caused by attacks of swarming cells, likely through the transfer and action of a highly potent but uncharacterised neurotoxin. The copepods grazed and reproduced on a diet of K. armiger at densities below 1000, cells ml⁻¹, but above 3500 cells ml⁻¹ the mixotrophic dinoflagellates immobilised, fed on and killed the copepods. Switching the trophic role of the microalgae from prey to predator of copepods couples population growth to reduced grazing pressure, promoting the persistence of blooms at high densities. K. armiger also fed on three other metazoan organisms offered, suggesting that active predation by mixotrophic dinoflagellates may be directly involved in causing mortalities at several trophic levels in the marine food web.     

Trade-Offs between Predation Risk and Growth Benefits in the Copepod Eurytemora affinis with Contrasting Pigmentation

Intraspecific variation in body pigmentation is an ecologically and evolutionary important trait; however, the pigmentation related trade-offs in marine zooplankton are poorly understood. We tested the effects of intrapopulation phenotypic variation in the pigmentation of the copepod Eurytemora affinis on predation risk, foraging, growth, metabolic activity and antioxidant capacity. Using pigmented and unpigmented specimens, we compared (1) predation and selectivity by the invertebrate predator Cercopagis pengoi, (2) feeding activity of the copepods measured as grazing rate in experiments and gut fluorescence in situ, (3) metabolic activity assayed as RNA:DNA ratio in both experimental and field-collected copepods, (4) reproductive output estimated as egg ratio in the population, and (5) total antioxidant capacity. Moreover, mitochondrial DNA (mtDNA) COI gene variation was analysed. The pigmented individuals were at higher predation risk as evidenced by significantly higher predation rate by C. pengoi on pigmented individuals and positive selection by the predator fed pigmented and unpigmented copepods in a mixture. However, the antioxidant capacity, RNA:DNA and egg ratio values were significantly higher in the pigmented copepods, whereas neither feeding rate nor gut fluorescence differed between the pigmented and unpigmented copepods. The phenotypic variation in pigmentation was not associated with any specific mtDNA genotype. Together, these results support the metabolic stimulation hypothesis to explain variation in E. affinis pigmentation, which translates into beneficial increase in growth via enhanced metabolism and antioxidant protective capacity, together with disadvantageous increase in predation risk. We also suggest an alternative mechanism for the metabolic stimulation via elevated antioxidant levels as a primary means of increasing metabolism without the increase in heat absorbance. The observed trade-offs are relevant to evolutionary mechanisms underlying plasticity and adaptation and have the capacity to modify strength of complex trophic interactions.     

Individual variations of the phospholipid compositions of the organs of arthropods, echinoderms, and tunicates from peter the Great Bay of the Sea of Japan

Individual variations in the phospholipid compositions of the organs of marine invertebrates were investigated in the representatives of three phyla: Arthropoda [decapods: Paralithodes camtschaticus (Tilesius, 1815) and Erimacrus isenbeckii (Brandt, 1848)]; Echinodermata [starfishes: Distolasterias nipon (Doderlein, 1902) and Evasterias echinosoma Fisher, 1926]; and Tunicata [ascidians: Halocynthia aurantium (Pallas, 1787), H. roretzi (Drasche, 1884), and Styela clava Herdman, 1881]. The specificity of individual phospholipid variation was shown to be related to the systematic status and functional properties of the organs of marine invertebrates. Phosphatidylcholine and phosphatidylethanolamine were the least variable. Low variation of phospholipid composition was found in the liver, legs, and gills of decapods; in the liver of starfishes; and in the gonads and skin-muscular and gill sacs of ascidians. The contents of ceramide aminoethylphosphonate (CAEP) and phosphatidylglycerol (PG) showed the highest variation. These phospholipids were found but not in all individuals or organs studied. After 7 days of starvation, CAEP completely disappeared from the organs of the animals; thus, we suggest its exogenous origin.     

