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Marine protists

From Wikipedia, the free encyclopedia

Marine radiolarian – detail from Ernst Haeckel's Kunstformen der Natur,  1904
Marine radiolarian – detail from Ernst Haeckel's Kunstformen der Natur, 1904

Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Protists are eukaryotes that cannot be classified as plants, fungi or animals. They are usually single-celled and microscopic. Life originated as single-celled prokaryotes (bacteria and archaea) and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics, because they are paraphyletic (lacking a common ancestor).

Protists are highly diverse organisms currently organised into 18 phyla, but are not easy to classify.[1][2] Studies have shown high protist diversity exists in oceans, deep sea-vents and river sediments, suggesting a large number of eukaryotic microbial communities have yet to be discovered.[3][4] There has been little research on mixotrophic protists, but recent studies in marine environments found mixotrophic protests contribute a significant part of the protist biomass.[5] Since protists are eukaryotes they possess within their cell at least one nucleus, as well as organelles such as mitochondria and Golgi bodies. Protists are asexual but can reproduce rapidly through mitosis or by fragmentation.

In contrast to the cells of prokaryotes, the cells of eukaryotes are highly organised. Plants, animals and fungi are usually multi-celled and are typically macroscopic. Most protists are single-celled and microscopic. But there are exceptions. Some single-celled marine protists are macroscopic. Some marine slime molds have unique life cycles that involve switching between unicellular, colonial, and multicellular forms.[6] Other marine protist are neither single-celled nor microscopic, such as seaweed.

Protists have been described as a taxonomic grab bag of misfits where anything that doesn't fit into one of the main biological kingdoms can be placed.[7] Some modern authors prefer to exclude multicellular organisms from the traditional definition of a protist, restricting protists to unicellular organisms.[8][9] This more constrained definition excludes many brown, multicellular red and green algae, and slime molds.[10]

By trophic mode

Protists can be broadly divided into four groups depending on whether their nutrition is plant-like, animal-like, fungal-like,[11] or a mixture of these.[12]

Protists according to how they get food
Type of protist Description Example Some other examples
Plant-like Autotrophic protists that make their own food without needing to consume other organisms, usually by using photosynthesis
Pyramimonas sp.jpg
Green algae, Pyramimonas Red and brown algae, diatoms and some dinoflagellates. Plant-like protists are important components of phytoplankton discussed below.
Animal-like Heterotrophic protists that get their food consuming other organisms (bacteria, archaea and small algae)
Haeckel Spumellaria detail.png
Radiolarian protist as drawn by Haeckel Foraminiferans, and some marine amoebae, ciliates and flagellates.
Fungal-like Saprotrophic protists that get their food from the remains of organisms that have broken down and decayed
Aplanonet3.jpg
Marine slime nets form labyrinthine networks of tubes in which amoeba without pseudopods can travel Marine lichen
Mixotrophs
Various
(see below)
Mixotrophic and osmotrophic protists that get their food from a combination of the above
Euglena mutabilis - 400x - 1 (10388739803) (cropped).jpg
Euglena mutabilis, a photosynthetic flagellate Many marine mixotrops are found among protists, particularly among ciliates and dinoflagellates[5]
micrograph
cell schematic
Choanoflagellates, unicellular "collared" flagellate protists, are thought to be the closest living relatives of the animals.[13]

Mixotrophs

Mixotrophic radiolarians
Acantharian radiolarian hosts Phaeocystis symbionts
White Phaeocystis algal foam washing up on a beach

A mixotroph is an organism that can use a mix of different sources of energy and carbon, instead of having a single trophic mode on the continuum from complete autotrophy at one end to heterotrophy at the other. It is estimated that mixotrophs comprise more than half of all microscopic plankton.[16] There are two types of eukaryotic mixotrophs: those with their own chloroplasts, and those with endosymbionts—and others that acquire them through kleptoplasty or by enslaving the entire phototrophic cell.[17]

