Dinoflagellate

The dinoflagellates (Greek δῖνος dinos "whirling" and Latin flagellum "whip,
Dinoflagellata
scourge") are a large group of flagellate eukaryotes that constitute the phylum
Dinoflagellata. Most are marine plankton, but they also are common in Temporal range: 240–0 Ma [1]
freshwater habitats. Their populations are distributed depending on sea surface PreЄ Є OS D C P T J K Pg N
temperature, salinity, or depth. Many dinoflagellates are known to be Triassic or earlier–Present
photosynthetic, but a large fraction of these are in fact mixotrophic, combining
photosynthesis with ingestion of prey (phagotrophy).[5] In terms of number of
species, dinoflagellates are one of the largest groups of marine eukaryotes,
although this group is substantially smaller than diatoms.[6] Some species are
endosymbionts of marine animals and play an important part in the biology of
coral reefs. Other dinoflagellates are unpigmented predators on other protozoa,
and a few forms are parasitic (for example,oodinium and pfiesteria). Some
dinoflagellates produce resting stages, called dinoflagellate cysts or dinocysts, as
part of their lifecycles.

Dinoflagellates are considered to be protists, with their own division,

Dinoflagellata.[7]

About 1,555 species of free-living marine dinoflagellates are currently

described.[8] Another estimate suggests about 2,000 living species, of which more
than 1,700 are marine (free-living, as well as benthic) and about 220 are from
fresh water.[9] The latest estimates suggest a total of 2,294 living dinoflagellate
species, which includes marine, freshwater, and parasitic dinoflagellates.[2] Ceratium sp.

A bloom of certain dinoflagellates can result in a visible coloration of the water Scientific classification
colloquially known as red tide, which can cause shellfish poisoning if humans eat
Domain: Eukaryota
contaminated shellfish. Some dinoflagellates also exhibit bioluminescence—
primarily emitting blue-green light. (unranked): SAR
(unranked): Alveolata
Phylum: Dinoflagellata
Bütschli 1885 [1880-
Contents
1889] sensu Gomez
History
2012 [2][3][4]
Morphology
Theca structure and formation Classes
The dinoflagellate nucleus: dinokaryon
Classification Ellobiophyceae
Identification
Psammosea
Ecology and physiology
Habitats
Oxyrrhea
Endosymbionts Pronoctilucea
Nutritional strategies
Harmful algal blooms
Duboscquellea
Bioluminescence Syndiniophyceae
Lipid and sterol production
Noctiluciphyceae
Transport
 Life cycle Dinophyceae
Genomics
Synonyms
Evolutionary history
Examples Cilioflagellata Claparède &
See also
Lachmann, 1868
References
Dinophyta Dillon, 1963
Bibliography
External links Dinophyceae sensu Pascher,
1914

Pyrrophyta Pascher 1914

History Pyrrhophycophyta Papenfuss
In 1753, the first modern dinoflagellates were described by Henry Baker as 1946
"Animalcules which cause the Sparkling Light in Sea Water",[10] and named by
Arthrodelen Flagellaten Stein
Otto Friedrich Müller in 1773.[11] The term derives from the Greek word δῖνος
(dinos), meaning whirling, and Latin flagellum, a diminutive term for a whip or
1883
scourge. Dinomastigota Margulis &
Sagan, 1985
In the 1830s, the German microscopist Christian Gottfried Ehrenberg examined
many water and plankton samples and proposed several dinoflagellate genera that Dinophyta Dillon, 1963
are still used today includingPeridinium, Prorocentrum, and Dinophysis.[12]

These same dinoflagellates were first defined by Otto Bütschli in 1885 as the flagellate order Dinoflagellida.[13] Botanists treated
them as a division of algae, named Pyrrophyta or Pyrrhophyta ("fire algae"; Greek pyrr(h)os, fire) after the bioluminescent forms,
or Dinophyta. At various times, the cryptomonads, ebriids, and ellobiopsids have been included here, but only the last are now
considered close relatives. Dinoflagellates have a known ability to transform from noncyst to cyst-forming strategies, which makes
recreating their evolutionary history extremely dif
ficult.

