Ingestion of the cyanobacterium Trichodesmium spp. by pelagic harpacticoid copepods Macrosetella,Miracia and Oculosetella

Trichodesmium is a filamentous, colonial nitrogen fixing cyanobacteria, ubiquitous in tropical and subtropical regions of the world's oceans. Trichodesmium fixes atmospheric nitrogen and can comprise a significant fraction of total primary production in oceanic surface waters. Therefore, the consumption and fate of Trichodesmium has important consequences for understanding carbon and nitrogen cycling in the open ocean. The pelagic harpacticoid copepod Macrosetella gracilis uses Trichodesmium not only as a physical substrate for juvenile development, but also as a food source. Several different types of pelagic copepods (including several species of calanoids, harpacticoids and a poecilostomatoid species) were tested for ingestion of Trichodesmium by labelling the cyanobacteria with ¹⁴C. Only the pelagic harpacticoids ingested Trichodesmium. Here we report the first grazing rates based on ¹⁴C-uptake measurements for Macrosetella gracilis (0.173 µg C copepod^–1 h^–1), and the first quantitative measurements of both Miracia efferata (0.402 µg C copepod^–1 h^–1) and Oculosetella gracilis (0.126 µg C copepod^–1 h^–1) ingesting this cyanobacteria. Ingestion rates of M. gracilis and M. efferata on the two different species of Trichodesmium, T. thiebautii and T. erythraeum, as well as the two different colonial morphologies of T. thiebautii, spherical-shaped (puffs) and fusiform (tufts), were also compared. Both Miracia and Macrosetella had higher ingestion rates on the puff colonies than the tuft colonies of T. thiebautii.. Both also had higher ingestion rates of T. erythraeum than T. thiebautii. Trichodesmium thiebautii contains a previously reported neurotoxin which may be an important factor in determining trophodynamic interactions. Our results suggest that pelagic harpacticoid copepods can be quantitatively important in determining the fate of Trichodesmium carbon and nitrogen.     

GENOTYPIC RELATIONSHIPS IN TRICHODESMIUM (CYANOPHYCEAE) BASED ON nifH SEQUENCE COMPARISONS1

The DNA sequence of a fragment of nifH was compared to natural populations of the marine cyanobacteria Trichodesmium thiebautii and T. erythraeum from the Caribbean Sea and the unialgal culture Trichodesmium sp. NIBB 1067, which was isolated from the Kuroshio waters (Japan). Through replication Of amplification, cloning, and sequencing, four nucleotides in a 359-bp fragment were identified that were identical in sequence to Trichodesmium sp. NIBB 1067 and natural populations of T. erythraeum but were distinctly different in sequence from T. thiebautii. The data indicate that Trichodesmium sp. NIBB 1067 is more closely related to T. erythraeum than to T. thiebautii.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

相似文献   

Fitness consequences for copepods feeding on a red tide dinoflagellate: deciphering the effects of nutritional value, toxicity, and feeding behavior

Phytoplankton exhibit a diversity of morphologies, nutritional values, and potential chemical defenses that could affect the feeding and fitness of zooplankton consumers. However, how phytoplankton traits shape plant–herbivore interactions in the marine plankton is not as well understood as for terrestrial or marine macrophytes and their grazers. The occurrence of blooms of marine dinoflagellates such as Karenia brevis suggests that, for uncertain reasons, grazers are unable to capitalize on, or control, this phytoplankton growth—making these systems appealing for testing mechanisms of grazing deterrence. Using the sympatric copepod Acartia tonsa, we conducted a mixed diet feeding experiment to test whether K. brevis is beneficial, toxic, nutritionally inadequate, or behaviorally rejected as food relative to the palatable and nutritionally adequate phytoplankter Rhodomonas lens. On diets rich in K. brevis, copepods experienced decreased survivorship and decreased egg production per female, but the percentage of eggs that hatched was unaffected. Although copepods showed a 6–17% preference for R. lens over K. brevis on some mixed diets, overall high ingestion rates eliminated the possibility that reduced copepod fitness was caused by copepods avoiding K. brevis, leaving nutritional inadequacy and toxicity as remaining hypotheses. Because egg production was dependent on the amount of R. lens consumed regardless of the amount of K. brevis eaten, there was no evidence that fitness costs were caused by K. brevis toxicity. Copepods limited to K. brevis ate 480% as much as those fed only R. lens, suggesting that copepods attempted to compensate for low food quality with increased quantity ingested. Our results indicate that K. brevis is a poor food for A. tonsa, probably due to nutritional inadequacy rather than toxicity, which could affect bloom dynamics in the Gulf of Mexico where these species co-occur.     

