Gyrodinium undulans Hulburt, a marine dinoflagellate feeding on the bloom-forming diatomOdontella aurita, and on copepod and rotifer eggs

The marine dinoflagellateGyrodinium undulans was discovered as a feeder on the planktonic diatomOdontella aurita. Every year, during winter and early spring, a certain percentage of cells of this bloom-forming diatom, in the Wadden Sea along the North Sea coast, was regularly found affected by the flagellate. Supplied with the food diatomO. aurita the dinoflagellate could be maintained successfully in clonal culture. The vegetative life cylce was studied, mainly by light microscopy on live material, with special regard to the mode of food uptake. Food is taken up by a so-called phagopod, emerging from the antapex of the flagellate. Only fluid or tiny prey material could be transported through the phagopod. Larger organelles like the chloroplasts ofOdontella are not ingested and are left behind in the diatom cell. Thereafter, the detached dinoflagellate reproduces by cell division, occasionally followed by a second division. As yet, stages of sexual reproduction and possible formation of resting cysts could not be recognized, neither from wild material nor from laboratory cultures. Palmelloid stages (sometimes with a delicate wall) occurring in ageing cultures may at least partly function as temporary resting stages. The winter speciesG. undulans strongly resemblesSyltodinium listii, a summer species feeding on copepod and rotifer eggs. Surprisingly, in a few cases this prey material was accepted byG. undulans as well, at least under culture conditions. When fed with copepod eggs, the dinoflagellate developed into a large trophont, giving rise thereafter by repeated binary fission to 4, 8 or 16 flagellates, as a result of a single feeding act. A re-examination of both species under simultaneous culture conditions is planned.     

PHAGOTROPHIC FEEDING AND ITS IMPORTANCE TO THE LIFE CYCLE OF THE HOLOZOIC DINOFLAGELLATE,GYMNODINIUM FUNGIFORME1

The holozoic dinoflagellate, Gymnodinium fungiforme Anissimova, has been observed in both asexually and sexually reproducing cultures. Asexual reproduction is characterized by zoosporangium formation and subsequent new cell release. Sexuality is gametic, and planozygotes and hypnozygotes are present. The life cycle is highly dependent on feeding, and in food-depleted cultures the swimming cells rapidly disappear. These are replaced with resistant long-term resting cysts. Despite its small size (8.5–19 μm), G. fungiforme can feed on prey as large as the ciliated protozoan, Condylostoma magnum Spiegel (600–1000 μm in length), or small injured metazoans, and has been cultured phagotrophically with the chlorophyte, Dunaliella salina Teodoresco as a food source. Eleven additional species of algae including 1 chlorophyte, 7 chrysophytes and 3 rhodophytes, however, were not suitable as food sources. Feeding is characterized by the formation of ‘dynamic aggregations’ of hundreds of dinoflagellates that attach to the surface of a prey organism by a peduncle. G. fungiforme ingests the cytoplasm or body fluids of its prey and a feeding aggregation can ingest a C. magnum in 20–30 minutes.     

AMPHIDINIUM CRYOPHILUM SP. NOV. (DINOPHYCEAE) A NEW FRESHWATER DINOFLAGELLATE. II. ULTRASTRUCTURE1

The dinoflagellate Amphidinium cryophilum sp. nov. is one of the few gymnodinians to be studied at the ultrastructural level. It resembles other dinoflagellates in the structure of the nucleus, trichocysts, storage materials, flagella, mitochondria, and microbodies. Other features of A. cryophilum less commonly observed in related organisms include a network of small interconnected vesicles, a system of large, peripheral vacuoles, chloroplasts bound by two rather than three membranes, an accumulation body, thylakoid-associated plastoglobuli, a vesiculated nuclear envelope, a complex tubular pusule, striated flagellar collars, collared pits, and a peduncle. The occurrence of a peduncle, a structure implicated in phagotrophy, in this autotrophic organism is noteworthy. The ultrastructure of the peduncle of A. cryophilum differs significantly from that reported in another dinoflagellate.     