Lethal and Sublethal Toxicity Comparison of BFRs to Three Marine Planktonic Copepods: Effects on Survival,Metabolism and Ingestion

The estuarine planktonic copepods have a wide geographical distribution and commendable tolerance to various kinds of contaminants. The primary aim of the present study was to contrast the impacts of model POPs (TBBPA and HBCD) on three common estuarine planktonic copepods (Oithona similis, Acartia pacifica and Pseudodiaptomus inopinus) and establish a protocol for the assessment of acute toxicity of marine organic pollutants. We first quantified the 96h-LC₅₀ (0.566, 0.04 and 0.257 mg/L of TBBPA to the three subjects above respectively and 0.314 mg/L of HBCD to P. inopinus; all reported concentrations are nominal values). In the sub-lethal toxicity tests, it was turned out that the effects of copepods exposed to TBBPA could product different influences on the energy ingestion and metabolism. Different type of pollutions, meanwhile, could also bring varying degree effect on the target copepods. In general, the indicators (the rate of oxygen consumption, ammonia excretion, food ingestion and filtration) in higher concentration groups showed marked significant difference compared with controls as well a dose-effect relationship. The study also extended the research on the joint toxicity of TBBPA and HBCD based on the survival rate of P.inopinus. Whether 1:1 concentration or 1:1 toxic level, the research showed synergy effect relative to single exposure conditions. The result indicated that current single ecological testing used for environmental protection activities may underestimate the risk for copepods. It was also demonstrated that short-term sub-lethal experiment could be a standard to evaluate the sensitivity of copepods to POPs.     

Ocean Acidification Affects the Phyto-Zoo Plankton Trophic Transfer Efficiency

The critical role played by copepods in ocean ecology and biogeochemistry warrants an understanding of how these animals may respond to ocean acidification (OA). Whilst an appreciation of the potential direct effects of OA, due to elevated pCO₂, on copepods is improving, little is known about the indirect impacts acting via bottom-up (food quality) effects. We assessed, for the first time, the chronic effects of direct and/or indirect exposures to elevated pCO₂ on the behaviour, vital rates, chemical and biochemical stoichiometry of the calanoid copepod Acartia tonsa. Bottom-up effects of elevated pCO₂ caused species-specific biochemical changes to the phytoplanktonic feed, which adversely affected copepod population structure and decreased recruitment by 30%. The direct impact of elevated pCO₂ caused gender-specific respiratory responses in A.tonsa adults, stimulating an enhanced respiration rate in males (> 2-fold), and a suppressed respiratory response in females when coupled with indirect elevated pCO₂ exposures. Under the combined indirect+direct exposure, carbon trophic transfer efficiency from phytoplankton-to-zooplankton declined to < 50% of control populations, with a commensurate decrease in recruitment. For the first time an explicit role was demonstrated for biochemical stoichiometry in shaping copepod trophic dynamics. The altered biochemical composition of the CO₂-exposed prey affected the biochemical stoichiometry of the copepods, which could have ramifications for production of higher tropic levels, notably fisheries. Our work indicates that the control of phytoplankton and the support of higher trophic levels involving copepods have clear potential to be adversely affected under future OA scenarios.     

Distribution,diversity and density of wolbachial infections in cladocerans and copepods from Thailand

Species of the genus Wolbachia comprise a group of Rickettsia-like, maternally-inherited bacteria that cause several reproductive alterations in arthropod hosts. The best known are cytoplasmic incompatibility and feminization. Here, the first systematic surveys of wolbachial infections in cladocerans and copepods from six geographic regions of Thailand, including Northern, Northeastern, Western, Central, Eastern and Southern are reported. Using gene amplification assays with wsp and groE primers, wolbachiae were detected in 239 (4 spp.) of 1885 (57 spp.) copepods and cladocerans from all regions of Thailand surveyed. Screening results obtained with wsp primers or groE primers were similar in all cases. The presence of wolbachiae was only detected in copepods, not in cladocerans. Sex ratio analyses of the progeny of two species of copepods, Mesocyclops aspericornis and Mesocyclops thermocyclopoides, naturally or artificially infected with wolbachiae showed infection causes feminization (female-bias). The relative density if infection in naturally infected populations of three copepod species, M. thermocyclopoides, Heliodiaptomus elegans and Neodiaptomus blachei, were determined using real-time quantitative PCR assay based on the wsp gene. The density of wolbachiae in M. thermocyclopoides was significantly higher than in the other two species. These results suggest that wolbachial infections are distributed throughout Thailand, and that possibly the natural occurrence of these in copepods may be due to their predation on mosquito larvae. This apparent novel biology may have importance as a genetic drive system for control of vector borne diseases in the future.     

