PIGMENTS,FATTY ACIDS,AND STEROLS OF THE TOXIC DINOFLAGELLATE GYMNODINIUM CATENATUM1

Cultures and field samples of the toxic dinoflagellate Gymnodinium catenatum Graham from Tasmania, Australia, were analyzed for pigment, fatty acid, and sterol composition. Gymnodinium catenatum contained the characteristic pigments of photosynthetic dinoflagellates, including chlorophyll a, chlorophyll c₂, and the carotenoids peridinin, dinoxanthin, diadinoxanthin, diatoxanthin, and β,β-carotene. In midlogarithmic and early stationary phase cultures, the chlorophyll a content ranged 50–72 pg · cell^?1, total lipids 956–2084 pg · cell^?1, total fatty acids 426–804 pg · cell^?1, and total sterols 8–20 pg · cell^?1. The major fatty acids (in order of decreasing abundance) were 16:0, 22:6(n-3), and 20:5(n-3) (collectively 65–70% of the total fatty acids), followed by 16:1(n-7), 18:2(n-6), and 14:0. This distribution is characteristic of most dinoflagellates, except for the low abundance (<3%) of the fatty acid 18:5(n-3), considered by some authors to be a marker for dinoflagellates. The three major sterols were 4α-methyl-5α-cholest-7-en-3β-ol, 4α,23,24-trimethyl-5α-cholest-22E-en-3β-ol (the dinoflagellate sterol, dinosterol), and 4α,23,24-trimethyl-5α-cholest-7-en-3β-ol. These three sterols comprised about 75% of the total sterols in both logarithmic and early stationary phase cultures, and they were also found in high proportions (22–25%) in natural dinoflagellate bloom samples. 4-Desmethyl sterols, which are common in most microalgae, were only present in trace amounts in G. catenatum. The chemotaxonomic affinities of G. catenatum and the potential for using specific signature lipids for monitoring toxic dinoflagellate blooms are discussed.     

THE UNIQUE,MICRORETICULATE CYST OF THE NAKED DINOFLAGELLATE GYMNODINIUM CATENATUM1

Gymnodinium catenatum Graham is an unarmored dinoflagellate responsible for episodes of paralytic shellfish poisoning. This species forms a resting cyst that is unique in several ways. The outer surface of the spherical, brownish cyst is microreticulate and composed of hundreds of 1-3 μm polygons. In several regions, these polygons are smaller, more uniform in shape, and oriented in distinct bands that define morphological features. These features on the cyst reflect the cingulum, sulcus, flagellar pore complex, and acrobase of the motile stage precursor to the cyst. The archeopyle is irregularly but extensively developed. Its margin is generally smooth and extends almost completely around the circumference of the cyst, though not consistently in the plane of the equator. The cyst wall is resistant to acetolysis and standard palynological preparation techniques. Gymnodinium catenatum Graham is emended to include the details of the cyst stage. The significance of this cyst is that it is the first described cyst of a naked dinoflagellate that bears oriented surface ornamentation reflecting features of the motile dinoflagellate. Its microreticulate surface ornamentation is unique to dinocysts, naked or armored, living or fossilized. Resistance of the cyst wall to harsh processing techniques suggests the presence of sporopollenin-like material commonly associated with cysts of armored dinoflagellates. From an ecological standpoint, the existence of a G. catenatum cyst has important implications with respect to species bloom dynamics and geographic distribution. In addition, the distinct differences between this cyst and those of the armored saxitoxin-producing gonyaulacoid species argues against a proposed evolutionary linkage.     