Morphology
Dinoflagellates are unicellular and possess two dissimilar flagella arising from the ventral cell side (dinokont flagellation). They have
a ribbon-like transverse flagellum with multiple waves that beats to the cell's left, and a more conventional one, the longitudinal
flagellum, that beats posteriorly.[14][15][16] The transverse flagellum is a wavy ribbon in which only the outer edge undulates from
base to tip, due to the action of the axoneme which runs along it. The axonemal edge has simple hairs that can be of varying lengths.
The flagellar movement produces forward propulsion and also a turning force. The longitudinal flagellum is relatively conventional
in appearance, with few or no hairs. It beats with only one or two periods to its wave. The flagella lie in surface grooves: the
transverse one in the cingulum and the longitudinal one in the sulcus, although its distal portion projects freely behind the cell. In
dinoflagellate species with desmokont flagellation (e.g., Prorocentrum), the two flagella are differentiated as in dinokonts, but they
are not associated with grooves.

Dinoflagellates have a complex cell covering called an amphiesma or cortex, composed of a series of membranes, flattened vesicles
called alveolae (= amphiesmal vesicles) and related structures.[17][18] In armoured dinoflagellates, these support overlapping
cellulose plates to create a sort of armor called the theca, as opposed to athecate dinoflagellates. These occur in various shapes and
arrangements, depending on the species and sometimes on the stage of the dinoflagellate. Conventionally, the term tabulation has
been used to refer to this arrangement of thecal plates. The plate configuration can be denoted with the plate formula or tabulation
formula. Fibrous extrusomes are also found in many forms. Together with various other structural and genetic details, this
organization indicates a close relationship between the dinoflagellates, the Apicomplexa, and ciliates, collectively referred to as the
alveolates.[19]

Dinoflagellate tabulations can be grouped into six "tabulation types": gymnodinoid, suessoid, gonyaulacoid–peridinioid,
nannoceratopsioid, dinophysioid, and prorocentroid.
The chloroplasts in most photosynthetic dinoflagellates are bound by three
membranes, suggesting they were probably derived from some ingested algae. Most
photosynthetic species contain chlorophylls a and c2, the carotenoid beta-carotene,
and a group of xanthophylls that appears to be unique to dinoflagellates, typically
peridinin, dinoxanthin, and diadinoxanthin. These pigments give many
dinoflagellates their typical golden brown color. However, the dinoflagellates
Karenia brevis, Karenia mikimotoi, and Karlodinium micrum have acquired other
pigments through endosymbiosis, including fucoxanthin.[20] This suggests their
chloroplasts were incorporated by several endosymbiotic events involving already
colored or secondarily colorless forms. The discovery of plastids in the Apicomplexa
has led some to suggest they were inherited from an ancestor common to the two
groups, but none of the more basal lines has them. All the same, the dinoflagellate
cell consists of the more common organelles such as rough and smooth endoplasmic
reticulum, Golgi apparatus, mitochondria, lipid and starch grains, and food vacuoles.
Some have even been found with a light-sensitive organelle, the eyespot or stigma,
or a larger nucleus containing a prominent nucleolus. The dinoflagellate
Erythropsidium has the smallest known eye.[21]
Longitudinal (l.f.) and transverse
Some athecate species have an internal skeleton consisting of two star-like siliceous flagellum (t.f.); sack pusule (s.p.);
elements that has an unknown function, and can be found as microfossils. nucleus (n).
Tappan[22] gave a survey of dinoflagellates with internal skeletons. This included
the first detailed description of the pentasters in Actiniscus pentasterias, based on
scanning electron microscopy. They are placed within the order Gymnodiniales,
suborder Actiniscineae.[7]

Theca structure and formation

The formation of thecal plates has been studied in detail through ultrastructural
studies.[18]