The distinction between plants and animals often breaks down in very small organisms. Possible combinations are photo- and chemotrophy, litho- and organotrophy, auto- and heterotrophy or other combinations of these. Mixotrophs can be either eukaryotic or prokaryotic.[18] They can take advantage of different environmental conditions.[19]

Recent studies of marine microzooplankton found 30–45% of the ciliate abundance was mixotrophic, and up to 65% of the amoeboid, foram and radiolarian biomass was mixotrophic.[5]

Phaeocystis is an important algal genus found as part of the marine phytoplankton around the world. It has a polymorphic life cycle, ranging from free-living cells to large colonies.[20] It has the ability to form floating colonies, where hundreds of cells are embedded in a gel matrix, which can increase massively in size during blooms.[21] As a result, Phaeocystis is an important contributor to the marine carbon[22] and sulfur cycles.[23] Phaeocystis species are endosymbionts to acantharian radiolarians.[24][25]

Mixotrophic plankton that combine phototrophy and heterotrophy – table based on Stoecker et. al., 2017 [26]
General types Description Example Further examples
Bacterioplankton Photoheterotrophic bacterioplankton
Cholera bacteria SEM.jpg
Vibrio cholerae Roseobacter spp.
Erythrobacter spp.
Gammaproteobacterial clade OM60
Widespread among bacteria and archaea
Phytoplankton Called constitutive mixotrophs by Mitra et. al., 2016.[27] Phytoplankton that eat: photosynthetic protists with inherited plastids and the capacity to ingest prey.
Ochromonas.png
Ochromonas species Ochromonas spp.
Prymnesium parvum
Dinoflagellate examples: Fragilidium subglobosum,Heterocapsa triquetra,Karlodinium veneficum,Neoceratium furca,Prorocentrum minimum
Zooplankton Called nonconstitutive mixotrophs by Mitra et. al., 2016.[27] Zooplankton that are photosynthetic: microzooplankton or metazoan zooplankton that acquire phototrophy through chloroplast retentiona or maintenance of algal endosymbionts.
Generalists Protists that retain chloroplasts and rarely other organelles from many algal taxa
Halteria.jpg
Most oligotrich ciliates that retain plastidsa
Specialists 1. Protists that retain chloroplasts and sometimes other organelles from one algal species or very closely related algal species
Dinophysis acuminata.jpg
Dinophysis acuminata Dinophysis spp.
Myrionecta rubra
2. Protists or zooplankton with algal endosymbionts of only one algal species or very closely related algal species
Noctiluca scintillans varias.jpg
Noctiluca scintillans Metazooplankton with algal endosymbionts
Most mixotrophic Rhizaria (Acantharea, Polycystinea, and Foraminifera)
Green Noctiluca scintillans
aChloroplast (or plastid) retention = sequestration = enslavement. Some plastid-retaining species also retain other organelles and prey cytoplasm.

By locomotion

Another way of categorising protists is according to their mode of locomotion. Many unicellular protists, particularly protozoans, are motile and can generate movement using flagella, cilia or pseudopods. Cells which use flagella for movement are usually referred to as flagellates, cells which use cilia are usually referred to as ciliates, and cells which use pseudopods are usually referred to as amoeba or amoeboids. Other protists are not motile, and consequently have no movement mechanism.

Protists according to how they move
Type of protist Movement mechanism Description Example Other examples
Motile Flagellates
Locomotion by flagellum.jpg
A flagellum (Latin for whip) is a lash-like appendage that protrudes from the cell body of some protists (as well as some bacteria). Flagellates use from one to several flagella for locomotion and sometimes as feeding and sensory organelle.
CSIRO ScienceImage 6743 SEM Cryptophyte.jpg
Cryptophytes All dinoflagellates and nanoflagellates (choanoflagellates, silicoflagellates, most green algae)[28][29]
(Other protists go through a phase as gametes when they have temporary flagellum – some radiolarians, foraminiferans and Apicomplexa)
Ciliates
Locomotion by cilia.jpg
A cilium (Latin for eyelash) is a tiny flagellum. Ciliates use multiple cilia, which can number in many hundreds, to power themselves through the water.
Stichotricha secunda - 400x (14974779356).jpg
Paramecium bursaria
click to see cillia
Foraminiferans, and some marine amoebae, ciliates and flagellates.
Amoebas
(amoeboids)
Locomotion by pseudopod.jpg
Amoeba have the ability to alter shape by extending and retracting pseudopods (Greek for false feet).[30] Amoeba Found in every major protist lineage. Amoeboid cells occur among the protozoans, but also in the algae and the fungi.[31][32]
Not motile
none
Diatom Diatoms, coccolithophores, and non‐motile species of Phaeocystis[29] Among protozoans the parasitic Apicomplexa are non‐motile.