ULTRASTRUCTURAL ASPECTS OF SEXUAL REPRODUCTION IN THE RED TIDE DINOFLAGELLATE GONYAULAX TAMARENSIS1

The marine dinoflagellate Gonyaulax tamarensis Lebour is best known for its propensity to form blooms known as red tides in coastal waters worldwide. This paper examines the sexual cycle of this organism using light and electron microscopy. Sexual reproduction begins with contact between thecate gametes which subsequently shed their thecae to fuse along their pellicular layers. Nuclear fusion occurs well after cytoplasmic fusion and is characterized by several distinctive features: a highly vesiculate nucleoplasm without microtubules; nucleoli and V-shaped chromosomes abut the nuclear envelope distal to the region of nuclear contact; and each chromosome possesses a longitudinal line, the central chromosomal axis. Fusion results in a planozygote with numerous cytoplasmic storage products and a slightly thickened layer beneath the pellicle. Subsequent loss of thecal plates and a thickening of the sub-pellicular layer results in a non-motile hypnozygote. A newly-formed hypnozygote possesses numerous minute papillae along its outer surface, formed by the up-folding of the accumulating wall layer. Maturation of the hypnozygote wall results in a smooth three-layered wall, the outermost layer of which is the pellicular layer. Hypnozygote germination produces a large quadriflagellate plan-omeiocyte with a single nucleus and thecal plates identical to vegetative cells. Two subsequent divisions, presumably meiotic, result in Jour cells morphologically identical to vegetative cells.     

EFFECTS OF TURBULENCE ON THE MARINE DINOFLAGELLATE GYMNODINIUM NELSONII1

Laboratory experiments were conducted to study the effects of agitation on growth, cell division, and nucleic acid dynamics of the dinoflagellate Gymnodinium nelsonii Martin. When cultures were placed on an orbital shaker at 100 rpm, cell division was prevented, cellular volume increased up to 1.5 times that of the nonperturbed cells, the form and location of the cell nucleus were modified, and the RNA and DNA concentrations per cell increased up to 10 times those of the controls. When shaking was stopped after 10 days, cells divided immediately at about 2/3 of the division rate of the unshaken populations, and all the altered parameters were restored. If the agitation continued for more than 20 days, total cell death and disintegration occurred. Several cellular types differing in size and shape were observed in the control and shaken cultures. One possible hypothesis for these results is that failure of the cell to divide results from physical disturbance of the microtubule assemblage associated with chromosome separation during mitosis. My study suggests that small-scale oceanic turbulence of sufficient intensity may inhibit growth of individual dinoflagellate cells, but immediate development of the population may continue when calm weather follows the active mixing period.     

SHORT-TIME SCALE DEVELOPMENT OF A GYMNODINIUM CATENATUM POPULATION IN THE RIA DE VIGO (NW SPAIN)1

Wind direction and fresh water runoff determine the circulation pattern of the Ría de Vigo (NW Spain), which in turn influence the selection and distribution of its phytoplankton populations. Coastal winds with a south–southwesterly component reverse the positive estuarine circulation in the Ría, causing an off-shore to in-shore flow of surface waters and, consequently, the outflow of inner waters via deeper layers. We found that this reversal imposed a selective force on the phytoplankton population: diatoms, which could not counteract the sinking movement of the surface waters, were diminished, while dinoflagellates remained in the water column. From the end of September to the beginning of October 1993, an accumulation of Gymnodimium catenatum Graham was observed coinciding with an intrusion of coastal water induced by westerly winds which provoked a reversal in the circulation of the Ría. The slow reestablishment of the positive estuarine circulation pattern, which was due to a weak coastal upwelling and considerable fresh water runoff, allowed the population of G. catenatum to flourish.     

GROWTH AND CELL CYCLE OF TWO CLOSELY RELATED RED TIDE-FORMING DINOFLAGELLATES: GYMNODINIUM NAGASAKIENSE AND G. CF. NAGASAKIENSE1

Small-sized vegetative cells were found to co-occur with normal-sized cells in populations of the European bloom-forming dinoflagellate Gymnodinium cf. nagasakiense Takayama et Adachi, currently known as Gyrodinium aureolum Hulburt, but not in populations of the closely related Japanese species Gymnodiniumn agasakiense. We examined how cell size differentiation may influence growth and cell cycle progression under a 12:12-h light: dark cycle in the European taxon, as compared to the Japanese one. Cell number and volume and chlorophyll red fluorescence in both species varied widely during the photocycle. These variations generally appeared to be related lo the division period, which occurred at night, as indicated by the variations of the fraction of binucleated cells (mitotic index) as well as the distribution of cellular DNA content. “Small” cells of G. cf. nagasakiense divided mainly during the first part of the dark period, although a second minor peak of dividing cells could occur shortly before light onset. In contrast, “large” cells displayed a sharp division peak that occurred 9 h after the beginning of the dark period. The lower degree of synchrony of “small” cells could be a consequence of their faster growth. Alternatively, these data may suggest that cell division is lightly controlled by an endogenous clock in “large” cells and much more loosely controlled in “small” cells. Cells of the Japanese species, which were morphologically similar to “large” cells of the European taxon, displayed an intermediate growth pattern between the two cell types of G. cf. nagasakiense, with a division period that extended to most of the dark period.     