The life cycle of Cyclops vicinus in Lake Søbygård: new aspects derived from sediment analyses

Cyclops vicinus is the only copepod species in the pelagic zone of Lake Søbygård and can be found there all year round. We studied the population dynamics of this copepod over a one-year period. In contrast to earlier studies we included the copepods in the pelagic zone as well as the copepods resting in the sediment. Cyclops vicinuswas found not only in the open water, but also in the sediment throughout the year. From the fluctuations of the abundances in both habitats we suggest that the life cycle is more complex than known so far: different diapause pattern appear within the population and the induction of diapause is different than in other populations. We assume that these features contribute to the predominance of C. vicinusin Lake Søbygård.     

The trophic importance of algal turfs for coral reef fishes: the crustacean link

On coral reefs, the epilithic algal matrix (EAM) is widely recognised as an important resource for herbivorous and detritivorous fishes. In comparison, little is known of the interaction between benthic carnivores and the EAM, despite the abundance of Crustacea within the EAM. The trophic importance of the EAM to fishes was investigated in Pioneer Bay, Orpheus Island, Great Barrier Reef. Fish densities were quantified using visual and clove oil censuses, and gut content analyses conducted on abundant fish species. Crustaceans were found to be an important dietary category, contributing between 49.5 and 100 % of the gut contents, with harpacticoid copepods being the dominant component. Of the benthic carnivores, the goby Eviota zebrina was found to consume the most harpacticoids with a mean of 249 copepods m^?2 day^?1. This represents approximately 0.1 % of the available harpacticoid population in the EAM. In a striking comparison, herbivorous parrotfishes were estimated to consume over 12,000 harpacticoids m^?2 day^?1, over 27 times more than all benthic carnivores surveyed, representing approximately 5.3 % of the available harpacticoid copepod population each day. The high consumption of harpacticoid copepods by benthic carnivores and parrotfishes indicates that harpacticoids form an important trophic link between the EAM and higher trophic levels on coral reefs.     

Mesozooplankton Grazing on Picocyanobacteria in the Baltic Sea as Inferred from Molecular Diet Analysis

Our current knowledge on the microbial component of zooplankton diet is limited, and it is generally assumed that bacteria-sized prey is not directly consumed by most mesozooplankton grazers in the marine food webs. We questioned this assumption and conducted field and laboratory studies to examine picocyanobacteria contribution to the diets of Baltic Sea zooplankton, including copepods. First, qPCR targeting ITS-1 rDNA sequence of the picocyanobacteria Synechococcus spp. was used to examine picocyanobacterial DNA occurrence in the guts of Baltic zooplankton (copepods, cladocerans and rotifers). All field-collected zooplankton were found to consume picocyanobacteria in substantial quantities. In terms of Synechococcus quantity, the individual gut content was highest in cladocerans, whereas biomass-specific gut content was highest in rotifers and copepod nauplii. Moreover, the gut content in copepods was positively related to the picocyanobacteria abundance and negatively to the total phytoplankton abundance in the water column at the time of sampling. This indicates that increased availability of picocyanobacteria resulted in the increased intake of this prey and that copepods may rely more on picoplankton when food in the preferred size range declines. Second, a feeding experiments with a laboratory reared copepod Acartia tonsa fed a mixture of the picocyanobacterium Synechococcus bacillaris and microalga Rhodomonas salina confirmed that copepods ingested Synechococcus, even when the alternative food was plentiful. Finally, palatability of the picocyanobacteria for A. tonsa was demonstrated using uptake of ¹³C by the copepods as a proxy for carbon uptake in feeding experiment with ¹³C-labeled S. bacillaris. These findings suggest that, if abundant, picoplankton may become an important component of mesozooplankton diet, which needs to be accounted for in food web models and productivity assessments.     