Assessing the diversity of New Zealand freshwater harpacticoid copepods (Crustacea: Copepoda) using mitochondrial DNA (COI) barcodes

Taxonomic and ecological studies of freshwater harpacticoid copepods are limited globally by the ability to easily and accurately identify specimens. Here, we test the use of the mitochondrial cytochrome c oxidase subunit I (COI) gene locus as a tool for assessing the diversity of freshwater Harpacticoida. We obtained sequences from New Zealand harpacticoid copepods, representing two families, five genera and nine species, including the non-indigenous Elaphoidella sewelli. All species were delineated by the COI gene. However, high intraspecific diversity was evident among populations of Elaphoidella bidens (>12%), and between North and South Island populations of Bryocamptus pygmaeus (>18%), potentially indicating the presence of morphologically cryptic taxa. We suggest that mitochondrial DNA (COI) sequences can provide a useful tool for the routine identification of freshwater harpacticoid copepods. Applications of these data will include assessing species diversity and biogeography as well as assisting with the detection of non-indigenous species.     

The formation and degradation of cyanobacterium Aphanizomenon flos-aquae blooms: the importance of pH, water temperature, and day length

Investigation of annual changes in phytoplankton community structure in a small artificial eutrophic pond was carried out from May 2002 to April 2003. A heavy bloom of Aphanizomenon flos-aquae var. klebahnii Elenk. (Cyanobacteria) persisted in most of the water column from June to the end of October. In November, the A. flos-aquae bloom suddenly crashed and green algae were predominant until the end of spring. Weekly monitoring suggested strong involvement of the changes in abiotic factors in the cyanobacterial bloom degradation. To clarify the effects of pH, water temperature, and day length on the growth of A. flos-aquae, laboratory batch experiments were conducted. The results showed that A. flos-aquae could not grow below pH 7.1 and 11°C, and the growth tended to be suppressed under a 10L:14D photoperiod. pH, water temperature, and day length are vital factors in the growth of A. flos-aquae and, additionally, grazing by cyclopoid copepods also seemed important in bloom collapse.     

Seasonal variations in copepod body length: a comparison between different species in the Lagoon of Venice

The seasonal variation in length of the most abundant copepod species in the lagoon of Venice were compared and the relative influence of temperature, chlorophyll concentration and salinity examined. Temperature seems to be the primary factor influencing copepod body length in the lagoon of Venice, but the different species vary markedly in their response. Calanoid copepods (Acartia clausi, A. tonsa, Paracalanus parvus and Centropages ponticus) showed a more definite trend of size variation with temperature than the cyclopoids (Oithona nana and O. similis) and the harpacticoid Euterpina acutifrons. The size of the poecilostomatoids Oncaea media and O. subtilis was not affected by temperature, and was almost constant over time. Differences in the metabolic rates, longevity and specialization of calanoid, cyclopoid and harpacticoid copepods could explain their different responses to temperature.     

Lacustrine profundal meiobenthos as an environmental indicator

Organic loading and eutrophy is indicated at profundal depths by large numbers of resting copepodid stages of cyclopoid copepods, by the occurrence of the naidid oligochaete species, Amphichaeta leydigii and Specaria josinae, and the harpacticoid species Canthocamptus staphylinus, and by a low meiobenthos/macrobenthos biomass ratio. An oligotrophic environment is indicated by the occurrence of the aeolosomatid oligochaetes Aeolosoma quaternarium, A. hemprichi and Rheomorpha neiswestnovae, the naidid oligochaete Chaetogaster langi and the harpacticoid species Attheyella crassa and a high meiobenthos/macrobenthos biomass ratio.     

