Short-term variation of nutritive and metabolic parameters in <Emphasis Type="Italic">Temora longicornis</Emphasis> females (Crustacea,Copepoda) as a response to diet shift and starvation

Changes in fatty acid patterns, digestive and metabolic enzyme activities and egg production rates (EPR) were studied in the small calanoid copepod Temora longicornis. Female copepods were collected in spring 2005 off Helgoland (North Sea). In the laboratory one group of copepods was fed with the cryptophycean Rhodomonas baltica for a period of 3 days. Another group of copepods was maintained without food. According to the fatty acid patterns, animals from the field were feeding on a more detrital, animal-based and to a minor extent to a diatom-based diet. Under laboratory conditions, females rapidly accumulated fatty acids such as 18:4 (n-3), 18:3 (n-3) and 18:2 (n-6) which are specific of R. baltica. Diatom-specific fatty acids such as 16:1 (n-7) were strongly reduced. In fed animals the activities of digestive and metabolic enzymes remained constant and egg production rates were highest on day 2. Starving animals, in contrast, showed significantly reduced faecal pellet production and EPR. Proteolytic enzyme activity decreased rapidly within 24 h and remained at a low level until the end of the experiment. Citrate synthase decreased continuously as well. T. longicornis rapidly reacts to dietary changes and food depletion. It has limited energy stores and, thus, strongly depends on continuous food supply.     

Importance of copepod faecal pellets to the fate of the DSP toxins produced by Dinophysis spp.

An ingestion experiment was carried out in Rı́a de Pontevedra (Spain) with the copepod Temora longicornis in order to determine ingestion rates of the DSP toxin-producers, Dinophysis spp. (Dinophyceae), and the excretion rate of Dinophysis spp. cells within the faecal pellets. Ingestion rate was a function of dinoflagellate abundance and did not vary with either the amount, or the composition of the co-occurring phytoplankton species in the food suspension. Faecal pellet production increased at higher food concentrations. Intact Dinophysis spp. cells representing 34.4% of the total Dinophysis cells ingested by the copepods were found within the pellets. T. longicornis was the only dominant copepod species in the area that fed on Dinophysis spp., thus the pellets produced by T. longicornis were the main source of copepod “toxic” pellets in the media during blooms of Dinophysis spp. These “toxic” pellets might contribute to the maintenance of the toxic algal blooms, if the cells inside the pellets remain viable, can spread the potential toxicity of the toxic dinoflagellates throughout the pelagic food web due to coprophagy, and/or be an important toxic vector into the benthic food web. However, during a Dinophysis spp. bloom, the percentage of cells excreted daily within the pellets was lower than 1% of the total dinoflagellate population and, moreover, copepod faecal pellets represent a small fraction of the sinking material in this area. Although it was not possible to measure the amount of toxins in the pellets, we concluded that copepod faecal pellets do not have an important role in the transport of DSP toxins through the food web in this area.     

Trophic modification of food quality by heterotrophic protists: species-specific effects on copepod egg production and egg hatching

Recent literature suggests that heterotrophic protists could improve the biochemical constituents of poor quality algae for subsequent use by higher trophic levels, a phenomenon dubbed “trophic upgrading”. We conducted experiments to test trophic upgrading effects on the omnivorous calanoid copepod Acartia tonsa. Two heterotrophic protists, Oxyrrhis marina and Gymnodinium dominans, were grown on three algal species of different nutritional qualities: Dunaliella tertiolecta, Isochrysis galbana and Rhodomonas salina. In parallel incubations, the copepods were fed the heterotrophic protists or the algae that the protists grew on. We measured the ingestion rate, egg production rate and egg hatching success of the copepods in each treatment. Comparison of the egg production efficiency (EPE; egg carbon production/carbon ingestion×100%) between parallel diet treatments allowed us to calculate the trophic upgrading index (TUI; ratio of EPE-heterotrophic protist to EPE-alga). The copepods ingested similar amount, or 20-30% more algae than heterotrophic protists. Daily rations ranged 50-122% of body C per day. Copepods feeding on O. marina that grew on D. tertiolecta had higher egg production rate and egg hatching success than copepods feeding on D. tertiolecta directly, and the resultant TUI between the two treatments was 19.7. Similarly, the copepod egg production was improved by G. dominans that grew on D. tertiolecta and the resultant TUI was 23.3. TUI was near 1 between I. galbana treatment and the parallel heterotrophic protist treatments, indicating no or little trophic upgrading effects on the copepod's egg production. However, egg hatching success was significantly lower with G. dominans growing on I. galbana. O. marina that grew on R. salina induced a lower EPE relative to the alga itself, yielding a TUI of less than 1. Biochemical data showed that the heterotrophic protists contained eicosapentanoic acid (EPA) and docosahexanoic acid (DHA) even when they fed on alga that lacked these essential fatty acids, which may explain some of the observed trophic upgrading effects and species-specific differences in our experiments. However, our data also suggested that dietary EPA and DHA could not be the sole factors regulating the copepod's reproductive output, and that egg production and egg hatching appeared to have different nutritional requirements and respond differently to the copepod's diets.     