Flagellates include bacteria as well as protists. The rotary motor model used by bacteria uses the protons of an electrochemical gradient in order to move their flagella. Torque in the flagella of bacteria is created by particles that conduct protons around the base of the flagellum. The direction of rotation of the flagella in bacteria comes from the occupancy of the proton channels along the perimeter of the flagellar motor.[33]

Ciliates generally have hundreds to thousands of cilia that are densely packed together in arrays. During movement, an individual cilium deforms using a high-friction power stroke followed by a low-friction recovery stroke. Since there are multiple cilia packed together on an individual organism, they display collective behavior in a metachronal rhythm. This means the deformation of one cilium is in phase with the deformation of its neighbor, causing deformation waves that propagate along the surface of the organism. These propagating waves of cilia are what allow the organism to use the cilia in a coordinated manner to move. A typical example of a ciliated microorganism is the Paramecium, a one-celled, ciliated protozoan covered by thousands of cilia. The cilia beating together allow the Paramecium to propel through the water at speeds of 500 micrometers per second.[34]

Algae

Algae is an informal term for a widespread and diverse group of photosynthetic protists which are not necessarily closely related and are thus polyphyletic. Marine algae can be divided into six groups: green, red and brown algae, euglenophytes, dinoflagellates and diatoms.

Dinoflagellates and diatoms are important components of marine algae and have their own sections below. Euglenophytes are a phylum of unicellular flagellates with only a few marine members.

Not all algae are microscopic. Green, red and brown algae all have multicellular macroscopic forms that make up the familiar seaweeds. Green algae, an informal group, contains about 8,000 recognised species.[35] Many species live most of their lives as single cells or are filamentous, while others form colonies made up from long chains of cells, or are highly differentiated macroscopic seaweeds. Red algae, a (disputed) phylum contains about 7,000 recognised species,[36] mostly multicellular and including many notable seaweeds.[36][37] Brown algae form a class containing about 2,000 recognised species,[38] mostly multicellular and including many seaweeds such as kelp. Unlike higher plants, algae lack roots, stems, or leaves. They can be classified by size as microalgae or macroalgae.

Microalgae are the microscopic types of algae, not visible to the naked eye. They are mostly unicellular species which exist as individuals or in chains or groups, though some are multicellular. Microalgae are important components of the marine protists discussed above, as well as the phytoplankton discussed below. They are very diverse. It has been estimated there are 200,000-800,000 species of which about 50,000 species have been described.[39] Depending on the species, their sizes range from a few micrometers (µm) to a few hundred micrometers. They are specially adapted to an environment dominated by viscous forces.

Macroalgae are the larger, multicellular and more visible types of algae, commonly called seaweeds. Seaweeds usually grow in shallow coastal waters where they are anchored to the seafloor by a holdfast. Like microalgae, macroalgae (seaweeds) can be regarded as marineprotists since they are not true plants. But they are not microorganisms, so they are not within the scope of this article.