The dinoflagellate nucleus: dinokaryon

Most dinoflagellates have a peculiar form of nucleus, called a dinokaryon, in which the chromosomes are attached to the nuclear
membrane. These carry reduced number of histones. In place of histones, dinoflagellate nuclei contain a novel, dominant family of
nuclear proteins that appear to be of viral origin, thus are called dinoflagellate/ viral nucleoproteins (DVNPs) which are highly basic,
bind DNA with similar affinity to histones, and occur in multiple posttranslationally modified forms.[23] Dinoflagellate nuclei remain
condensed throughout interphase rather than just during mitosis, which is closed and involves a uniquely extranuclear mitotic
spindle.[24] This sort of nucleus was once considered to be an intermediate between the nucleoid region of prokaryotes and the true
nuclei of eukaryotes, so were termed mesokaryotic, but now are considered advanced rather than primitive traits.

Classification
Dinoflagellates are protists which have been classified using both the International Code of Botanical Nomenclature (ICBN, now
renamed as ICN) and the International Code of Zoological Nomenclature (ICZN). About half of living dinoflagellate species are
autotrophs possessing chloroplasts and half are nonphotosynthesising heterotrophs.

Most (but not all) dinoflagellates have a dinokaryon, described below (see: Life cycle, below). Dinoflagellates with a dinokaryon are
classified under Dinokaryota, while dinoflagellates without a dinokaryon are classified underSyndiniales.
Although classified as eukaryotes, the dinoflagellate nuclei are not characteristically
eukaryotic, as some of them lack histones and nucleosomes, and maintain
continually condensed chromosomes during mitosis. The dinoflagellate nucleus was
termed ‘mesokaryotic’ by Dodge (1966),[25] due to its possession of intermediate
characteristics between the coiled DNA areas of prokaryotic bacteria and the well-
defined eukaryotic nucleus. This group, however, does contain typically eukaryotic
[26]
organelles, such as Golgi bodies, mitochondria, and chloroplasts.

Jakob Schiller (1931–1937) provided a description of all the species, both marine
and freshwater, known at that time.[27] Later, Alain Sournia (1973, 1978, 1982,
1990, 1993) listed the new taxonomic entries published after Schiller (1931–
1937).[28][29][30][31][32] Sournia (1986) gave descriptions and illustrations of the
marine genera of dinoflagellates, excluding information at the species level.[33] The
latest index is written by Gómez.[2]

Identification
English-language taxonomic monographs covering large numbers of species are
published for the Gulf of Mexico,[34] the Indian Ocean,[35] the British Isles,[36] the
Mediterranean[37] and the North Sea.[38]

The main source for identification of freshwater dinoflagellates is the Süsswasser

1. Ornithocercus; 2. diagram; 3.
Flora.[39] Exuviaeella; 4. Prorocentrum; 5, 6.
[40] Ceratium; 7. Pouchetia; 8.
Calcofluor-white can be used to stain thecal plates in armoured dinoflagellates.
Citharistes; 9. Polykrikos

Ecology and physiology

Habitats
Dinoflagellates can occur in all aquatic environments: marine, brackish, and fresh water, including in snow or ice. They are also
common in benthic environments and sea ice.

Endosymbionts
All Zooxanthellae are dinoflagellates and most of them are members within the genus Symbiodinium.[41] The association between
Symbiodinium and reef-building corals is widely known. However, endosymbiontic Zooxanthellae inhabit a great number of other
invertebrates and protists, for example many sea anemones, jellyfish, nudibranchs, the giant clam Tridacna, and several species of
radiolarians and foraminiferans.[42] Many extant dinoflagellates are parasites (here defined as organisms that eat their prey from the
inside, i.e. endoparasites, or that remain attached to their prey for longer periods of time, i.e. ectoparasites). They can parasitize
animal or protist hosts. Protoodinium, Crepidoodinium, Piscinoodinium, and Blastodinium retain their plastids while feeding on their
zooplanktonic or fish hosts. In most parasitic dinoflagellates, the infective stage resembles a typical motile dinoflagellate cell.