DIEL PHASING OF THE CELL-CYCLE IN THE FLORIDA RED TIDE DINOFLAGELLATE, GYMNODINIUM BREVE

The diel cycle is a key regulator of the cell-cycle in many dinoflagellates, but the mechanisms by which the diel cycle entrains the cell-cycle remain poorly understood. In this study, we describe diel phasing of the cell-cycle in the Florida red tide dinoflagellate Gymnodinium breve Davis, determine the diel cue which serves to entrain the cell-cycle, and provide evidence for the presence of cyclin-dependent kinase (CDK), a cell-cycle regulator which may be responsive to this cue. Four laboratory isolates from the West Coast of Florida were compared. When grown on a 16:8 h LD cycle, all isolates displayed phased cell division, with the S-phase beginning 6–8 h into the light phase, and mitosis following 12–14 h later, as determined by flow cytometry. A naturally occurring bloom of G. breve, studied over one diel cycle, displayed diel cell-cycle phasing similar to that in the laboratory cultures, with the S-phase beginning during daylight and the peak of mitosis occurring approximately 4 h after sunset. In the laboratory cultures, the dark/light "dawn" transition was found to provide the diel cue which serves to entrain the G. breve cell-cycle, whereas the light/ dark "dusk" transition did not appear to be involved. Evidence for the presence of CDK in G. breve was obtained using two approaches: (1) identification of a 34-kDa protein, immunoreactive to an antibody against a conserved amino acid sequence (α-PSTAIR) unique to the CDK protein family and (2) inhibition of the cell-cycle by olomoucine, a selective CDK inhibitor. Together, these results provide the basis from which one can begin addressing mechanisms by which the diel cycle regulates the cell-cycle in G. breve.     

OBSERVATIONS ON THE MORPHOLOGY AND SEXUAL REPRODUCTION OF COOLIA MONOTIS (DINOPHYCEAE)1

The surface morphology of the dinoflagellate Coolia monotis Meunier was compared with the surface morphology of Ostreopsis, The apical pore of C. monotis is similar in architecture to that of Ostreopsis but considerably longer (12 μm) than in O. heptagona (8–9 μm) and O. ovata (6–7 μm). A ventral pore in C. monotis is located on the right ventral margin between apical plate l′ and precingular plate 6″ and is similar in appearance and location to the ventral pore of O. ovata. The longitudinal flagellum (20 μm) in C. monotis is longer than in O. ovata (12 μ). Although Coolia and Ostreopsis appear to be distinctly different and should remain as two separate genera, they appear to be related. Cells of C. monotis divided by binary fission. Doubling time was 3–4 days in the logarithmic phase of growth at 23°C, 12:12 h L:D, 30–90 μE-m^?2·s^?1, and a salinity of 36%. Cultures reached cell densities of 2.5 × 10³ cells·L^?1 after 15 days of growth. The sexual process in C. monotis occurred in Erdschreiber's medium when Danish soil extract was substituted with mangrove sediment extract under the culture conditions described above. Gamete fusion produced large biflagellated planozygotes (70–75 μm diam). Planozygote maturation involved cytoplasmic reorganization, loss of motility, development of a spherical shape (80–90 μm diam), and two to three orange accumulation bodies. The cells at this stage appeared to be thin-walled cysts. Further development included reorganization of cyst contents, emergence of non-motile gametes, and development of chloroplasts, sulcus, and girdle. The nucleus of the newly formed cells occupied 50% or more of the total cell volume. Meiosis occurred in the cyst, but nuclear cyclosis was not observed. Four daughter cells were produced within 36–48 h, and motile gametes developed. The gametes exhibited sexuality for 2 months and completed the sexual life cycle by going through a thin-walled cyst stage.     