Changes in the abundance and composition of cyclopoid copepods following fish manipulation in eutrophic Lake Væng, Denmark

1. The seasonal cycle of cyclopoid copepods during and following an approximately 50% reduction in planktivorous fish biomass was studied in shallow, eutrophic Lake Vaeng, Denmark, from 1986 to 1990. 2. The dominant cyclopoid copepods changed from Cyclops vicinus and Mesacyclops leuckarti during 1986–1989 to M. leuckarti and Megacyclops viridis in 1990. The abundance of cyclopoid copepods gradually increased from 1986 to 1988–89, decreased in autumn 1989 and markedly decreased in 1990. 3. The increase in the abundance of cyclopoid copepods from 1986 to 1988 is attributed mainly to the reduction in fish predation pressure, there being no concomitant increase in edible phytoplankton. The appearance of M. viridis in 1990, and the general decrease in cyclopoid copepod density in autumn 1989 and in 1990, are attributed to the appearance of submerged macrophytes. 4. Temperature, predation and availability of edible phytoplankton appear to determine whether C. vicinus or M. leuckarti dominates the cyclopoid copepod population of eutrophic Lake Væng.     

Diel and seasonal vertical distribution of meiobenthic copepods in muddy sediments of a eutrophic lagoon (fish ponds of Arcachon Bay)

The vertical distribution of meiobenthic copepods was investigated within muddy sediments of a eutrophic lagoon (fish ponds of Arcachon Bay, France). The aim of the study was to determine if in muddy sediments, as previously established in sandy sediments, meiobenthic copepods migrate vertically according to the seasons or diel periods. Two experimental approaches were used, viz: a three-season comparison was made of the diel vertical distribution of the harpacticoid Canuella perplexa T. & A. Scott (1893) and secondly the depth distribution of a meiobenthic copepod assemblage was followed for a 24 h period, in shallow water subtidal locations. The harpacticoid C. perplexa vertically migrated through the top three centimeters of the sediment, showing diel and seasonal variations in depth distribution. The differential vertical distributions shown by the dominant meiobenthic populations suggest that emergence into the water column may mainly concern surface dwelling copepods. The physical and biological factors affecting seasonal and diel changes in the copepod assemblage of the fish ponds are discussed.     

Trichodesmium thiebautii; structure of a nitrogen-fixing marine blue-green alga (Cyanophyta)

Summary Trichodesmium thiebautii was collected, as floating bundles composed of uniseriate filaments aligned in parallel, from the Kuroshio waters off Shikoku Island, Japan. The ultrastructure of this alga had basically the same general features as the related speciesT. erythraeum first described byvan Baalen andBrown (1969). InT. thiebautii long electron dense fibers and concentrically lamellated bodies were observed which were either not reported previously, or did not occur inT. erythraeum. The peripheral wall layers were generally typical ofOscillatoria-type blue-green algae, but with a distinctive finely striated outer layer. Thylakoids per cell volume were very sparse compared to most other blue-green algae. Phycobilisomes, apparently hemidiscoidal in shape, typically occurred on the stromal side of the thylakoid surface. Large gas vesicle areas occupied the main volume of the cell, including cells which seemed to be actively growing. The gas vesicle areas were distributed throughout the cell, not only in the cell periphery as inT. erythraeum. Considerable complexity was suggested by the apparent cell compartmentation, particularly because the gas vesicle areas were delimited by one to several thylakoids. Only rarely were the gas vesicle areas traversed by thylakoids. Electron dense fibers (ca. 25 nm diameter) were always observed between the gas vesicles and were usually oriented parallel with them, but they were not rigid appearing as were the gas vesicles. The gas vesicles had a smaller diameter (ca. 45 nm) than most blue-greens. Concentrically lamellated bodies (ca. 1.0 m diameter) were observed in cells of some of the bundles. Each concentric layer was ca. 1.3 nm wide. These concentrically lamellated bodies may be characteristic of older cells. Cylindrical bodies were considerably smaller (ca. 120 nm diameter) and less complex than those reported forT. erythraeum.     