Role of Zooplankton Diversity in Vibrio cholerae Population Dynamics and in the Incidence of Cholera in the Bangladesh Sundarbans

Vibrio cholerae, a bacterium autochthonous to the aquatic environment, is the causative agent of cholera, a severe watery, life-threatening diarrheal disease occurring predominantly in developing countries. V. cholerae, including both serogroups O1 and O139, is found in association with crustacean zooplankton, mainly copepods, and notably in ponds, rivers, and estuarine systems globally. The incidence of cholera and occurrence of pathogenic V. cholerae strains with zooplankton were studied in two areas of Bangladesh: Bakerganj and Mathbaria. Chitinous zooplankton communities of several bodies of water were analyzed in order to understand the interaction of the zooplankton population composition with the population dynamics of pathogenic V. cholerae and incidence of cholera. Two dominant zooplankton groups were found to be consistently associated with detection of V. cholerae and/or occurrence of cholera cases, namely, rotifers and cladocerans, in addition to copepods. Local differences indicate there are subtle ecological factors that can influence interactions between V. cholerae, its plankton hosts, and the incidence of cholera.     

Selection and viability after ingestion of vegetative cells, resting spores and resting cells of the marine diatom, Chaetoceros pseudocurvisetus, by two copepods

Feeding response of two copepods Neocalanus flemingeri and Calanus sinicus on cultures of three life-forms; vegetative cells, resting spores and resting cells, of Chaetoceros pseudocurvisetus was investigated. N. flemingeri fed heavily on the vegetative cells but scarcely responded to feed on the resting spores. C. sinicus showed significantly higher filtering rate on the vegetative cells and resting cells than on the resting spores. Survival of the three life-forms of C. pseudocurvisetus after gut passage of the copepods was also studied. The resting spores could germinate from fecal pellets of both N. flemingeri and C. sinicus; however, both the vegetative cells and the resting cells could not survive ingestion by the copepods. These results suggest that resting spore forming diatoms, such as C. pseudocurvisetus form spores which have a low nutritional value and during gut passage are largely indigestible due to the heavily silicified frustules and thus minimize the effects of grazing by copepods.     

The copepod Centropages abdominalis as a carrier of the stalked ciliate Zoothamnium

Adult specimens of the calanoid copepod Centropages abdominalis collected from February through April 1986 in Shinhamako, a saline lake connected to Tokyo Bay, were heavily infested with the stalked ciliate Zoothamnium. The number of copepods and those infested varied during the infestation period; in February when the copepod population was large, the incidence of infestation was low, whereas in March when the copepod population was small, copepods infested with Zoothamnium were proportionally more abundant. Numerical variations of infested copepods may correspond to changes in the Zoothamnium population. No specimens of Acartia omorii, which coexists with C. abdominalis, were infested in 1986, although a small number of A. omorii were infested with peritrich ciliates in 1985. Some shrimp larvae were also covered with the same peritrichs as copepods. Nevertheless, as a whole, the relationship between the phoront and its carrier seems to be specific. The highest incidence of infested copepods was approximately 30% similar to that in 1985. The sex ratio of C. abdominalis changed on each sampling date but in general females were more numerous than males. The percentage of infested males was higher than that of infested females, suggesting that the former can be attached to more easily than the latter.     

Predation by Acartia tonsa Dana on planktonic ciliates and rotifers

In laboratory experiments, adults and nauplii of the calanoid copepod Acartia tonsa Dana feed on planktonic ciliates and rotifers. Adults have a higher clearing rate for planktonic ciliates and rotifers than for phytoplankton. Adult copepods have a maximum clearing rate of ≈200 ml copepod⁻¹ · d⁻¹ for large ciliates, with lower clearing rates for small ciliates. Nauplii have higher clearance for small than for large ciliates. Addition of ciliates or rotifers to the diet of A. tonsa enhances egg production; this effect is due to improved food quality. Microzooplankton may be an important component of the diet of A. tonsa even when phytoplankton are plentiful. Selective predation by copepods probably influences the population dynamics of planktonic ciliates and rotifers in coastal waters.     

Impacts of very warm temperature on egg production rates of three Acartiidae (Crustacea,Copepoda) in a Northern African lagoon

The daily Egg Production Rate (EPR) of the three Acartiidae copepods namely Acartia clausi, Paracartia latisetosa and Paracartia grani in the North Lagoon of Tunis was compared to temperature and food availability every season from March, 2003 to February, 2004 corresponding to the major seasonal peaks of their abundance and over a 20-days period. The daily EPR was evaluated by assuming a direct correlation with chlorophyll a, particulate organic carbon (POC), and some easily extractable macromolecular compounds from the seston, such as proteins, carbohydrates and lipids. The results showed significant differences in EPR between seasons and species. Temperature and salinity negatively affected the fecundity of A. clausi and positively that of P. latisetosa and P. grani. The A. clausi EPR was supported by chlorophyll a and sestonic proteins while that of P. latisetosa and P. grani correlated with POC suggesting that these two taxa feed on microzooplankton and detritus more than on phytoplankton. Overall, EPR data showed potential population recruitment evidently the highest in P. latisetosa in summer, P. grani in autumn and A. clausi in winter. Our results indicate that temperature and food quality are the determining factors of both abundance and reproduction of the copepods under study.     