Nutritional strategies
Three nutritional strategies are seen in dinoflagellates: phototrophy, mixotrophy, and heterotrophy. Phototrophs can be
photoautotrophs or auxotrophs. Mixotrophic dinoflagellates are photosynthetically active, but are also heterotrophic. Facultative
mixotrophs, in which autotrophy or heterotrophy is sufficient for nutrition, are classified as amphitrophic. If both forms are required,
the organisms are mixotrophic sensu stricto. Some free-living dinoflagellates do not have chloroplasts, but host a phototrophic
endosymbiont. A few dinoflagellates may use alien chloroplasts (cleptochloroplasts), obtained from food (kleptoplasty). Some
dinoflagellates may feed on other organisms as predators or parasites.[43]

Food inclusions contain bacteria, bluegreen algae, small dinoflagellates, diatoms, ciliates, and other
dinoflagellates.[44][45][46][47][48][49][50]

Mechanisms of capture and ingestion in dinoflagellates are quite diverse. Several dinoflagellates, both thecate (e.g. Ceratium
hirundinella,[49] Peridinium globulus[47] ) and nonthecate (e.g. Oxyrrhis marina,[45] Gymnodinium sp.[51] and Kofoidinium spp.[52] ),
draw prey to the sulcal region of the cell (either via water currents set up by the flagella or via pseudopodial extensions) and ingest
the prey through the sulcus. In several Protoperidinium spp., e.g. P. conicum, a large feeding veil — a pseudopod called the pallium
— is extruded to capture prey which is subsequently digested extracellularly (= pallium-feeding).[53][54] Oblea, Zygabikodinium, and
Diplopsalis are the only other dinoflagellate genera known to use this particular feeding mechanism[54][55][56] ). Katodinium
(Gymnodinium) fungiforme, commonly found as a contaminant in algal or ciliate cultures, feeds by attaching to its prey and ingesting
prey cytoplasm through an extensible peduncle.[57] The feeding mechanisms of the oceanic dinoflagellates remain unknown,
although pseudopodial extensions were observed inPodolampas bipes.[58]

Harmful algal blooms

Dinoflagellates sometimes bloom in concentrations of more than a million cells per millilitre. Under such circumstances, they can
produce toxins (generally called dinotoxins) in quantities capable of killing fish and accumulating in filter feeders such as shellfish,
which in turn may be passed on to people who eat them. This phenomenon is called a red tide, from the color the bloom imparts to
the water. Some colorless dinoflagellates may also form toxic blooms, such as Pfiesteria. Some dinoflagellate blooms are not
dangerous. Bluish flickers visible in ocean water at night often come from blooms of bioluminescent dinoflagellates, which emit
short flashes of light when disturbed.

The same red tide mentioned above is more specifically produced when
dinoflagellates are able to reproduce rapidly and copiously on account of the
abundant nutrients in the water. Although the resulting red waves are an unusual
sight, they contain toxins that not only affect all marine life in the ocean, but the
people who consume them, as well.[59] A specific carrier is shellfish. This can
introduce both nonfatal and fatal illnesses. One such poison is saxitoxin, a powerful
paralytic neurotoxin. Human inputs of phosphate further encourage these red tides,
so strong interest exists in learning more about dinoflagellates, from both medical
and economic perspectives. The ecology of harmful algal blooms is extensively
Algal bloom (akasio) by Noctiluca
studied.[60]
spp. in Nagasaki

Bioluminescence
At night, water can have an appearance of sparkling light due to the bioluminescence of dinoflagellates.[61][62] More than 18 genera
of dinoflagellates are bioluminescent,[63] and the majority of them emit a blue-green light.[64] These species contain scintillons,
individual cytoplasmic bodies (about 0.5 µm in diameter) distributed mainly in the cortical region of the cell, outpockets of the main
cell vacuole. They contain dinoflagellate luciferase, the main enzyme involved in dinoflagellate bioluminescence, and luciferin, a
chlorophyll-derived tetrapyrrole ring that acts as the substrate to the light-producing reaction. The luminescence occurs as a brief (0.1
sec) blue flash (max 476 nm) when stimulated, usually by mechanical disturbance. Therefore, when mechanically stimulated—by
[65]
boat, swimming, or waves, for example—a blue sparkling light can be seen emanating from the sea surface.