Effects of food on bacterial community composition associated with the copepod Acartia tonsa Dana

The estuarine copepod Acartia tonsa naturally carried diverse strains of bacteria on its body. The bacterial community composition (BCC) remained very conservative even when the copepod was fed different axenic algal species, indicating that the food per se did not much affect BCC associated with the copepod. In xenic algal treatments, however, copepod-associated BCC differed with each alga fed, even though the same bacterial source was used to inoculate the algae. In addition, starved copepods taken at the same location but at different times significantly differed in their BCC. Algal species composition and copepod life history therefore serve to regulate BCC associated with copepods, and spatial and temporal variations in algal species composition and copepod origin would alter bacteria–copepod interactions.     

Are the diatoms Navicula sp. and Thalassiosira fluviatilis suitable to be fed to the benthic harpacticoid copepod Tisbe biminiensis?

The objective of this study was to compare the influence of the diets of two diatoms, Navicula sp. (benthic) and Thalassiosira fluviatilis (planctonic), on the development, fecundity and survival of the harpacticoid copepod Tisbe biminiensis. In order to determine the optimal concentration of food, 35 egg-bearing females were submitted to six algal concentrations and controls (without food). After 24 h, the content of the recipients was fixed with 4% formalin and then fecal pellets produced by each female were counted and measured. The larval development was studied by surveying 50 nauplii on each diet individually until the adult stage, at intervals of 6 h. The cast exoskeletons were removed to count the number of segments and for measurement. The fecundity was obtained counting the naupliar production every 48 h of groups containing 10 females in different algal concentrations in both diets. The diet influence on fecundity was tested by submitting four groups of 10 females fed on optimal algal concentrations based on the fecal pellet experiments. In the Navicula sp. concentration of 0.4 μg Chl-a/ml, considered to be optimal for fecal pellets production, the diatoms were shown toxic, resulting in a low survival rate and inhibiting the egg production of copepods. The optimal concentration considering fecundity was estimated to be 0.1 μg Chl-a/ml for both diets. Copepods fed on Navicula sp. presented a faster development rate and higher naupliar production compared to copepods fed on T. fluviatilis. Size and survival did not vary significantly among diets. The algal concentration interfered significantly in the reproductive success of females. Both very low and very high algal concentrations reduced reproductive success. Concluding, the benthic diatom Navicula sp. was more favorable to the copepod T. biminiensis than the planktonic diatom T. fluviatilis.     

Omnivory in cyclopoid copepods: comparisons of algae and invertebrates as food for three, differenfly sized species

We examined the importance of algal versus invertebrate preyfor three cyclopoid copepods by comparing reproductive successfor females fed on three diets: exclusively algal food, exclusivelyinvertebrate prey and a combination of algal plus invertebrateprey. The three cyclopoid species represent a distinct gradientin body size: Tropocyclops prasinus mexicanus (small body size),Diacyclops thomasi (medium body size) and Mesocyclops edax (largebody size). Our results provide strong evidence for the potentialimportance of algae in the diet of adult cyclopoid copepods.However, the role of herbivory is dependent on species bodysize. We found that algae are the key dietary factor for reproductivesuccess of the small T.p.mexicanus. In contrast, the availabilityof invertebrate prey had a greater influence on reproductionby the two larger species. Overall, cyclopoid copepods shouldbe generally considered as having the potential to derive theirenergy from either animal or algal sources. This has importantconsequences for interpretations of the forces influencing zooplanktoncommunity diversity as well as for general food web theory.     