人为干扰对乌海市四合木小灌木景观的影响

以乌海市四合木(Tetraena mongolica Maxim)为研究对象,利用2009年Quickbird高分辨率影像,结合四合木群落野外调查数据,对乌海市四合木分布区在不同人为干扰强度下的景观格局和群落进行研究。结果表明:人为干扰越强烈,四合木生境破碎化程度越严重。在植株生理年龄相近,生境相同时,四合木的高度、冠幅以及重要值随人为干扰加剧,变化趋势明显。受人为干扰程度越强地区,其物种丰富度指数越小,生物多样性越低。干扰强度发生变化时,Pielou均匀度指数均在0.77—0.90间。     

Tolerance of copepods to short-term thermal stress caused by coastal power stations

The tolerance of marine copepods to short-term thermal stress was measured by the median lethal temperature (LT₅₀) tests in laboratory. Experiments on LT₅₀ of copepods from different acclimation and acclimatization conditions collected from the Yueqing Bay were carried out under heat exposure for 15, 30 and 45 min. The LT₅₀ of copepods decreased with increasing exposure time but increased with rising acclimation and acclimatization temperatures. However, the differences in copepod LT₅₀ decreased with rising acclimatization temperatures, which suggested that entrained copepod mortality increased with raised water temperature due to the acute thermal stress of coastal power stations. Results also revealed that the thermal tolerance of Labidocera euchaeta was much higher than that of Calanus sinicus in spring. The thermal tolerances of different copepod species in summer were in the order, Pseudodiaptomus marinus, Acartia spinicauda, Acartia pacifica and L. euchaeta.     

What Drives the Occurrence of the Melioidosis Bacterium Burkholderia pseudomallei in Domestic Gardens?

Melioidosis is an often fatal infectious disease affecting humans and animals in tropical regions and is caused by the saprophytic environmental bacterium Burkholderia pseudomallei. Domestic gardens are not only a common source of exposure to soil and thus to B. pseudomallei, but they also have been found to contain more B. pseudomallei than other environments. In this study we addressed whether anthropogenic manipulations common to gardens such as irrigation or fertilizers change the occurrence of B. pseudomallei. We conducted a soil microcosm experiment with a range of fertilizers and soil types as well as a longitudinal interventional study over three years on an experimental fertilized field site in an area naturally positive for B. pseudomallei. Irrigation was the only consistent treatment to increase B. pseudomallei occurrence over time. The effects of fertilizers upon these bacteria depended on soil texture, physicochemical soil properties and biotic factors. Nitrates and urea increased B. pseudomallei load in sand while phosphates had a positive effect in clay. The high buffering and cation exchange capacities of organic material found in a commercial potting mix led to a marked increase in soil salinity with no survival of B. pseudomallei after four weeks in the potting mix sampled. Imported grasses were also associated with B. pseudomallei occurrence in a multivariate model. With increasing population density in endemic areas these findings inform the identification of areas in the anthropogenic environment with increased risk of exposure to B. pseudomallei.     

Bacteria Associated with the Surface and Gut of Marine Copepods

Little is known about the nature of bacteria associated with the surface and gut of marine copepods, either in laboratory-reared animals or in the natural environment. Nor is it known whether such animals possess a gut flora. The present report deals with studies of microorganisms isolated from healthy, laboratory-reared copepods of the species Acartia tonsa Dana, from several species of wild copepods collected from a marine or estuarine environment, and from laboratory dishes containing moribund copepods. Evidence for a unique gut flora in laboratory-reared animals is presented; the predominant bacteria were represented by the genus Vibrio. Other organisms such as Pseudomonas and Cytophaga were found less abundantly associated with the copepods and not specifically associated with the gut.