Ultrastructural characterisation of cells specialised for nitrogen fixation in a non-heterocystous cyanobacterium,Trichodesmium spp.

Summary Trichodesmium is the first described example of a filamentous cyanobacterium without heterocysts that contains cells specialised for nitrogen fixation. The ultrastructure of cells with and without nitrogenase were compared using primarilyTrichodesmium tenue Wille, but alsoT. thiebautii Gomont andT. erythraeum Ehrenberg et Gomont. Immunohistochemistry demonstrated that the cytoplasm of certain cells was densely labelled with antibodies against Fe-protein (dinitrogenase reductase). Comparative TEM-image analysis revealed that these cells were also distinguished by a denser thylakoid network, dividing the vacuole-like space into smaller units. The nitrogenase-containing cells also exhibited less extensive gas vacuoles as well as fewer and smaller cyanophycin granules compared to cells which lacked nitrogenase. Carboxysomes were present in both cell types in equal proportion. Longitudinal sections showed that cells with nitrogenase were arranged adjacent to each other, and that groups of cells with and without nitrogenase may coexist in the same trichome. The correlation between modifications in ultrastructure and the presence of nitrogenase suggests a new type of cyanobacterial cell specialisation related to nitrogen fixation. The results obtained also question the systematic affiliation of the genusTrichodesmium.     

Chemical Response of the Toxic Dinoflagellate Karenia mikimotoi Against Grazing by Three Species of Zooplankton

We investigated the toxicity of Karenia mikimotoi toward three model grazers, the cladoceran Moina mongolica, the copepod Pseudodiaptomus annandalei, and the crustacean Artemia salina, and explored its chemical response upon zooplankton grazing. An induction experiment, where K. mikimotoi was exposed to grazers or waterborne cues from the mixed cultures revealed that K. mikimotoi might be toxic or nutritionally inadequate toward the three grazers. In general, direct exposure to the three grazers induced the production of hemolytic toxins and the synthesis of eicosapentaenoic acid (EPA). Both EPA and the hemolytic toxins from K. mikimotoi decreased the survival rate of the three grazers. In addition, the survival rates of M. mongolica, P. annandalei, and A. salina in the presence of induced K. mikimotoi that had previously been exposed to a certain grazer were lower than their counterparts caused by fresh K. mikimotoi, suggesting that exposure to some grazers might increase the toxicity of K. mikimotoi. The chemical response and associated increased resistance to further grazing suggested that K. mikimotoi could produce deterrents to protect against grazing by zooplankton and that the substances responsible might be hemolytic toxins and EPA.     

Top‐down regulation of filamentous cyanobacteria varies among a raptorial versus current feeding copepod across multiple prey generations

1. Although considered a key functional trait, little is known about how zooplankton feeding mode affects top‐down regulation of phytoplankton communities. Indeed, copepods are expected to promote the dominance of toxic phytoplankton by selective removal of their edible competitors; however, empirical evidence comparing the effect among calanoid and cyclopoid copepods is lacking.
2. We compared the top‐down effects of two copepods with contrasting feeding modes—the calanoid Notodiaptomus iheringi (current feeder) and the cyclopoid Thermocyclops decipiens (ambush feeder) — on the relative and absolute biomass of the filamentous cyanobacterium Raphidiopsis raciborskii co‐cultured with the nutritious eukaryotic phytoplankton Cryptomonas obovata in a week‐long laboratory assay.
3. The current feeder had a stronger top‐down effect on the biomass of both prey throughout the experiment, with mass‐specific clearance rates 3–5× higher than ambush feeder. By the end of the experiment, the current feeder significantly reduced cyanobacteria biomass compared to controls while the ambush feeder did not. During the week‐long experiment, the current feeder switched from grazing on edible prey to cyanobacteria as the former became less abundant.
4. Contrary to expectation, neither of the copepod species promoted cyanobacterial dominance by the end of the experiment. This is because both grazers, but especially the current feeder, initially increased but subsequently decreased the relative contribution of cyanobacteria to total phytoplankton biomass. Moreover, both copepods decreased the length of cyanobacteria filaments by c. 70%
5. Current feeders can switch from edible prey to cyanobacteria when the abundance of shortened filaments surpasses the abundance of edible prey. While top‐down regulation of phytoplankton can be stronger for current feeding copepods, ambush feeding copepods can have a significant role during blooms by shortening cyanobacterial filaments. Hence, the broader role of contrasting copepod feeding traits on phytoplankton communities merits further study.