甲藻的异养营养型

综述了甲藻的异养类型。目前已知异养营养型在甲藻中广泛存在,只有很少几种甲藻营严格自养营养方式。有近一半的甲藻物种是没有色素体的,还有很多甲藻即使具有色素体也会有异养营养需求,称为兼养营养类型。这些兼养类群不一定主要以有机物作为其获取碳的来源,而仅仅是补充一些生长必需的有机物如维生素、生物素等。兼养类群以渗透营养和腐食营养方式进行,同时也可以寄生方式和共生方式进行兼养生活。无色素体的甲藻以有机物作为碳的唯一来源,仅仅依靠异养方式生存,属于严格异养营养方式,又称有机营养型。它们是甲藻异养营养型的主体,其主要类型有寄生、渗透营养和吞噬营养。由于吞噬营养是甲藻异养的主要类型,因此论述了3种吞噬营养型:吞噬营养方式、捕食茎营养方式和捕食笼营养方式。吞噬营养方式在无甲类和具甲类甲藻中都有存在,主要通过甲藻细胞的纵沟或底部对猎物进行吞噬,也有研究发现吞噬部位为顶孔或片间带。捕食茎营养方式是通过捕食茎刺穿猎物细胞膜并吸食其细胞质来获取营养,在异养甲藻中也较常见。捕食笼营养方式只在原多甲藻属(Protoperidinium)和翼藻属(Diplopsalis)里发现,是甲藻通过鞭毛孔分泌细胞质到胞外形成捕食笼将猎物包裹并进行消化来摄食的。甲藻摄食对象尺寸范围变化较大,小至几微米,大至几百微米。有些甲藻具有摄食选择性,通过感应猎物释放的化学物质来判断猎物的位置并进行摄食,摄食完成后由于体积的增加经常会发生细胞分裂和蜕鞘。对于甲藻异养的其他形式如拦截摄食营养方式、伪足摄食营养方式、口足摄食营养方式、触手摄食营养方式等只作简单介绍。还就甲藻异养的研究方法、其生态学意义和进化学意义进行简要论述,并对相关研究进行展望。     

Phagotrophic Mechanisms and Prey Selection in Free-living Dinoflagellates1

Three types of feeding mechanisms are known in dinoflagellates: pallium feeding, tube feeding, and direct engulfment. Pallium feeding has only been described for heterotrophic thecate species (Protoperidinium, Diplopsalis group). Tube feeding is commonly found among both naked and thecate species of mixotrophic and heterotrophic dinoflagellates (e.g. Amphidinium, Dinophysis, Gyrodinium, Peridiniopsis). Direct engulfment is mainly found among naked species (e.g. Gymnodinium, Gyrodinium, Noctiluca): recently, however, some thecate species have been shown to use this feeding mechanism as well. Feeding behavior in dinoflagellates involves several steps prior to actual ingestion, including precapture, capture, and prey manipulation. As feeding mechanisms allow the ingestion of relatively large prey or parts thereof, dinoflagellates are regarded as raptorial feeders. While prey size plays an important role in the ability of dinoflagellates to ingest food, this alone cannot explain observed prey preferences. Some dinoflagellate species can be very selective in their choice of prey, while others show a remarkable versatility.     

AMPHIDINIUM CRYOPHILUM SP. NOV. (DINOPHYCEAE) A NEW FRESHWATER DINOFLAGELLATE. I. SPECIES DESCRIPTION USING LIGHT AND SCANNING ELECTRON MICROSCOPY1

Amphidinium cryophilum sp. nov. was found in the fall of 1979 in a small pond near Madison, Wisconsin. During the ensuing winter, it became the dominant phytoplankter. Cell numbers remained high despite a thick layer of ice and snow. After the ice melted in the spring the organism disappeared from plankton samples. A successful culture of A. cryophilum was established only when isolates were incubated at 5–7° C. It is compared with two morphologically similar species, A. amphidinioides (Geitler) Schiller and Gymnodinium inversum Nygaard. Amphidinium cryophilum is distinguished from the former by its pigmentation (golden-yellow vs. blue-green), the location of the cingulum, and its lack of an eyespot. It differs from the latter in cell shape, the route of the sulcus and position of the nucleus.     