Copepod feeding behaviour and egg production during a dinoflagellate bloom in the North Sea

Feeding strategies of copepods were studied during a dinoflagellate-dominated bloom in the North Sea in August 2001. The aim of this study was to evaluate the importance of mesozooplankton grazing as a biological loss factor of harmful algal blooms under natural conditions. Therefore, ingestion, egestion and egg production experiments were performed with the most abundant copepod species Calanus helgolandicus, Temora longicornis and Acartia sp. feeding on the natural phytoplankton community. Dinophysis norvegica and Ceratium furca were the most abundant dinoflagellate species at the time of the experiments. Grazing experiments as well as examination of fecal pellet content revealed C. helgolandicus fed efficiently on D. norvegica. Ingestion rates up to 47 cells female⁻¹ h⁻¹ were measured and a large proportion of the C. helgolandicus fecal pellets contained intact D. norvegica cells. Dinophysis cells were rarely seen in fecal pellets produced by T. longicornis, and never observed in pellets produced by Acartia sp. The ingestion rate of C. furca, which was the dominating Ceratium species, mimicked that of D. norvegica. C. helgolandicus grazed significantly on C. furca (16 cells female⁻¹ h⁻¹), while the ingestion rate of T. longicornis was low and Acartia sp. was not able to graze on C. furca. Egg production experiments revealed that 92% of the C. helgolandicus females produced eggs. The specific egg production rate and the proportion of females producing eggs among T. longicornis were low. This field experiment clearly shows that some copepod species feed efficiently on D. norvegica and C. furca under natural conditions, which may affect the bloom development of these dinoflagellates.     

Foraging behaviors of two sympatric ant species in response to lizard eggs

The trade-off between behavioral dominance and resource discovery ability represents a mechanism which could facilitate the coexistence of species, but evidence of the existence of this trade-off is limited and is often derived from experiments using artificial bait. In this study, I performed a field experiment to investigate the outcome of potential food competition between an encounter species (Paratrechina longicornis) and an exploitative one (Pheidole taivanensis) and to examine the factors that may explain the behavior of P. taivanensis when obtaining food (lizard eggs) without being attacked by P. longicornis. When P. longicornis was experimentally introduced to eggs occupied by P. taivanensis for 1 day, it displaced P. taivanensis. However, P. longicornis ignored lizard eggs which had been occupied by P. taivanensis for 2 or more days, and did not displace P. taivanensis, because by that time the eggshells had been damaged by P. taivanensis so they could no longer be used by P. longicornis. Eggshells were damaged more quickly by P. taivanensis at Santimen, southwestern Taiwan, than at four other study sites where there were lower intensities of food competition between P. taivanensis and P. longicornis. The displacement percentage was higher at Santimen which had higher ant population densities. The present study shows that lizard eggs may constitute a natural, ephemeral resource for which ants compete in space and time. Comparisons between study sites with and without ants suggest the existence of a trade-off between resource discovery and territorial defense.     

A comparative approach to study inhibition of grazing and lipid composition of a toxic and non-toxic clone of Chrysochromulina polylepis (Prymnesiophyceae)

Since the massive bloom in 1988 in the North Sea, the prymnesiophyte flagellate Chrysochromulina polylepis Manton et Parke has been known for its ichtyotoxicity. Laboratory experiments using two different clones of C. polylepis were conducted in a comparative approach. Both clones were nearly similar in size and shape, but differed in their toxicity, as demonstrated by the Artemia bioassay. In order to study the effects of toxic C. polylepis on protozooplankton grazers, grazing experiments were performed with the heterotrophic dinoflagellate Oxyrrhis marina Dujardin as grazer. A first experiment was carried out in order to follow batch culture growth and initial grazing of O. marina when fed toxic or non-toxic clones of C. polylepis. Ingestion of the toxic clone was 27% of ingestion when fed with the non-toxic clone. When O. marina was fed with the toxic clone, vacuoles within O. marina contained fewer food particles per cell and the cells attained slower division rate (58% of the division rate estimated for the non-toxic clone). A second experiment was conducted to determine the grazing and growth response of O. marina as a function of algal food concentration. Profound differences in ingestion, clearance, division and gross growth efficiency of O. marina when fed the two clones of C. polylepis again were apparent. However, even at algal concentrations of 400×10³ ml⁻¹, O. marina is not killed by the presence or by the ingestion of toxic C. polylepis, indicating that the toxin deters grazers. In addition to grazing experiments, lipid classes and fatty acids of both algal clones were analysed and compared in order to follow the hypothesis that toxicity of C. polylepis is caused by liposaccharides, lipids, or fatty acids. However, the chemical composition with respect to lipid classes and fatty acids of both clones were quite similar, making an involvement of these substances in the toxicity towards Artemia and O. marina unlikely.     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