     

Response of algae and zooplankton to C18 fatty acids of Chlamydomonas reinhardtii

Three C₁₈ fatty acids were assayed for their activity against a number of algae and zooplankton. The three acids, lenolenic, lenoleic, and oleic, reduced the growth of Haematococcus lacustris, Synechococcus leopoliensis, and Botrydiopsis alpina by 50% of control growth in concentrations below 7 ppm. Calanoid and cyclopoid copepods in mixed cultures inoculated with lenolenic and lenoleic acid had LD₅₀ values below 10 ppm. An increase in copepod mortality was observed with increases in cyclopoid density and decreased with increases in calanoid density. Eucyclops agilis inoculated with lenolenic acid had a LD₅₀ value of 4 ppm.     

Feeding,egg production,and egg hatching success of the copepods Acartia tonsa and Temora longicornis on diets of the toxic diatom Pseudo-nitzschia multiseries and the non-toxic diatom Pseudo-nitzschia pungens

In 1987, there was an episode of shellfish poisoning in Canada with human fatalities caused by the diatom Pseudo-nitzschia multiseries, which produced the toxin domoic acid. In order to examine whether domoic acid in this diatom serves as a grazing deterrent for copepods, we compared feeding rates, egg production rates, egg hatching success and mortality of the calanoid copepods Acartia tonsa and Temora longicornis feeding on unialgal diets of the toxic diatom P. multiseries and the similarly-sized non-toxic diatom Pseudo-nitzschia pungens. Copepods were collected in summers of 1994, 1995 and 1996 from Shediac Bay, New Brunswick, Canada, near Prince Edward Island, the site of the 1987 episode of domoic acid shellfish poisoning. Rates of ingestion of the toxic versus the non-toxic diatom by A. tonsa and T. longicornis were similar, with only one significantly different pair of values obtained in 1994, for which A. tonsa had a higher mean rate of ingestion of the toxic than the non-toxic diatom. Thus, domoic acid did not appear to retard grazing. Analyses of copepods with high performance liquid chromatography (HPLC) revealed that copepods accumulated domoic acid when feeding on P. multiseries. Egg production rates of copepods when feeding on P. multiseries and P. pungens were very low, ranging from 0 to 2.79 eggs female^–1 d^–1. There did not appear to be differential egg production or egg hatching success on diets of the toxic and non-toxic diatoms. Mortality of females on the toxic diet was low, ranging from 0 to 20%, with a mean of 13%, and there was no apparent difference between mortality of copepods feeding on toxic versus non-toxic diatoms. Egg hatching success on both diets, although based on few eggs, ranged between 22% and 76%, with a mean percentage hatching of 45%. Diets of the non-toxic diatom plus natural seawater assemblages supplemented with dissolved domoic acid, revealed similar rates and percentages when compared to previous experiments. In summary, none of the variables measured indicated adverse effects on copepods feeding on the toxic compared to the non-toxic diatom.     

