Ultrastructure and ontogeny of a new type of eyespot in dinoflagellates (original) (raw)
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Reflective Properties of Different Eyespot Types in Dinoflagellates
Protist, 1999
The reflective properties of different types of dinoflagellate eyespots were investigated using confocal laser scanning microscopy in the epireflection contrast mode. Although the eyespots studied differed with respect to localization (cytosol or plastid) and organization of the globule layer(s), all types effectively absorbed and reflected blue-green laser light (principal lines of 488/514 nm). The relative orientation of the eyespot surface towards the light source strongly influenced the reflective properties. Maximal reflection occurred when the eyespot surface was approximately perpendicular to the light source and rapidly decreased at increasing angles of light incidence. Horizontal and vertical optical sectioning of live and fixed cells resolved differences in the reflection patterns. Focusing of reflected light on the basal portion of the longitudinal flagellum was observed for the cytosolic eyespot of Glenodinium sp. and the triple membrane-bounded eyespot of Peridinium foliaceum, presumably a vestige of a host plastid. This flagellum is thought to be mainly involved in mediating orientational movement responses. In contrast, the reflection patterns obtained from the eyespot of Woloszynskia pascheri, which represents the third and most commonly observed dinoflagellate eyespot type within a plastid, point to only minor focusing. Reflection signals could be followed a considerable distance into the sulcus in all cases, indicating that in dinoflagellate eyespots, irrespective of the presumed receptor location (plasma membrane overlying the eyespot and/or the basal part of the longitudinal flagellum), back reflection of non-absorbed light can enhance the excitation probability of the photoreceptor(s). Such a combined reflection/absorption screen allows maximal contrast modulation and will, in conjunction with the specialized geometry of the dinoflagellate eyespots, increase the directionality of these eyespot apparatuses considerably.
Little is known about the morphological versatility of the unarmoured dinoflagellates. The morphology of an unarmoured dinoflagellate with a distinctive H-shaped contour is described from sub-surface waters of the Strait of Gibraltar (NE Atlantic) and the northern Philippine Sea (NW Pacific). The cell body was slightly hexagonal with two rounded-tip apical arms and two antapical sharp-pointed horns. A carina with a straight groove was located between the two apical arms. The cingulum was excavated, with a descending displacement of more than one-half body length and had an overhang. A round nucleus was located in the left hyposome. These forms may correspond to a gymnodiniacean cell that is able to project body extensions under unfavourable environmental conditions.
The marine dinoflagellate Erythropsidinium possesses an ocelloid, the most elaborate photoreceptor organelle known in a unicellular organism, and a piston, a fast contractile appendage unknown in any other organism. The ocelloid is able to rotate, often before the cell swims. The ocelloid contains lenses that function to concentrate light. The flagellar propulsion is atrophied, and the piston is responsible for locomotion through successive extensions and contractions. During the “locomotion mode”, the contraction is ~4 times faster than the extension. The piston attained up to 50 mm s 1 and the cell jumps backwards at 4 mm s 1, while during the piston extension the cell moves forwards. The net speed of ~1 mm s 1 is faster than other dinoflagellates. The piston usually moved in the “static mode” without significant cell swimming. This study suggests that the piston is also a tactile organelle that scans the surrounding waters for prey. Erythropsidinium feeds on copepod eggs by engulfing. The end of the piston possesses a “suction cup” able to attach the prey and place it into the posterior cavity for engulfing. The cylindrical shape of Erythropsidinium, and the anterior position of the ocelloid and nucleus, are morphological adaptations that leave space for the large vacuole. Observations are provided on morphological development during cell division. Most of the described species of Erythropsidinium apparently correspond to distinct life stages of known species, and the genus Greuetodinium (=Leucopsis) corresponds to an earlier division stage. Dinoflagellata; Erythropsis; eyespot; motile behavior; motility; Nematodinium; peduncle feeding; photoreceptor organelle; protist locomotion; Warnowia
The flagellar apparatus and cytoskeleton of the dinoflagellates
Protoplasma, 1991
Modem microscopical approaches have allowed more accurate investigations of the three-dimensional nature of the dinoflagellate flagellar apparatus (FA) and several other cytoskeletal protein complexes. Our presentation overviews the nature of the dinoflagellate FA and cytoskeleton in a number of taxa and compares them with those of other protists. As with other protists, the FA of the dinoflagellates can be characterized by the presence of fibrous and microtubular components. Our studies and others indicate that the dinoflagellate FA can be expected to possess a striated fibrous root on the basal body of the transverse flagellum and a multimembered microtubular root on the basal body of the longitudinal flagellum. Two other features that appear widespread in the group are the transverse striated root associated microtubule (tsrm) and the transverse microtubular root (tmr). The tsrm extends at least half the length of the transverse striated root while the tmr extends from the transverse basal body toward the exit aperture of the transverse flagelhun. In most cases, the tmr gives rise to several cytoplasmic microtubules at a right angle. The apparent conserved nature of these roots leads us to the conclusion that the dinoflagellate FA can be compared to the FA of the cryptomonads, chrysophytes, and the ciliates for phylogenetic purposes. Of these groups, the chrysophytes possess an FA with the most structures in common with the dinoflagellates. Our immunomicroscopical investigations of the microtubular, actin and centrin components of the dinoflagellate cytoskeleton point to the comparative usefulness of these cytological features.