Mixotrophy in the sand-dwelling dinoflagellate Thecadinium kofoidii

Thecadinium kofoidii is a marine sand-dwelling dinoflagellate that sometimes forms dense blooms. This species was previously thought to be an exclusively autotrophic dinoflagellate, and its mixotrophic ability has not been explored yet. By investigating its ecophysiology, its trophic mode should be revealed. We explored the mixotrophic ability of T. kofoidii by examining its protoplasm under light and transmission electron microscopes with diverse algal prey species. Furthermore, the feeding mechanism of T. kofoidii and prey species on which it feeds were investigated. In addition, the growth and ingestion rates of T. kofoidii as a function of prey concentration were determined when feeding on the benthic cryptophyte Rhodomonas salina. Thecadinium kofoidii was able to feed on R. salina and the dinoflagellate Symbiodinium voratum, which had equivalent spherical diameters (ESDs) ≤ 10.1?µm, while it did not feed on the benthic dinoflagellates Levanderina fissa, Prorocentrum concavum or Ostreopsis cf. ovata, which had ESDs ≥ 15?µm. Thecadinium kofoidii fed on the edible prey cells using the peduncle. The maximum ingestion rate of T. kofoidii on R. salina was 1.3 cells predator^?1 d^?1. However, feeding on R. salina did not significantly increase the growth rate of T. kofoidii. The low ingestion rate of T. kofoidii on R. salina may have partially resulted in the lack of significant increase in its growth rate due to mixotrophy. The present study discovered predator–prey relationships between T. kofoidii and R. salina and S. voratum, which may change our view of the energy flow and carbon cycling in marine benthic food webs.     

Phosphatase activity in the heterotrophic dinoflagellate Pfiesteria shumwayae

The ELF-97 phosphatase substrate was used to examine phosphatase activity in four strains of the estuarine heterotrophic dinoflagellate, Pfiesteria shumwayae. Acid and alkaline phosphatase activities also were evaluated at different pH values using bulk colorimetric methods. Intracellular phosphatase activity was demonstrated in P. shumwayae cells that were actively feeding on a fish cell line and in food limited cells that had not fed on fish cells for 3 days. All strains, whether actively feeding or food limited showed similar phosphatase activities. P. shumwayae cells feeding on fish cells showed ELF-97 activity near, or surrounding, the food vacuole. Relatively small, spherical ELF-97 deposits were also observed in the cytoplasm and sometimes near the plasma membrane. ELF-97 fluorescence was highly variable among cells, likely reflecting different stages in digestion and related metabolic processes. The location of enzyme activity and supporting colorimetric measurements suggest that, as in other heterotrophic protists, acid phosphatases predominate in P. shumwayae and have a general catabolic function.     

Feeding by the Newly Described Mixotrophic Dinoflagellate Paragymnodinium shiwhaense: Feeding Mechanism,Prey Species,and Effect of Prey Concentration

ABSTRACT. To investigate the feeding by the newly described mixotrophic dinoflagellate Paragymnodinium shiwhaense (GenBank accession number=AM408889), we explored the feeding process and the kinds of prey species that P. shiwhaense is able to feed on using several different types of microscopes, including a transmission electron microscope and high‐resolution video‐microscopy. In addition, we measured the growth and ingestion rates of P. shiwhaense on its optimal algal prey Amphidinium carterae as a function of prey concentration. We also measured these parameters for edible prey at a single concentration at which the growth and ingestion rates of P. shiwhaense on A. carterae were saturated. Paragymnodinium shiwhaense feed on algal prey using a peduncle after anchoring the prey by a tow filament. Among the algal prey offered, P. shiwhaense ingested small algal species that had equivalent spherical diameters (ESDs) ≤11 μm (e.g. the prymnesiophyte Isochrysis galbana, the cryptophytes Teleaulax sp. and Rhodomonas salina, the raphidophyte Heterosigma akashiwo, and the dinoflagellates Heterocapsa rotundata and A. carterae). However, it did not feed on larger algal species that had ESDs ≥12 μm (e.g. the dinoflagellates Prorocentrum minimum, Heterocapsa triquetra, Scrippsiella trochoidea, Alexandrium tamarense, Prorocentrum micans, Gymnodinium catenatum, Akashiwo sanguinea, and Lingulodinium polyedrum) or the small diatom Skeletonema costatum. The specific growth rates for P. shiwhaense feeding upon A. carterae increased rapidly with increasing mean prey concentration before saturating at concentrations of ca. 350 ng C/ml (5,000 cells/ml). The maximum specific growth rate (i.e. mixotrophic growth) of P. shiwhaense on A. carterae was 1.097/d at 20 °C under a 14:10 h light–dark cycle of 20 μE/m²/s, while its growth rate (i.e. phototrophic growth) under the same light conditions without added prey was ?0.224/d. The maximum ingestion and clearance rates of P. shiwhaense on A. carterae were 0.38 ng C/grazer/d (5.4 cells/grazer/d) and 0.7 μl/grazer/h, respectively. The calculated grazing coefficients for P. shiwhaense on co‐occurring Amphidinium spp. was up to 0.07/h (i.e. 6.7% of the population of Amphidinium spp. was removed by P. shiwhaense populations in 1 h). The results of the present study suggest that P. shiwhaense can have a considerable grazing impact on algal populations.     