Feeding habits of juvenile <Emphasis Type="Italic">Trematomus newnesi</Emphasis> (Pisces,Nototheniidae) at Potter Cove,South Shetland Islands,Antarctica

The dietary composition of juvenile Trematomus newnesi, trawled at Potter Cove, South Shetland Islands, in the summers 2004–2005 and 2005–2006 was analysed using frequency of occurrence (F%) and dietary coefficient Q (%) methods. The samples consisted exclusively of immature specimens in the range 4–15.4 cm (total length) and are complementary to those of larger fish from the same site, including adults, which were analysed in previous work. Benthic-demersal organisms such as gammaridean amphipods and harpacticoid copepods were the main (coefficient Q) and most frequent (F%) prey. The importance of the smaller and larger main prey diminished and increased, respectively, during ontogeny. Pelagic krill, being negligible in the diet of the small and medium size fish categories, became secondary food, but only for fish larger than 12 cm. Other taxonomic groups occurred scarcely and constituted occasional food. They were mostly benthic, such as gastropods, bivalves, isopods, cumaceans, and algae, with the exception of an insignificant number of pelagic ostracods and calanoid copepods. Unlike the more pelagic/planktivorous mode of life known for late juvenile-adult stages of T. newnesi, including cryopelagy, present results indicate that early juvenile fish remain sheltered among macroalgae beds preying on the associated community of demersal-benthic organisms.     

Copepod grazing during an iron-induced diatom bloom in the Antarctic Circumpolar Current (EisenEx): I. Feeding patterns and grazing impact on prey populations

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics.     

Is diatom size selection by harpacticoid copepods related to grazer body size?

Copepods are known as important consumers of primary production and are eaten by larger animals. They therefore form a main link to higher trophic levels. While feeding pathways and specificity of planktonic copepods have been well studied, the selectivity of the benthic harpacticoid copepods is far less documented. A better knowledge of the functional ecology of harpacticoids as important grazers on primary producers may have consequences for the re-evaluation of basic energy flow in benthic ecosystems.We tested whether size selectivity for diatoms exists in harpacticoid copepods. We hypothesized that size selectivity of harpacticoid copepod species is strongly related to body size. Because of morphological constraints, we expected smaller copepods to prefer smaller diatoms while larger copepods should be able to consume both small and large diatoms. We tested this hypothesis in four harpacticoid copepod species of varied body size: Tigriopus brevicornis, Harpacticus obscurus, Amphiascus minutus and Paramphiascella fulvofasciata. As food source we used two ¹³C labelled strains of the benthic diatom Seminavis robusta with a four-fold difference in cell biovolume.Three out of four harpacticoid species showed size selectivity: H. obscurus and A. minutus preferred the larger Seminavis cells, while P. fulvofasciata selected the smaller Seminavis cells. Based on monoclonal treatments, there was no clear preference found for T. brevicornis although there was a small preference for large cells in the mixed treatments. Except for P. fulvofasciata, all species showed a lower uptake when offered the mixed diet (both small and large cells). Although most species showed a size selectivity, our results suggest that this selectivity was not related to their body size. However, the only species that ate significantly more of small diatoms was characterised by comparatively small mouthparts in relation to its body size.     

Differential responses of populations of the copepod Acartia hudsonica to toxic and nutritionally insufficient food algae

Nutritional insufficiency and toxicity are deleterious effects of phytoplankton on grazers. We hypothesize that toxic food is likely to have stronger evolutionary selective effects on grazers than nutritionally insufficient food. We explore this hypothesis in comparative studies of egg production and egg hatching of the copepod Acartia hudsonica challenged with both a toxic and a nutritionally insufficient alga. Experiments lasting 6 days, in which mixtures of different proportions of the suspect and a control alga were offered as food to female copepods, showed that the dinoflagellate Alexandrium fundyense, which bears paralytic shellfish toxins, was toxic to A. hudsonica. In contrast, the diatom Phaeodactylum tricornutum was nutritionally insufficient to A. hudsonica. In another set of experiments, the effects of A. fundyense and P. tricornutum, respectively, as sole foods on egg production and egg hatching success of two geographically separated populations (Maine and Connecticut) of the copepod A. hudsonica were examined in common-environment experiments, after being raised under identical conditions for two generations. The location in Maine regularly experiences toxic blooms of Alexandrium sp. whereas the location in Connecticut does not. During a 6-day period, A. fundyense reduced the egg production rates of the Connecticut copepod population, but not of the Maine population. In contrast, the diatom P. tricornutum reduced the egg production of both populations. These results of this study are consistent with the hypothesis of local adaptation to toxic food, but not to nutritionally insufficient food.