POTENTIAL IMPORTANCE OF BENTHIC CYSTS OF GONYAULAX TAMARENSIS AND G. EXCAVATA IN INITIATING TOXIC DINOFLAGELLATE BLOOMS1,2,3

Thick-walled, nonmotile cysts (termed hypnocysts) of two dinoflagellates were isolated from estuarine sediments in Cape Cod, Massachusetts, and germinated to produce their respective motile, thecate stages. Hypnocysts from Orleans district were identified as Gonyaulax excuvata (Braarud) Balech sensu Loeblich & Loeblich. Visually identical hypnocysts from Falmouth district were provisionally identified as Gonyaulax tamarensis Lebour. Both species were toxic. A geographic survey in September detected hypnocysts in only the sediments of locations where toxic blooms developed the preceding and following Spring. Laboratory incubation (16 C) of hypnocysts from sediment samples stored in the dark (5 C) for 6 mo initiated excystment by the temperature increase, with no appreciable effect from light regime, nutrient, or chelator concentrations. Motility of excysted germlings was optimum in highly chelated medium and in the presence of light. We conclude that hypnocysts of both tasa are important in seeding recurrent annual blooms, synchronizing early bloom development with vernal warning of seawater and increasing the geographic range of the species. We suggest that many red tides in New England and eastern Canadian waters are initiated through the displacement of motile estuarine populations into nearshore area by tidal advection and surface runoff, although the potential existence and importance of offshore cyst reservoirs cannot be discounted. Evidence is presented that hypnocysts are probable sexual zygotes whereas the thin-walled cysts readily formed in laboratory cultures (pellicle cyst) are asexual. Pellicle cysts are of limited durability, do not overwinter in nature, and therefore do not play a significant role in initiating toxic blooms.     

MORPHOLOGICAL AND BIOCHEMICAL VARIABILITY OF THE TOXIC DINOFLAGELLATE PROROCENTRUM LIMA ISOLATED FROM THREE LOCATIONS AT HERON ISLAND, AUSTRALIA

Seventeen clones of the toxic, epiphytic-benthic dinoflagellate Prorocentrum lima ( Ehrenberg) Dodge isolated from three separate sites on the reef flats of Heron Island, Australia, were acclimated under the same set of environmental conditions. Morphological features examined for each clone included cell surface configuration, size, and dry weight. Physiological and biochemical features determined for each clone included reproduction rates, pigments (chlorophyll a, chlorophyll c₂, peridinin, and other xanthophylls), toxins (okadaic acid and methyl-okadaic acid), and macromolecular compounds (total protein, lipid, and carbohydrate). Variation in morphological features and reproduction rates of clones within and between sites was minimal and not significant. Also, variation in biochemical features within an individual site was low, but pronounced differences existed among sites, the most notable of which was toxin content (okadaic acid and methyl-okadaic acid). The greatest difference in biochemical features was between clones isolated from the southern site and clones isolated from the northern and southeastern sites. Results of a cluster analysis of clonal characters support the view that these two groups represent distinct genotypes. We suggest that these groups originated from separate seed sources and that the genetic integrity of each is maintained through asexual reproduction .     

A NOVEL ASSOCIATION BETWEEN AN ENDEMIC STICKLEBACK AND A PARASITIC DINOFLAGELLATE. 2. MORPHOLOGY AND LIFE CYCLE1

An unusual dinoflagellate has been discovered in association with an endemic population of stickleback, Gasterosteus (L.), from the Queen Charlotte Islands, Canada. The dinoflagellate spends most of its life cycle as a coccoid vegetative cyst, not as a parasitic trophont. The vegetative cyst is unique in containing a rigid fenestrated matrix, which is penetrated by cytoplasmic process that emanate from a central area containing the dinokaryotic nucleus and associated chloroplasts. Some pores in the matrix are filled by oil droplets or starch granules. Intracellular bacteria are found throughout the cyst, sometimes in association with the nucleus. The cytoplasm contains accumulation bodes, microbodies, polyhedral crystals, chloroplasts and polyvesicular bodes. The encysted dinoflagellate has several potential strategies. It can 1) shed its wall and become amoeboid; 2) undergo sporogenesis and give rise to both regular and resistant spores; 3) divide mitotically, with a gradual reduction in the size of daughter cells down to 20 μm; and 4) apparently form a resting cyst, during which it secretes a thick outer wall composed of five layers. Taxonomically, this unusual dinoflagellate appears to be a new member of the Blastodiniales, although its position will become clearer when details of the motile stage are known.     