The encystment of a freshwater dinoflagellate: A light and electron-microscopical study
British Phycological Journal, 1972
The process of encystment, or resting spore formation, in a freshwater dinoflageltate (Woloszynskia tylota nov. comb.) has been studied with both light and electron microscopy. The main features of the process are as follows: (i) the replacement of the theca by a thin, amorphous outer wall, which gradually thickens by the deposition of material on its inner face; (ii) the appearance of a layer of closely-packed lipid droplets at the cytoplasmic margin of the mature cyst, resembling a granular 'inner wall' in the light microscope; (iii) the reduction in size or disappearance of cytoplasmic structures such as chloroplasts, Golgi bodies and pusule; and (iv) the enlargement of a central 'accumulation body' and cytoplasmic vacuoles containing crystals. Comparisons are made with light-microscope studies of encystment of other dinoflagellates, with ultrastructural studies of non-motile division stages, with zooxanthellae and with fossil dinoflagellate cysts or hystrichospheres.
Ultrastructural Features of the Benthic Dinoflagellate Ostreopsis cf. ovata (Dinophyceae)
Protist, 2014
The toxic benthic dinoflagellate Ostreopsis cf. ovata has considerably expanded its distribution range in the last decade, posing risks to human health. Several aspects of this species are still poorly known. We studied ultrastructural features of cultivated and natural populations of Ostreopsis cf. ovata from the Gulf of Naples (Mediterranean Sea) using confocal laser scanning, and scanning and transmission electron microscopy. New information on the morphology and location of several sulcal plates was gained and a new plate designation is suggested that better fits the one applied to other Gonyaulacales. The microtubular component of the cytoskeleton, revealed using an anti--tubulin antibody, consisted of a cortical layer of microtubules arranged asymmetrically in the episome and in the hyposome, complemented by a complex inner microtubular system running from the sulcal area towards the internal part of the cell. The conspicuous canal was delimited by two thick, burin-shaped lobes ending in a tubular ventral opening. The canal was surrounded by mucocysts discharging their content into it. A similar structure has been reported in other benthic and planktonic dinoflagellates and may be interpreted as an example of convergent evolution in species producing large amounts of mucus.
Dinoflagellates: a remarkable evolutionary experiment
American Journal of Botany, 2004
In this paper, we focus on dinoflagellate ecology, toxin production, fossil record, and a molecular phylogenetic analysis of hosts and plastids. Of ecological interest are the swimming and feeding behavior, bioluminescence, and symbioses of dinoflagellates with corals. The many varieties of dinoflagellate toxins, their biological effects, and current knowledge of their origin are discussed. Knowledge of dinoflagellate evolution is aided by a rich fossil record that can be used to document their emergence and diversification. However, recent biogeochemical studies indicate that dinoflagellates may be much older than previously believed. A remarkable feature of dinoflagellates is their unique genome structure and gene regulation. The nuclear genomes of these algae are of enormous size, lack nucleosomes, and have permanently condensed chromosomes. This chapter reviews the current knowledge of gene regulation and transcription in dinoflagellates with regard to the unique aspects of the nuclear genome. Previous work shows the plastid genome of typical dinoflagellates to have been reduced to single-gene minicircles that encode only a small number of proteins. Recent studies have demonstrated that the majority of the plastid genome has been transferred to the nucleus, which makes the dinoflagellates the only eukaryotes to encode the majority of typical plastid genes in the nucleus. The evolution of the dinoflagellate plastid and the implications of these results for understanding organellar genome evolution are discussed.