WIDESPREAD PHAGOCYTOSIS OF CILIATES AND OTHER PROTISTS BY MARINE MIXOTROPHIC AND HETEROTROPHIC THECATE DINOFLAGELLATES1

An electron microscopic examination of large amorphous inclusions located in a variety of photosynthetic thecate dinoflagellates (Alexandrium ostenfeldii (Paulsen) Balech et Tangen, Gonyaulax diegensis Kofoid, Scrippsiella sp., Ceratium longipes (Bailey) Gran, and Prorocentrum micans Ehrenberg) and a nonphotosynthetic thecate species (Amylax sp.) revealed each inclusion to be a food vacuole, the majority of which were ingested ciliate prey. Recognizable features of these ciliates included linear arrays of basal bodies and cilia consistent with oligotrich polykinetid structure, characteristic macronuclei, chloroplasts (evidently kleptoplastids), cup-shaped starch plates, and cylindrical extrusomes. Three species contained (apparent) nonciliate prey: Scrippsiella sp., whose food vacuoles consistently contained unusual and complex extrusome-like cylindrical bodies having a distinctive six-lobed, multilayered structure; P. micans, which contained an unidentified encysted cell; and a single A. ostenfeldii cell, containing a Dinophysis sp. dinoflagellate cell. Several food vacuoles of ciliate origin had a red hue. This, together with the resemblance of A. ostenfeldii cells to planozygotes, suggests that similar structures previously identified as accumulation bodies may in fact be food vacuoles and that feeding may in some cases be associated with sexual processes.     

Feeding by the newly described, nematocyst-bearing heterotrophic dinoflagellate Gyrodiniellum shiwhaense

We explored the feeding ecology of the newly described, nematocyst-bearing heterotrophic dinoflagellate Gyrodiniellum shiwhaense (GenBank accession number=FR720082). Using several different types of microscopes and high-resolution video-microscopy, we investigated feeding behavior and types of prey species that G. shiwhaense feeds upon. Additionally, we measured its growth and ingestion rates on its optimal algal prey, the cryptophyte Teleaulax sp. and the dinoflagellate Amphidinium carterae, as a function of prey concentration. These rates were measured for other edible prey at single prey concentrations at which the growth and ingestion rates of G. shiwhaense were saturated. After anchoring the prey with a tow filament, G. shiwhaense fed using a peduncle, ingesting small algal species with equivalent spherical diameters (ESDs) of <13 μm. However, it did not feed on larger algal species that had ESDs≥13 μm or the small diatom Skeletonema costatum. The specific growth rates for G. shiwhaense feeding upon Teleaulax sp. and A. carterae increased rapidly with increasing mean prey concentration before saturating at concentrations of ca. 180-430 ng C/ml. The maximum specific growth rate of G. shiwhaense on Teleaulax sp. and A. carterae were 1.05 and 0.82/d, respectively. However, Heterosigma akashiwo did not support positive growth of G. shiwhaense. The maximum ingestion rates of G. shiwhaense on Teleaulax sp. and A. carterae were 0.35 and 0.54 ng C/grazer/d, respectively. The calculated grazing coefficients attributable to G. shiwhaense on co-occurring cryptophytes and Amphidinium spp. were 0.01-1.87/d and 0.08-2.60/d, respectively. Our results suggest that G. shiwhaense can have a considerable grazing impact on algal populations.     