THE ECOLOGY OF CHONDRUS CRISPUS AT PLYMOUTH,MASSACHUSETTS. I. ONTOGENY,VEGETATIVE ANATOMY,REPRODUCTION, AND LIFE CYCLE

Carpospores and tetraspores germinate and the germlings develop similarly. A prostrate holdfast of regular or irregular outline is formed if the spore makes direct contact with the substrate. If not, rhizoids grow from the few-celled germling unilaterally in no apparent relation to light or gravity, forming a prostrate holdfast if they encounter a solid substrate, and, in any case, eventually an erect shoot of different anatomy. Tetrasporophytes have been grown to reproductive maturity from single spore cultures in the laboratory. The ontogeny of the procarp and subsequent postfertilization events are described from field material. Only a single putative male plant was found when plants bearing procarps were numerous at Plymouth. Evidence from this study and from other reported observations places in doubt the expected role of fertilization in the diphasic, cytological alternation of Chondrus.     

LIFE HISTORIES OF BLIDINGIA MINIMA (CHLOROPHYCEAE), ESPECIALLY SEXUAL REPRODUCTION1

Blidingia minima (Näg. ex Kütz.) Kylin from Muroran, Hokkaido, Japan, has been shown to exhibit four patterns of life history in culture. Sexual reproduction, reported fully for the first time in the genus, occurs in two of them. I. An isomorphic-heteromorphic complex in which erect, tubular sporophytes alternate with dioecious gametophytes of the same forms, with the irregular production by both phases of discoid or pulvinate (pincushion-like) microthalli capable of both sexual and asexual reproduction. II. An alternation of heteromorphic phases in which an erect, tubular sporophyte alternated irregularly with a gametophytic microthallus. III. An asexual alternation of heteromorphic phases in which an erect, tubular frond alternates irregularly with a microthallus by means of quadriflagellate zoospores. IV. An asexual, monophasic life cycle in which erect, tubular fronds are perpetuated by quadriflagellate zoospores. The first pattern occurred in spring populations from one of three sites. The second occurred in populations from two sites. The third and fourth occurred in all populations tested. The life history of Blidingia minima from Muroran is similar to that of Kornmannia zostericola from Muroran.     

SYMBIOSIS IN THE PLANKTONIC FORAMINIFER,ORBULINA UNIVERSA,AND THE ISOLATION OF ITS SYMBIOTIC DINOFLAGELLATE,GYMNODINIUM BÉII SP. NOV.1

The symbiotic association of the spinose planktonic foraminifer, Orbulina universa, with the dinoflagellate, Gymnodinium béii sp. nov., was examined with light and electron microscopy, and the symbiont was isolated into unialgal culture. The intact association is characterized by a diurnal movement of the symbionts from the distal regions of the spines during the day, to perialgal vacuoles within the host cytoplasm at night. This diurnal migration involves a daily endo- exocytotic cycle. Gymnodinium béii is non-motile and spindle-shaped within the host, whereas it is motile and gymnodinoid in shape when in culture. Ultrastructural examination revealed two or more stalked pyrenoids penetrated by lamellae, a typical dinokaryon nucleus and no trichocysts. A distinct ‘flange’projects over the sulcus from the hypocone. The swimming behavior of this dinoflagellate was characterized by intermittent darting events. Swimming speeds during a dart reached velocities of 770 μm. s^?1 as compared to a mean, non-darting swimming velocity of 126 μm. s^?1. Gymnodinium béii is eurythermal and division rates ranged between 0.16 and 0.65 divisions day^?1 for culture temperatures between 6.5 and 25° C respectively.     

BIOGEOGRAPHY OF ASEXUAL AND SEXUAL REPRODUCTION IN CALOGLOSSA (DELESSERIACEAE,RHODOPHYTA) FROM AUSTRALIA AND NEW ZEALAND

Caloglossa species are widely distributed in mangroves and salt marshes around the world and their life history patterns are being investigated in laboratory culture. In Australia all isolates of C. monosticha, C. postiae and C. ogasawaraensis have Polysiphonia‐type (P‐type) sexual life histories. Among the 70 C. leprieurii isolates from Australia and New Zealand P‐type sexual reproduction also is dominant. However, ten isolates of C. leprieurii from the Spencer Gulf and the Gulf of St. Vincent in South Australia give rise to successive tetrasporphyte generations without gametophytes. Moreover, one isolate from Queensland is asexual. Only one South Australia isolate, obtained from Lake Alexandrina at the mouth of the Murray River, is sexual. South Australia and Pacific Mexico are two regions in which asexual reproduction is dominant. In another mangrove dwelling red alga Bostrychia moritiziana (Rhodomelaceae) non‐sexual reproduction also is frequent in Australia, New Caledonia and Bali (Indonesia). This asexual reproductive pattern of tetrasporophytic recycling appears to have arisen independently among individual populations of various red algal species in different regions. Investigations are underway on the molecular phylogeny of the Caloglossa leprieurii isolates.     