Dinoflagellate bioluminescence is controlled by a circadian clock and only occurs at night.[66] Luminescent and nonluminescent
strains can occur in the same species. The number of scintillons is higher during night than during day, and breaks down during the
[67]
end of the night, at the time of maximal bioluminescence.
The luciferin-luciferase reaction responsible for the bioluminescence is pH
sensitive.[65] When the pH drops, luciferase changes its shape, allowing luciferin,
more specifically tetrapyrrole, to bind.[65] Dinoflagellates can use bioluminescence
as a defense mechanism. They can startle their predators by their flashing light or
they can ward off potential predators by an indirect effect such as the "burglar
alarm". The bioluminescence attracts attention to the dinoflagellate and its attacker,
[65]
making the predator more vulnerable to predation from higher trophic levels.

Bioluminescent dinoflagellate ecosystem bays are among the rarest and most
fragile,[68] with the most famous ones being the Bioluminescent Bay in La Parguera, Long exposure image of
Lajas, Puerto Rico; Mosquito Bay in Vieques, Puerto Rico; and Las Cabezas de San bioluminescence of N. scintillans in
the yacht port of Zeebrugge, Belgium
Juan Reserva Natural Fajardo, Puerto Rico. Also, a bioluminescent lagoon is near
[69]
Montego Bay, Jamaica, and bioluminescent harbors surround Castine, Maine.

Lipid and sterol production

Dinoflagellates produce characteristic lipids and sterols.[70] One of these sterols is
typical of dinoflagellates and is calleddinosterol.

Transport Play media

Kayaking in the Bioluminescent Bay,
Dinoflagellate theca can sink rapidly to the seafloor inmarine snow.[71] Vieques, Puerto Rico

Life cycle
Dinoflagellates have a haplontic life cycle, with the possible
exception of Noctiluca and its relatives.[7] The life cycle usually
involves asexual reproduction by means of mitosis, either through
desmoschisis or eleuteroschisis. More complex life cycles occur,
more particularly with parasitic dinoflagellates. Sexual reproduction
also occurs,[72] though this mode of reproduction is only known in a
small percentage of dinoflagellates.[73] This takes place by fusion of
two individuals to form a zygote, which may remain mobile in
typical dinoflagellate fashion and is then called a planozygote. This
zygote may later form a resting stage orhypnozygote, which is called
a dinoflagellate cyst or dinocyst. After (or before) germination of the
cyst, the hatchling undergoes meiosis to produce new haploid cells.

Genomics Dinoflagellata life cycle: 1-mitosis, 2-sexual

reproduction, 3-planozygote, 4-hypnozygote, 5-
One of their most striking features is the large amount of cellular planomeiocyte
DNA that dinoflagellates contain. Most eukaryotic algae contain on
average about 0.54 pg DNA/cell, whereas estimates of dinoflagellate
DNA content range from 3–250 pg/cell,[24] corresponding to roughly 3000–215 000 Mb (in comparison, the haploid human genome
is 3180 Mb and hexaploidTriticum wheat is 16 000 Mb).Polyploidy or polyteny may account for this large cellular DNA content,[74]
but studies of DNA reassociation kinetics do not support this hypothesis.

In addition to their disproportionately large genomes, dinoflagellate nuclei are unique in their morphology, regulation, and
composition.
The dinoflagellates share an unusual mitochondrial genome organisation with their relatives, the Apicomplexa.[75] Both groups have
very reduced mitochondrial genomes (around 6 kilobases (kb) in the Apicomplexa vs ~16kb for human mitochondria). The genes on
the dinoflagellate genomes have undergone a number of reorganisations, including massive genome amplification and recombination
which have resulted in multiple copies of each gene and gene fragments linked in numerous combinations. Loss of the standard stop
codons, trans-splicing of mRNAs for the mRNA of cox3, and extensive RNA editing recoding of most genes has occurred. The
reasons for this transformation are unknown.