Plastid-associated galactolipid composition in eyespot-containing dinoflagellates: a review
ALGAE
Relative to the large number of photosynthetic dinoflagellate species, only a select few possess proteinaceous, carotenoid-rich eyespots which have been demonstrated in other algae to act in phototactic responses. The proteins comprising the different categories of dinoflagellate eyespots are positioned in or near the peridinin-containing photosynthetic plastid membranes which are composed primarily of two galactolipids, mono-and digalactosyldiacylglycerol (MGDG and DGDG). Within eyespot-containing dinoflagellates, this arrangement occurs mostly in those with secondary plastids, although some dinoflagellates with tertiary plastids of diatom origin are known to possess eyespots. We here provide an examination of the MGDG and DGDG composition of eyespot-containing dinoflagellates with secondary, peridinin-containing plastids and tertiary plastids of diatom origin to address the fundamental question of whether eyespots and their component proteins and carotenoids are associated with alterations in galactolipid composition when compared to eyespot-lacking photosynthetic dinoflagellates. This is an important question because the dinoflagellate eyespot-plastid membrane system can be considered a more complicated and evolved state of plastid development. Included in this examination are data on the previously unexamined peridinin-and type A eyespot-containing dinoflagellate Margalefidinium polykrikoides, and the type D eyespot-containing, aberrant plastid "dinotom" Durinskia baltica. In addition, we have reviewed the galactolipid composition of algae from the Chlorophyceae, Cryptophyceae, and Euglenophyceae as a comparison to determine if algal classes apart from the Dinophyceae contain altered galactolipids in association with eyespots. We conclude that the presence of an eyespot in dinoflagellates and other algae is not associated with noticeable changes in galactolipid composition.
5 The origin of the dinoflagellate plastid
Journal of Phycology, 2003
The peridinin pigmented dinoflagellate chloroplasts are the result of a secondary endosymbiotic event between a photosynthetic eukaryote and a dinoflagellate. Dinoflagellate chloroplast and nuclear evolution were independent before this endosymbiotic event. To reconstruct the evolution of the dinoflagellate chloroplast, phylogenies were constructed with a chloroplast gene psbB. The gene phylogeny should reflect the evolution of the chloroplast and indicate the plastid donor lineage. Gene sequences derived from the dinoflagellate chloroplast were extremely divergent but suggested that the plastid donor could have been a haptophyte. In an attempt to find better genes for analysis and to further understand gene transfer about 4900 randomly selected expressed genes were sequenced from two dinoflagellates, Lingulodinium polyedra and Amphidinium carterae. From these genes, thirty typically plastid-encoded Copyright by Tsvetan Radoslavov Bachvaroff 2004 ii ACKNOWLEGEMENTS The author would like to acknowledge the following individuals for their contributions to this document. Torstein Tengs taught me valuable laboratory techniques and performed many sequencing reactions on my behalf. He also made substantial intellectual contributions to my understanding of plastid evolution. Dr. David Oldach of the Institute for Human Virology generously loaned equipment and provided encouragement. Professor Robert Rowan of the University of Guam challenged me to find better data. The psbB sequences from haptophytes were obtained from Kamran Shalchian-Tabrizi in Professor Jakobsen's lab at the University of Oslo, Norway. Kamran also contributed to my understanding of dinoflagellate nuclear phylogeny by generously sharing unpublished data. Three talented undergraduates participated in the EST data collection, Ernest Williams, Gregory Concepcion, and Carolyn Rogers. Greg also contributed substantially to the dinoflagellate EST database. A fourth undergraduate, Ali Tabatabai, worked on a dinoflagellate rubisco project not presented here. Finally, I would like to thank Maria Virginia Sanchez Puerta who has read and commented on many subunits of this document, and who has sequenced the organellar genomes of Emiliania.