The dinoflagellates (Greek δῖνος dinos "whirling" and Latin flagellum "whip, scourge") are a large group of flagellate protists. Most are marine plankton, but they are common in fresh water habitats, as well. Their populations are distributed depending on temperature, salinity, or depth. Many dinoflagellates are known to be photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey.[1] Dinoflagellates are the largest group of marine eukaryotes aside from the diatoms. Being primary producers makes them an important part of the aquatic food chain. Some species, called zooxanthellae, are endosymbionts of marine animals and play an important part in the biology of coral reefs. Other dinoflagellates are colorless predators on other protozoa, and a few forms are parasitic (see for example Oodinium, Pfiesteria). Dinoflagellates produce resting stages, called dinoflagellate cysts or dinocysts, as part of their life cycles.
Dinoflagellates are considered to be protists, with their own division, Dinoflagellata.[2]
About 1,555 species of free-living marine dinoflagellates are currently described.[3] Another estimate suggests ca. 2000 living species, of which more than 1700 are marine and about 220 are from freshwater.[4]
An algal bloom of dinoflagellates can result in a visible coloration of the water colloquially known as red tide.
Many reviews have been written on dinoflagellates.[2][5][6][7]
In 1753, the first modern dinoflagellates were described by Henry Baker as "Animalcules which cause the Sparkling Light in Sea Water",[8] and named by Otto Friedrich Müller in 1773.[9] The term derives from the Greek word δῖνος (dinos), meaning 'whirling,' and Latin flagellum, a diminutive term for a whip or scourge.
In the 1830s, the German microscopist C.G. Ehrenberg examined many water and plankton samples and proposed several dinoflagellate genera that are still used today including Peridinium, Prorocentrum and Dinophysis.
These same dinoflagellates were first defined by Otto Bütschli in 1885 as the flagellate order Dinoflagellida.[10] 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 difficult.
Dinoflagellates are protists which have been classified using both the International Code of Botanical Nomenclature (ICBN) and the International Code of Zoological Nomenclature (ICZN), approximately half living dinoflagellate species are autotrophs possessing chloroplasts and half are non-photosynthesising heterotrophs. It is now widely accepted that the ICBN should be used for their classification.
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 under Syndiniales.
Although classified as eukaryotes, the dinoflagellate nuclei are not characteristically eukaryotic, as they lack histones, nucleosomes and maintain continually condensed chromosomes during mitosis. In fact, Dodge (1966)[11] termed the dinoflagellate nucleus as ‘mesokaryotic’, 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 organelles, such as Golgi bodies, mitochondria and chloroplasts[12]
Jakob Schiller (1931–1937) provided a description of all the species, both marine and freshwater, known at that time.[13] Later Alain Sournia (1973, 1978, 1982, 1990, 1993) listed the new taxonomic entries published after Schiller (1931–1937)[14][15][16][17] .[18] Sournia (1986) gave descriptions and illustrations of the marine genera of dinoflagellates, excluding information at the species level.[19] The latest index is written by Gomez.[3]
English-language taxonomic monographs covering large numbers of species are published for the Gulf of Mexico,[20] the Indian Ocean,[21] the British Isles,[22] the Mediterranean[23] and the North Sea.[24]
The main source for identification of freshwater dinoflagellates is the Süsswasser Flora.[25]
The cyst of
Peridinium ovatum, showing the ecdysal opening
Dinoflagellates are unicellular forms with one to three flagellae. Usually, they possess two flagellae: one which extends towards the posterior, called the longitudinal flagellum, and the other forming a lateral circle, called the transverse flagellum. In many forms, these are set into grooves, called the sulcus and cingulum. The transverse flagellum is ribbon-like and coiled, provides most of the force propelling the cell, and often imparts to it a distinctive whirling motion, which is what gives them their name. The longitudinal flagellum acts mainly as a rudder, but provides a small amount of propulsive force, as well.
Dinoflagellates have a complex cell covering called an amphiesma, composed of flattened vesicles called alveoli. In armoured dinoflagellates, these support overlapping cellulose plates to create a sort of armor called the theca, as opposed to naked dinoflagellates. These come 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. 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, Apicomplexa, and ciliates, collectively referred to as the alveolates.[26]
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 goldenbrown color. However, some dinoflagellates have acquired other pigments through endosymbiosis, including fucoxanthin[27]. This suggests their chloroplasts were incorporated by several endosymbiotic events involving already colored or secondarily colorless forms. The discovery of plastids in Apicomplexa has led some to suggest they were inherited from an ancestor common to the two groups, but none of the more basal lines have 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.[28]
Some species have an internal skeleton consisting of two star-like siliceous elements that has an unknown function, and can be found as microfossils. Tappan[29] 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.
Most zooxanthellae are dinoflagellates. The association between dinoflagellates and reef-building corals is widely known, but dinoflagellate endosymbionts inhabit a great number of other invertebrates and protists, for example many sea anemones, jellyfish, nudibranchs, the giant clam Tridacna, as well as several species of radiolarians and foraminiferans.[30] 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.