COMPARATIVE ANALYSIS OF THE DINOFLAGELLATE FLAGELLAR APPARATUS IV. GYMNODINIUM ACIDOTUM1

Gymnodinium acidotum Nygaard is a freshwater dinoflagellate that is known to harbor a cryptomonad endosymbiont whose chloroplasls give the organism an overall blue-green color. The ultrastructure of G. acidotum was examined with particular attention being given to the three dimensional nature of the flagellar apparatus. The fiagellar apparatus is composed of two functional basal bodies that are slightly offset and lie at an angle of approximately 90° to one another. As in other dinoflagellates the transverse basal body is associated with a striated, fibrous root that extends from the proximal end of the basal body to the transverse flagellar opening. At least one microtubular root extends from the proximal end of the transverse basal body, and a multi-membered longitudinal microtubular root is associated with the longitudinal basal body. The most striking feature of the flagellar apparatus of G. acidotum is the large fibrous connective that extends from the region of the proximal ends of the basal bodies to the cingulum on the dorsal side of the cell. A similar structure has been reported from only one other dinoflagellate, Amphidinium cryophilum Wedemayer, Wilcox, and Graham. The presence of this structure as well as similarities in external morphology suggest thai these two species may be more closely related to each other than either is to other gymnodinioid taxa. The taxonomic importance of dinoflagellate flagellar apparatus components is discussed.     

Intraspecific variability in Karlodinium veneficum: Growth rates, mixotrophy, and lipid composition

We isolated eleven strains of the harmful algal bloom (HAB)-forming dinoflagellate Karlodinium veneficum during a bloom event in the NW Mediterranean coastal waters and we studied the inter-strain variability in several of their physiological and biochemical traits. These included autotrophic growth parameters, feeding capabilities (mixotrophy), lipid composition, and, in some cases, their responses to biotic and abiotic factors. The strains were found to differ in their growth rates (0.27–0.53 d⁻¹) and in the maximum cell concentrations achieved during stationary phase (6.1 × 10⁴–8.6 × 10⁴ cells mL⁻¹). Their ingestion performance, when offered Rhodomonas salina as prey, was also diverse (0.22–1.3 cells per K. veneficum per day; 8–52% of their daily ration). At least two strains survived for several months under strict heterotrophic conditions (no light, low inorganic nutrients availability, and R. salina as food source). These strains also showed very distinct fatty acid compositions, with very low contents of monounsaturated and polyunsaturated fatty acids. According to a Bray Curtis similarity analysis, three or four strain groups able to perform different roles in bloom development were identified. We further analyzed one strain from each of the two most distinct groups with respect to prey concentration, light intensity, nutrient availability, and we determined the functional responses (growth and feeding rates) to food concentration. Taken together, the results served to highlight the role of mixotrophy and clone variability in the formation of HABs.     

Copepod Foraging on the Basis of Food Nutritional Quality: Can Copepods Really Choose?

Copepods have been considered capable of selective feeding based on several factors (i.e., prey size, toxicity, and motility). However, their selective feeding behaviour as a function of food quality remains poorly understood, despite the potential impact of such a process on copepod fitness and trophodynamics. In this study, we aimed to evaluate the ability of copepods to feed selectively according to the nutritional value of the prey. We investigated the feeding performance of the calanoid copepod Acartia grani under nutritionally distinct diets of the dinoflagellate Heterocapsa sp. (nutrient-replete, N-depleted and P-depleted) using unialgal suspensions and mixtures of prey (nutrient-replete vs. nutrient-depleted). Despite the distinct cell elemental composition among algal treatments (e.g., C:N:P molar ratios) and the clear dietary impact on egg production rates (generally higher number of eggs under a nutrient-replete diet), no impact on copepod feeding rates was observed. All unialgal suspensions were cleared at similar rates, and this pattern was independent of food concentration. When the prey were offered as mixtures, we did not detect selective behaviour in either the N-limitation (nutrient-replete vs. N-depleted Heterocapsa cells) or P-limitation (nutrient-replete vs. P-depleted Heterocapsa cells) experiments. The lack of selectivity observed in the current study contrasts with previous observations, in which stronger nutritional differences were tested. Under normal natural circumstances, nutritional differences in natural prey assemblages might not be sufficiently strong to trigger a selective response in copepods based on that factor alone. In addition, our results suggest that nutritional quality might depend not only on the growing conditions but also on the inherent taxonomical properties of the prey.     