Effects of food diversity on diatom selection by harpacticoid copepods

The diversity of species assemblages which occupy a basal position in the trophic pyramid (typically unicellular algae in aquatic environments) is known to influence the interaction with organisms of higher trophic levels. A laboratory feeding experiment was conducted with cultures of three benthic diatom species (Navicula phyllepta, Grammatophora marina and Cylindrotheca closterium) as primary producers and three harpacticoid copepod species (Harpacticus obscurus, Paramphiascella fulvofasciata and Tigriopus brevicornis) as grazers to evaluate the effects of food diversity (and concomitant food availability) on grazing selectivity. This kind of selectivity experiments is singular for benthic harpacticoid copepods as so far, information on food selection of harpacticoids is scarce.Uptake of a unispecific food source by a single copepod species decreased as food diversity (and concomitant overall food concentration) increased. All three consumers reacted similarly to changing food diversity, but exhibited strong species-specific responses to food identity i.e. which diatom was added was crucial. Irrespective of level of food diversity, H. obscurus took up high amounts of G. marina, whereas both P. fulvofasciata and T. brevicornis preferred C. closterium when given the choice between different diatoms. As for zooplanktonic taxa, this experiment showed that in lower benthic marine food webs both prey organisms (primary producers) and grazers play a very specific role. Diversity of food and its identity are of critical importance at the base of the trophic pyramid, influencing trophic transfer from primary producers over grazers to higher trophic levels.     

Phenoloxidase and trypsin in germ-free larvae of Artemia fed with cooked unicellular diets: Examining the alimentary and protective effects of putative beneficial bacterium, yeast and microalgae against vibriosis

Three putative beneficial unicellular organisms, the marine bacterium Roseobacter sp, the yeast Saccharomyces cerevisiae mnn9 strain (SC-mnn9) and the microalga Tetraselmis suecica were cooked and offered separately as diets to developing germ-free (GF) Artemia larvae, in order to analyze their alimentary and protective effects. GF Artemia larvae were able to grow with cooked Tetraselmis and SC-mnn9 but failed with cooked Roseobacter. In spite of its high dietary quality, Tetraselmis failed to provide protection against Vibrio proteolyticus infection, while Roseobacter failed as food as well to provide protection. Cooked SC-mnn9 appeared to possess both values, dietary for growth and protective against Vibrio infection. GF Artemia larvae were apparently rapid adapted to dietary swaps; from yeast to algal and from algal to yeast. While the diets swap from algal or yeast, to bacterial diet appeared to be detrimental. Phenoloxidase-L (PO-L) and trypsin-L were used as biochemical indicators of defense and digestive functions, respectively. Developmental trypsin-L patterns were similar when fed on yeast and microalgae diets, suggesting a good digestive adaptation to plant or fungal substrates at early stages. On the contrary, diets swap or Vibrio infection affected PO-L and trypsin-L suggesting a sort of ‘alteration’ of digestive and defense functions.     

Assessing the In Situ Fertilization Status of Two Marine Copepod Species,Temora longicornis and Eurytemora herdmani; How Common Are Unfertilized Eggs in Nature?

We utilized an egg staining technique to measure the in situ fertilization success of two marine copepod species, Temora longicornis and Eurytemora herdmani from May to October 2008 in coastal Maine and correlated fertilization success with environmental conditions in their habitat. T. longicornis is a free spawning species that releases eggs into the ambient seawater after mating. In contrast, E. herdmani carries eggs in an egg sac until they hatch. The proportion of fertilized eggs within E. herdmani egg sacs was significantly higher than the freely spawned clutches of T. longicornis. This may be a result of the asymmetrical costs associated with carrying vs. spawning unfertilized eggs. T. longicornis frequently laid both fertilized and unfertilized eggs within their clutch. T. longicornis fertilization was negatively associated with chlorophyll concentration and positively associated with population density in their local habitat. The fertilization status of E. herdmani egg sacs was high throughout the season, but the proportion of ovigerous females was negatively associated with an interaction between predators and the proportion of females in the population. This study emphasizes that, in addition to population level processes, community and ecosystem level processes strongly influence the fertilization success and subsequent productivity of copepods.     