SEXUALITY AND CYST FORMATION IN PACIFIC STRAINS OF THE TOXIC D1NOFLAGELLATE GONYAULAX TAMARENSIS1,2

Sexuality has been established for a culture of Gonyaulax tarnarensis Lebour (strain NEPCC–71). The addition of a thick inoculum to a nitrogen–deprived medium results in the occurrence of anisogamous sexual fusion within the first three days in the new culture. Planozygotes, large “lumpy” cells recognizable by their four flagella, may persist up to 2 wk before forming a smooth–walled, oval hypnozygote. The latter resembles cysts released asexually by ecdysis but has a slightly thicker wall. Viable cysts resembling hypnozygotes (zygotic cysts), but with reduced photosynthetic pigmentation, have been isolated from natural murine sediments in Hidden Basin, British Columbia, and a culture (strain NEPCC–254) was initiated from excysted individuals. Zygotic cysts of NEPCC–71 remained encysted in the light at 17 C for 8 wk before excysting. The presence of a ventral pen with toxicity in the latter strain indicates that the taxonomy of G. tamarensis-like organisms is still in a stale of flux and the criteria for recognition of G. excavata (Braarud) Balech as a separate species are not satisfactory as presently formulated.     

REPRODUCTION AND LIFE HISTORY OF THE RED ALGA GALAXAURA OBLONGATA (NEMALIALES,GALAXAURACEAE)1

Culture and morphological studies showed that Galaxaura oblongata (Ellis et Solander) Lamouroux has a triphasic life history with conspicuous gametophytes and small filamentous tetrasporophytes. Development of male and female reproductive structures is very similar and both begin with the enlargement of a terminal cell of a filament branch occupying a normal vegetative position within the apical pit of a thallus branch. In male thalli this modified branch forms a conceptacle in which spermatangia are produced. In female thalli, this modified branch forms a three-celled carpogonial branch consisting of a carpogonium, hypogynous cell and basal cell. Filament branches from the basal cell form a pericarp and the gonimoblast develops directly from the carpogonium. Carposporangia are produced in conceptacles which resemble the male conceptacles. About the time the first carposporangia are produced, the carpogonium, hypogynous cell and basal cell form a large fusion cell. Released carpospores germinate in a unipolar or bipolar manner and form small filamentous thalli. Under short day conditions, cruciate tetrasporangia are produced in small clusters. Tetraspores germinate similarly to carpospores and also form small filamentous thalli. Under low nutrient conditions, small cylindrical thalli develop on the filaments and these appear similar to gametophytes collected in nature.     

ULTRASTRUCTURE OF THE FEEDING APPARATUS AND MYONEMAL SYSTEM OF THE HETEROTROPHIC DINOFLAGELLATE PROTOPERIDINIUM SPINULOSUM1

The feeding veil or pallium of the thecate heterotrophic dinoflagellate Protoperidinium spinulosum Schiller is a highly vesiculate membranous sac containing several arched, sometimes bifurcated microtubular ribbons. It originates from an internal microtubular basket, passes through a sphincter-like osmiophilic ring located inside the posterior flagellar pore, and emerges from the cell at that pore. The osmiophilic ring is part of an interconnected myonemal system (composed of two striated collars and several striated connectives) that is anchored to the pore plate and to two inward protrusions composed of minute sulcal plates. A related species, Protoperidinium punctulatum (Paulsen) Balech, also possesses a microtubular basket/osmiophilic ring complex. Elongate electron-dense bodies within the basket resemble digestive secretory granules found in other protists. Granular, electron-lucent microbodies clustered at the anterior end of the basket may also have a role in prey digestion. Dense membranous whorls observed within a P. spinulosum cell presented as it was preparing to initiate feeding indicate a condensed storage site for pallium membranes. A narrow microtubule-strengthened pseudopodal appendage found in two non-feeding cells constitutes the tow filament that serves as the initial linkage between the dinoflagellate and its food. The structures that constitute the pallium and pallium precursors, described here for the first time, are unlike those of other known protists, although some similarities with the dinoflagellate peduncle are evident. The existence of this unique system of organelles may have important ramifications in the search for evolutionary relationships among protists.