Feeding by Phototrophic Red-Tide Dinoflagellates on the Ubiquitous Marine Diatom Skeletonema costatum

ABSTRACT We investigated feeding by phototrophic red‐tide dinoflagellates on the ubiquitous diatom Skeletonema costatum to explore whether dinoflagellates are able to feed on S. costatum, inside the protoplasm of target dinoflagellate cells observed under compound microscope, confocal microscope, epifluorescence microscope, and transmission electron microscope (TEM) after adding living and fluorescently labeled S. costatum (FLSc). To explore effects of dinoflagellate predator size on ingestion rates of S. costatum, we measured ingestion rates of seven dinoflagellates at a single prey concentration. In addition, we measured ingestion rates of the common phototrophic dinoflagellates Prorocentrum micans and Gonyaulax polygramma on S. costatum as a function of prey concentration. We calculated grazing coefficients by combining field data on abundances of P. micans and G. polygramma on co‐occurring S. costatum with laboratory data on ingestion rates obtained in the present study. All phototrophic dinoflagellate predators tested (i.e. Akashiwo sanguinea, Amphidinium carterae, Alexandrium catenella, Alexandrium tamarense, Cochlodinium polykrikoides, G. polygramma, Gymnodinium catenatum, Gymnodinium impudicum, Heterocapsa rotundata, Heterocapsa triquetra, Lingulodinium polyedrum, Prorocentrum donghaiense, P. micans, Prorocentrum minimum, Prorocentrum triestinum, and Scrippsiella trochoidea) were able to ingest S. costatum. When mean prey concentrations were 170–260 ng C/ml (i.e. 6,500–10,000 cells/ml), the ingestion rates of G. polygramma, H. rotundata, H. triquetra, L. polyedrum, P. donghaiense, P. micans, and P. triestinum on S. costatum (0.007–0.081 ng C/dinoflagellate/d [0.2–3.0 cells/dinoflagellate/d]) were positively correlated with predator size. With increasing mean prey concentration of ca 1–3,440 ng C/ml (40–132,200 cells/ml), the ingestion rates of P. micans and G. polygramma on S. costatum continuously increased. At the given prey concentrations, the maximum ingestion rates of P. micans and G. polygramma on S. costatum (0.344–0.345 ng C/grazer/d; 13 cells/grazer/d) were almost the same. The maximum clearance rates of P. micans and G. polygramma on S. costatum were 0.165 and 0.020 μl/grazer/h, respectively. The calculated grazing coefficients of P. micans and G. polygramma on co‐occurring S. costatum were up to 0.100 and 0.222 h, respectively (i.e. up to 10% and 20% of S. costatum populations were removed by P. micans and G. polygramma populations in 1 h, respectively). Our results suggest that P. micans and G. polygramma sometimes have a considerable grazing impact on populations of S. costatum.     

Trophic Controls on Stage Transformations of a Toxic Ambush-Predator Dinoflagellate1