Incorporation of bacterial fatty acids and changes in a wax ester-based omnivory index during a long-term incubation experiment with Calanus glacialis Jaschnov

We traced the incorporation of fatty acid biomarkers into Calanus glacialis Jaschnov (CV) during a long-term incubation experiment using bacterivorous dinoflagellates and diatoms as food. Copepods fed Oxyrrhis marina Dujardin during a 3-week acclimation period developed an omnivorous lipid composition, relative to wild-captured copepods, characterized by significant losses of polyunsaturated fatty acids (PUFA) and diatom fatty acids [16:4(n−1), 20:5(n−3)], and increases in saturated fatty acids (SFA) and 18:1(n−7). Levels of a wax ester-based omnivory index [unsaturation coefficient (UC)], verified by gas chromatography (GC), also decreased in response to the relatively PUFA-poor dinoflagellate. After half of the copepods were switched to a diet comprised of the diatom Thalassiosira hispida Syvertsen (PUFA-rich), the data showed reversal to a more herbivorous lipid composition (increases in UC and relative amounts of PUFA and diatom fatty acids). We assert that UC, derived from routine thin-layer chromatography analysis (Iatroscan) can quickly determine in situ feeding strategies (i.e., degree of omnivory) of wax ester-storing copepods. None of the eight odd and/or branched fatty acids (OBFA) initially detected in C. glacialis increased in response to a diet of O. marina which was rich in these compounds [mainly iso (i)-15:0 and anteiso (ai)-15:0]. Lack of transfer of these and other fatty acids [e.g., 22:6(n−3)] could be related to the physiological state of the copepods (early diapause). We suggest that the bacterial fatty acid 18:1(n−7) may be more useful in inferring connections between Calanus spp. and the microbial food web than the odd and/or branched chains.     

Grazing by Talorchestia longicornis on an algal mat in a new England salt marsh

Grazing experiments using ¹⁴C and an analysis of fecal pellets and gut contents established that the gammaridean amphipod, Talorchestia longicornis Say, ingests blue-green algae on algal mats in a Massachusetts salt marsh. This grazing had a measurable effect on the lower algal mat, where the density of T. longicornis was high. Exclusion of amphipods resulted in increases in chlorophyll a content, carbon incorporation, and nitrogen fixation. This effect was not seen on the upper mat where T. longicornis was less abundant. The assimilation efficiency of T. longicornis feeding on a diet consisting mainly of blue-green algae was surprisingly high (67 %) considering that blue-green algae are usually considered as a poor quality food for herbivores. The population of T. longicornis seems to be annual, with growth of the overwintered juveniles in spring and early summer.     

A multi-generation Calanus finmarchicus culturing system for use in long-term oil exposure experiments

Crustacean zooplankton form the keystone link between primary producers and fish stocks along the Norwegian shelf and in the southern part of the Barents Sea. We have established a multi-generation cultivation system for zooplankton in order to perform future experiments on the biological effects of drilling discharges from the offshore petroleum industry. A population of the cold-water species, Calanus finmarchicus, was collected in March 2004 and maintained in a static system of 100 l polypropylene containers through all stages (Eggs-CVI). The population exhibited an average developmental time of 105 days which corresponds to realistic sub-Arctic to Arctic conditions (water temperatures = 5 °C (spring) and 8 °C (late summer)). A series of experiments was performed to examine copepod egg production as a function of different food sources and feeding concentrations. Only minor differences in egg production were observed for C. finmarchicus females fed with varying concentrations of the two diatom species Chaetoceros socialis and Thalassiosira weissflogii. In a separate experiment, the response to food availability was examined over time by tracking egg production and fecal pellet production (used as an indicator of food ingestion). Both hatching success and food ingestion were positively correlated with the food concentrations offered to adult females. Through this work we have demonstrated that C. finmarchicus populations can be maintained in the laboratory through multiple generations. In addition, methods to control egg production through changes in food availability have been established making it feasible to control the start date of exposure experiments or the timing of the collection of eggs to initiate a new generation.     

Selective grazing of Temora longicornis in different stages of a Phaeocystis globosa bloom—a mesocosm study

Selective grazing of a calanoid copepod Temora longicornis was measured during different stages of a Phaeocystis globosa bloom, in order to reveal (1) if T. longicornis feeds on single cells and/or colonies of P. globosa in the presence of alternative food sources, (2) if copepod food selection changes during the initiation, maintenance, collapse and decay of a P. globosa bloom and (3) if P. globosa dominated food assemblage provides a good diet for copepod egg production. Our results show low but constant feeding on small colonies of P. globosa, irrespective of the type or concentration of alternative food sources. In contrast, feeding on single cells was never significant, and the total contribution of P. globosa to carbon ingestion of T. longicornis was minor. T. longicornis fed most actively on the decaying colonies, whereas during the peak of the bloom copepods selected against P. globosa. Mostly, T. longicornis fed unselectively on different food particles: before the bloom, the major part of the diet consisted of diatoms, whereas during and after the bloom copepod diet was dominated by dinoflagellates and ciliates. Egg production was highest during the decay of the bloom, coinciding with highest proportional ingestion of heterotrophic organisms, but was not seriously reduced even during the peak of the bloom. We conclude that P. globosa blooms should not threaten survival of copepod populations, but the population recruitment may depend on the type (and concentration) of the dominant heterotrophs present during the blooms. Due to relatively unselective grazing, the impact of T. longicornis to the initiation of a Phaeocystis bloom is considered small, although grazing on decaying colonies may contribute to the faster termination of a bloom.     