The DNA of the plastid in the peridinin-containing dinoflagellates is contained in a series of small circles.[76] Each circle contains
one or two polypeptide genes. The genes for these polypeptides are chloroplast-specific because their homologs from other
photosynthetic eukaryotes are exclusively encoded in the chloroplast genome. Within each circle is a distinguishable 'core' region.
Genes are always in the same orientation with respect to this core region.

In terms of DNA barcoding, ITS sequences can be used to identify species,[77] where a genetic distance of p≥0.04 can be used to
delimit species.[78]

Evolutionary history
Dinoflagellates are mainly represented as fossils by fossil dinocysts, which have a long geological record with lowest occurrences
during the mid-Triassic,[79] whilst geochemical markers suggest a presence to the Early Cambrian.
[80]

Some evidence indicates dinosteroids in many Paleozoic and Precambrian rocks might be the product of ancestral dinoflagellates
(protodinoflagellates).[81][82]

Molecular phylogenetics show that dinoflagellates are grouped with ciliates and apicomplexans (=Sporozoa) in a well-supported
clade, the alveolates. The closest relatives to dinokaryotic dinoflagellates appear to be apicomplexans, Perkinsus, Parvilucifera,
syndinians, and Oxyrrhis.[83] Molecular phylogenies are similar to phylogenies based on morphology
.[84]

The earliest stages of dinoflagellate evolution appear to be dominated by parasitic lineages, such as perkinsids and syndinians (e.g.
Amoebophrya and Hematodinium).[85][86][87][88]

All dinoflagellates contain red algal plastids or remnant (nonphotosynthetic) organelles of red algal origin.[89] The parasitic
dinoflagellate Hematodinium however lacks a plastid entirely.[90]

Dinoflagellate evolution has been summarized into five principal organizational types: prorocentroid, dinophysoid, gonyaulacoid,
peridinioid, and gymnodinoid.[91] The transitions of marine species into fresh water have been infrequent events during the
, possibly as late as theCretaceous.[92]
diversification of dinoflagellates and in most cases have not occurred recently

Recently, the "living fossil" Dapsilidinium pastielsii was found inhabiting the Indo-Pacific Warm Pool, which served as a refugium
for thermophilic dinoflagellates.[93]

Examples
Alexandrium
Gonyaulax
Gymnodinium
Lingulodinium polyedrum
 Oxyrrhis marina Dinophysis acuminata Ceratium macroceros Ceratium furcoides
(Oxyrrhea) (Dinophyceae) (Dinophyceae) (Dinophyceae)

Unknown dinoflagellate Pfiesteria shumwayae Symbiodinium sp. Noctiluca scintillans

under SEM (Dinophyceae) (Dinophyceae): (Noctiluciphyceae)
(Dinophyceae) zooxanthella, a coral
endosymbiont

See also
Ciguatera
Paralytic shellfish poisoning
Yessotoxin

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Bibliography
Spector, D.L. (1984). Dinoflagellates. Academic Press. ISBN 978-0-323-13813-0.
Taylor, F.J.R. (1987). The Biology of Dinoflagellates. Botanical monographs.21. Blackwell Scientific.
ISBN 0632009152.

External links
International Society for the Study of Harmful Algae
Classic dinoflagellate monographs
Japanese dinoflagellate site
Noctiluca scintillans — Guide to the Marine Zooplankton of south eastern Australia
, Tasmanian Aquaculture &
Fisheries Institute
Tree of Life Dinoflagellates
Centre of Excellence for Dinophyte T axonomy CEDiT
Dinoflagellates
Judson O (5 January 2010). "A Tale of Two Flagella". New York Times.

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