ABSTRACT. The toxic dinoflagellate, Pfiesteria piscicida, was recently implicated as the causative agent for about 50% of the major fish kills occurring over a three-year period in the Albemarle-Pamlico Estuarine System of the southeastern USA. Transformations between life-history stages of this dinoflagellate are controlled by the availability of fresh fish secretions or fish tissues, and secondarily influenced by the availability of alternate prey including bacteria, algae, microfauna, and mammalian tissues. Toxic zoospores of P. piscicida subdue fish by excreting lethal neurotoxins that narcotize the prey, disrupt its osmoregulatory system, and attack its nervous system. While prey are dying, the zoospores feed upon bits of fish tissue and complete the sexual phase of the dinoflagellate life cycle. Other stages in the complex life cycle of P. piscidia include cryptic forms of filose, rhizopodial, and lobose amoebae that can form within minutes from toxic zoospores, gametes, or planozygotes. These cryptic amoebae feed upon fish carcasses and other prey and, thus far, have proven less vulnerable to microbial predators than flagellated life-history stages. Lobose amoebae that develop from toxic zoospores and planozygotes during colder periods have also shown ambush behavior toward live fish. In the presence of abundant flagellated algal prey, amoeboid stages produce nontoxic zoospores that can become toxic and form gametes when they detect what is presumed to be a threshold level of a stimulatory substance(s) derived from live fish. The diverse amoeboid stages of this fish “ambush-predator” and at least one other Pfiesteria-like species are ubiquitous and abundant in brackish waters along the western Atlantic and Gulf Coasts, indicating a need to re-evaluate the role of dinoflagellates in the microbial food webs of turbid nutrient-enriched estuaries.     

ULTRASTRUCTURE AND LSU RDNA‐BASED PHYLOGENY OF ESOPTRODINIUM GEMMA (DINOPHYCEAE), WITH NOTES ON FEEDING BEHAVIOR AND THE DESCRIPTION OF THE FLAGELLAR BASE AREA OF A PLANOZYGOTE1

A small, freshwater dinoflagellate with an incomplete cingulum, identified as Esoptrodinium gemma Javornický (=Bernardinium bernardinense sensu auctt. non sensu Chodat), was maintained in mixed culture and examined using light and serial section TEM. Vegetative flagellate cells, large cells with two longitudinal flagella (planozygotes), and cysts were examined. The cells displayed a red eyespot near the base of the longitudinal flagellum, made of two or three layers of pigment globules not bounded by a membrane. Yellow‐green, band‐shaped chloroplasts, bounded by three membranes and containing lamella with three thylakoids, were present in both flagellate cells and cysts. Most cells had food vacuoles, containing phagotrophically ingested chlamydomonads or chlorelloid green algae; ingestion occurred through the ventral area, involving a thin pseudopod apparently driven by the peduncle. The pusule was tubular, with numerous diverticula in its distal portion, and opened into the longitudinal flagellar canal. Three roots were associated with each pair of flagellar bases, both in vegetative cells and in a planozygote. The longitudinal microtubular root bifurcated around the longitudinal basal body. The planozygote contained a single peduncle and associated structures, and a single transverse flagellar canal with the two converging transverse flagella. Using two ciliates as outgroup species, phylogenetic analyses based on maximum parsimony, neighbor‐joining and posterior probability (Bayesian analysis) supported a clade comprising Esoptrodinium, Tovellia, and Jadwigia.     

Feeding by the Pfiesteria-like heterotrophic dinoflagellate Luciella masanensis

To explore the feeding ecology of the Pfiesteria-like dinoflagellate (PLD) Luciella masanensis (GenBank Accession no. AM050344, previously Lucy), we investigated the feeding behavior and the kinds of prey species that L. masanensis fed on and determined its growth and ingestion rates of L. masanensis when it fed on the dinoflagellate Amphidinium carterae and an unidentified cryptophyte species (equivalent spherical diam., ESD=5.6 microm), which were the dominant phototrophic species when L. masanensis and similar small heterotrophic dinoflagellates were abundant in Masan Bay, Korea in 2005. Additionally, these parameters were also measured for L. masanensis fed on blood cells of the perch Lateolabrax japonicus and the raphidophyte Heterosigma akashiwo in the laboratory. Luciella masanensis fed on prey cells by using a peduncle after anchoring the prey with tow filament, and was able to feed on diverse prey such as cryptophytes, raphidophytes, diatoms, mixotrophic dinoflagellates, and the blood cells of fish and humans. Among the prey species tested in the present study, perch blood cells were observed to be the optimal prey for L. masanensis. Specific growth rates of L. masanensis feeding on perch blood cells, A. carterae, H. akashiwo, and the cryptophyte, either increased continuously or became saturated with increasing the mean prey concentration. The maximum specific growth rate of L. masanensis feeding on perch blood cells (1.46/day) was much greater than that of A. carterae (0.59/day), the cryptophyte (0.24/day), or H. akashiwo (0.20/day). The maximum ingestion rate of L. masanensis on perch blood cells (2.6 ng C/grazer/day) was also much higher than that of A. carterae (0.32 ng C/grazer/day), the cryptophyte (0.44 ng C/grazer/day), or H. akashiwo (0.16 ng C/grazer/day). The kinds of prey species which L. masanensis is able to feed on were the same as those of Pfiesteria piscicida, but very different from those of another PLD Stoeckeria algicida. However, the maximum growth and ingestion rates of L. masanensis on perch blood cells, A. carterae, H. akashiwo, and the cryptophyte were considerably lower than those of P. piscicida. Therefore, these three dinoflagellates may occupy different ecological niches in marine planktonic communities, even though they have a similar size and shape and the same feeding mechanisms.     