Influence of Molting and Starvation on Digestive Enzyme Activities and Energy Storage in Gammarus fossarum

Among the many biological responses studied in ecotoxicology, energy-based biomarkers such as digestive enzyme activities and energy reserves appear to be useful predictive tools for detecting physiological disturbances in organisms. However, the use of these biological responses as biomarkers could be limited by the effects of confounding factors (biotic and abiotic) and physiological processes, such as the reproductive cycle. Thus, the optimal use of these biomarkers will be facilitated by understanding the effects of these factors on the energy metabolism of the sentinel species being studied. We considered abiotic factors (temperature and conductivity) in a previous study, whereas the present study investigated the effects of gender, the female reproductive stage, and food availability on the digestive enzyme activities and energy storage of Gammarus fossarum. The results indicated that, during the female reproductive cycle, the activities of digestive enzymes (amylase, cellulase, and trypsin) decreased significantly, whereas the levels of reserves (proteins, lipids, and sugar) increased until the last premolt stage. Restricted food diets only led to decreased amylase activities in both sexes. Food starvation also induced a decrease in the energy outcomes in females, whereas there were no effects in males. In general, the biochemical (digestive enzyme activities) and physiological (energy reserves) responses were more stable in males than in females. These results support the use of males fed ad libitum to limit the effects of confounding factors when using these energy biomarkers in Gammarus fossarum during biomonitoring programs.     

Direct Effects of Microalgae and Protists on Herring (Clupea harengus) Yolk Sac Larvae

This study investigated effects of microalgae (Rhodomonas baltica) and heterotrophic protists (Oxyrrhis marina) on the daily growth, activity, condition and feeding success of Atlantic herring (Clupea harengus) larvae from hatch, through the end of the endogenous (yolk sac) period. Yolk sac larvae were reared in the presence and absence of microplankton and, each day, groups of larvae were provided access to copepods. Larvae reared with microalgae and protists exhibited precocious (2 days earlier) and ≥ 60% increased feeding incidence on copepods compared to larvae reared in only seawater (SW). In the absence and presence of microalgae and protists, life span and growth trajectories of yolk sac larvae were similar and digestive enzyme activity (trypsin) and nutritional condition (RNA-DNA ratio) markedly declined in all larvae directly after yolk sac depletion. Thus, microplankton promoted early feeding but was not sufficient to alter life span and growth during the yolk sac phase. Given the importance of early feeding, field programs should place greater emphasis on the protozooplankton-ichthyoplankton link to better understand match-mismatch dynamics and bottom-up drivers of year class success in marine fish.     

Functional response,switching, and prey-stage preference of Scolothrips longicornis (Thysanoptera: Thripidae) on Schizotetranychus smirnovi (Acari: Tetranychidae)

The objective of this study was to evaluate the efficiency of Scolothrips longicornis Priesner (Thysanoptera: Thripidae) as a biocontrol agent of Schizotetranychus smirnovi Wainstein (Acari: Tetranychidae), a key pest of almond trees in the southwest of Iran. To achieve a strategy for the control of this pest, it is important to understand foraging behavior (Functional response, switching, and prey-stage preference) of S. longicornis. The predator exhibited a type III functional response when it was offered S. smirnovi protonymphs at seven densities (2, 4, 8, 16, 32 .64 and 128). Based on the random predator equation, the estimated attack rate (a), handling time (T_h), and maximum rate of predation were 0.0048 h^?1, 0.4816 h, and 49.84 per day, respectively. Using the Murdoch's model, switching behavior was observed in S. longicornis. The predator switched from one stage when it becomes rare to another more abundant stage. S. longicornis fed on all stages but preferred S. smirnovi larvae and protonymphs. The results of this study revealed that S. longicornis could be used in the integrated management of S. smirnovi. However, further field studies are needed to verify this hypothesis.