The formation of thecal plates has been studied in detail through ultrastructural studies.[31]
Dinoflagellates can occur in all aquatic environments: marine, brackish, and fresh water, including in snow or ice.
The dinoflagellates include autotrophs, phagotrophs, symbionts and parasites; photosynthetic species (autotrophs) account for about half of living genera, with the other half being nonphotosynthetic. Completely autotrophic species are however very rare.[32] Some taxa have more than one nutritional strategy (mixotrophic): for example, species of Protoperidinium are both parasitic and photosynthetic.[2]
Food inclusions contain bacteria, bluegreen algae, small dinoflagellates, diatoms, ciliates and other dinoflagellates.[33][34][35][36][37][38][39]
Mechanisms of capture and ingestion in dinoflagellates are quite diverse. Several dinoflagellates, both thecate (e.g. Ceratium hirundinella,;[38] Peridinium globulus,[36]) and nonthecate (e.g. Oxyrrhis marina,;[34] Gymnodinium sp.,;[40] and Kofoidinium spp.,[41]), 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. Protoperidinium conicum extrudes a large feeding veil to capture prey which is subsequently digested extracellularly.[42] 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.[43] The feeding mechanisms of the oceanic dinoflagellates remain unknown, although pseudopodial extensions were observed in Podolampas bipes.[44]
Dinoflagellata life cycle: 1-Binary fission, 2-Sexual reproduction, 3-planozygote, 4-hypnozygote, 5-planomeiocyte
Most dinoflagellates have a peculiar form of nucleus, called a dinokaryon, in which the chromosomes are attached to the nuclear membrane. These lack histones and remain condensed throughout interphase rather than just during mitosis, which is closed and involves a unique external spindle[45]. 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.
In most dinoflagellates, the nucleus is dikaryotic throughout the entire life cycle. They are usually haploid, and reproduce primarily through fission, but sexual reproduction also occurs.[46] This takes place by fusion of two individuals to form a zygote, which may remain mobile in typical dinoflagellate fashion or may form a resting stage, a dinoflagellate cyst or dinocyst, which later undergoes meiosis to produce new haploid cells.[clarification needed]
When conditions become unfavourable, usually when nutrients become depleted or there is insufficient light, some dinoflagellate species alter their life cycles dramatically. Two vegetative cells will fuse together, forming a planozygote. Next is a stage not much different from hibernation called a hypnozygote, when the organism takes in excess fat and oil. At the same time, its body enlarges and the shell gets harder. Sometimes even spikes are formed. When the weather allows it, these dinoflagellates break out of their shells and are in a temporary stage, the planomeiocyte, when they quickly reform their individual thecae and return to the dinoflagellates as at the beginning of the process.
Dinoflagellates sometimes bloom in concentrations of more than a million cells per millilitre. Some species produce neurotoxins, which in such quantities kill fish and accumulate in filter feeders such as shellfish, which in turn may pass them 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.[47] A specific carrier is shellfish. This can introduce both nonfatal and fatal illnesses. One such poison is saxitoxin, a powerful paralytic. Human inputs of phosphate further encourage these red tides, so there is a strong interest in learning more about dinoflagellates, from both medical and economic perspectives. The ecology of harmful algal blooms is extensively studied.[48]
Dinoflagellates produce characteristic lipids and sterols[49]. One of these sterols is typical of dinoflagellates and is called dinosterol.
At night, water can have an appearance of sparkling light due to the bioluminescence of dinoflagellates.[50][51] More than 18 genera of dinoflagellates are bioluminescent, and the majority of them (including Gonyaulax) emit a blue-green wavelength. Therefore, when mechanically stimulated—by boat, swimming or waves, for example—a blue sparkling light can be seen emanating from the sea surface.[52] The luciferin-luciferase reaction responsible for the bioluminescence is pH sensitive.[52] When the pH drops, luciferase changes its shape, allowing luciferin, more specifically tetrapyrrole, to bind.[52] 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".[52] The dinoflagellate can use its bioluminescence to attract attention to itself, thereby bringing attention to the predator and making the predator more vulnerable to predators from higher trophic levels.[52]
Bioluminescent dinoflagellate ecosystem bays are among the rarest and most fragile.[53]
Dinoflagellate theca can sink rapidly to the seafloor in marine snow.[54]
Dinoflagellates are represented by fossil dinocysts, which have a long geological record with lowest occurrences during the mid-Triassic.,[55] whilst geochemical markers suggest a presence to the Early Cambrian[56]
One of the most striking features is the large amount of cellular DNA that dinoflagellates contain. Most eukaryotic algae contain on average about 0.54 pg DNA/cell1, whereas estimates of dinoflagellate DNA content range from 3–250 pg/cell[57], corresponding to approximately 3000–215 000 Mb (in comparison, the haploid human genome is3180 Mb and hexaploid Triticum wheat is 16 000 Mb).It has been suggested that polyploidy or polyteny may account for this large cellular DNA content[58], 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.[59] Both groups have very reduced mitochondrial genomes (~6 kilobases in the Apicomplexa). 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.
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