Optimal foraging and ontogeny; food selection by Haplochromis piceatus

Summary We examined the influence of satiation level, prey density and light intensity on food uptake rate through the ontogeny of Haplochromis piceatus. Prey handling in the buccal cavity was found to be the main factor limiting prey uptake rate under light circumstances and at a sufficiently high prey density. Food uptake rate per unit body weight of different sizes of H. piceatus was equal when feeding on Chaoborus but decreased with increasing fish size when feeding on Daphnia magna. In choice experiments with Chaoborus and D. magna, prey selection by H. piceatus of all sizes was according to the predictions based on Charnov's 1976 model.     

Mixotrophy in the phototrophic harmful alga Cochlodinium polykrikoides (Dinophycean): prey species, the effects of prey concentration, and grazing impact

We first reported here that the harmful alga Cochlodinium polykrikoides, which had been previously known as an autotrophic dinoflagellate, was a mixotrophic species. We investigated the kinds of prey species and the effects of the prey concentration on the growth and ingestion rates of C. polykrikoides when feeding on an unidentified cryptophyte species (Equivalent Spherical Diameter, ESD = 5.6 microm). We also calculated grazing coefficients by combining field data on abundances of C. polykrikoides and co-occurring cryptophytes with laboratory data on ingestion rates obtained in the present study. Cocholdinium polykrikoides fed on prey cells by engulfing the prey through the sulcus. Among the phytoplankton prey offered, C. polykrikoides ingested small phytoplankton species that had ESD's < or = 11 microm (e.g. the prymnesiophyte Isochrysis galbana, an unidentified cryptophyte, the cryptophyte Rhodomonas salina, the raphidophyte Heterosigma akashiwo, and the dinoflagellate Amphidinium carterae). It did not feed on larger phytoplankton species that had ESD's > or = 12 microm (e.g. the dinoflagellates Heterocapsa triquetra, Prorocentrum minimum, Scrippsiella sp., Alexandrium tamarense, Prorocentrum micans, Gymnodinium catenatum, Akashiwo sanguinea, and Lingulodinium polyedrum). Specific growth rates of C. polykrikoides on a cryptophyte increased with increasing mean prey concentration, with saturation at a mean prey concentration of approximately 270 ng C ml(-1) (i.e. 15,900 cells ml(-1)). The maximum specific growth rate (mixotrophic growth) of C. polykrikoides on a cryptophyte was 0.324 d(-1), under a 14:10 h light-dark cycle of 50 microE m(-2) s(-1), while its growth rate (phototrophic growth) under the same light conditions without added prey was 0.166 d(-1). Maximum ingestion and clearance rates of C. polykrikoides on a cryptophyte were 0.16 ng C grazer(-1)d(-1) (9.4 cells grazer(-1)d(-1)) and 0.33 microl grazer(-1)h(-1), respectively. Calculated grazing coefficients by C. polykrikoides on cryptophytes were 0.001-0.745 h(-1) (i.e. 0.1-53% of cryptophyte populations were removed by a C. polykrikoides population in 1 h). The results of the present study suggest that C. polykrikoides sometimes has a considerable grazing impact on populations of cryptophytes.