Unicellular organisms are usually microscopic, less than one tenth of a millimeter long. There are exceptions. Mermaid's wineglass, a genus of subtropical green algae, is single-celled but remarkably large and complex in form with a single large nucleus, making it a model organism for studying cell biology.[41] Another single-celled algae, Caulerpa taxifolia, has the appearance of a vascular plant including "leaves" arranged neatly up stalks like a fern. Selective breeding in aquariums to produce hardier strains resulted in an accidental release into the Mediterranean where it has become an invasive species known colloquially as killer algae.[42]

Diatoms

Diatom shapes
          Drawings by Haeckel 1904 (click for details)

Diatoms form a (disputed) phylum containing about 100,000 recognised species of mainly unicellular algae. Diatoms generate about 20 percent of the oxygen produced on the planet each year,[14] take in over 6.7 billion metric tons of silicon each year from the waters in which they live,[43] and contribute nearly half of the organic material found in the oceans.

Diatoms
Diatoms have a silica shell (frustule) with radial (centric) or bilateral (pennate) symmetry

Diatoms are enclosed in protective silica (glass) shells called frustules. Each frustule is made from two interlocking parts covered with tiny holes through which the diatom exchanges nutrients and wastes.[44] The frustules of dead diatoms drift to the ocean floor where, over millions of years, they can build up as much as half a mile deep.[45]

Structure of a centric diatom frustule [47]
Structure of a centric diatom frustule [47]

Coccolithophores

Coccolithophores are minute unicellular photosynthetic protists with two flagella for locomotion. Most of them are protected by a shell covered with ornate circular plates or scales called coccoliths. The coccoliths are made from calcium carbonate. The term coccolithophore derives from the Greek for a seed carrying stone, referring to their small size and the coccolith stones they carry. Under the right conditions they bloom, like other phytoplankton, and can turn the ocean milky white.[48]

Coccolithophores
...have plates called coccoliths
...extinct fossil
Coccolithophores build calcite skeletons important to the marine carbon cycle[49]

Dinoflagellates

Dinoflagellate shapes
Unarmored dinoflagellates Kofoid (1921)
Haeckel Peridinea (1904)

Dinoflagellates are usually positioned as part of the algae group, and form a phylum of unicellular flagellates with about 2,000 marine species.[50] The name comes from the Greek "dinos" meaning whirling and the Latin "flagellum" meaning a whip or lash. This refers to the two whip-like attachments (flagella) used for forward movement. Most dinoflagellates are protected with red-brown, cellulose armour. Like other phytoplankton, dinoflagellates are r-strategists which under right conditions can bloom and create red tides. Excavates may be the most basal flagellate lineage.[28]

By trophic orientation dinoflagellates are all over the place. Some dinoflagellates are known to be photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey (phagotrophy).[51] Some species are endosymbionts of marine animals and other protists, and play an important part in the biology of coral reefs. Others predate other protozoa, and a few forms are parasitic. Many dinoflagellates are mixotrophic and could also be classified as phytoplankton.

The toxic dinoflagellate Dinophysis acuta acquire chloroplasts from its prey. "It cannot catch the cryptophytes by itself, and instead relies on ingesting ciliates such as the red Myrionecta rubra, which sequester their chloroplasts from a specific cryptophyte clade (Geminigera/Plagioselmis/Teleaulax)".[26]

A surf wave at night sparkles with blue light due to the presence of a bioluminescent dinoflagellate, such as Lingulodinium polyedrum
Suggested explanation for glowing seas[52]
Dinoflagellates
        Armoured
        Unarmoured
Traditionally dinoflagellates have been presented as armoured or unarmoured

Dinoflagellates often live in symbiosis with other organisms. Many nassellarian radiolarians house dinoflagellate symbionts within their tests.[53] The nassellarian provides ammonium and carbon dioxide for the dinoflagellate, while the dinoflagellate provides the nassellarian with a mucous membrane useful for hunting and protection against harmful invaders.[54] There is evidence from DNA analysis that dinoflagellate symbiosis with radiolarians evolved independently from other dinoflagellate symbioses, such as with foraminifera.[55]

Some dinoflagellates are bioluminescent. At night, ocean water can light up internally and sparkle with blue light because of these dinoflagellates.[56][57] Bioluminescent dinoflagellates possess scintillons, individual cytoplasmic bodies which contain dinoflagellate luciferase, the main enzyme involved in the luminescence. The luminescence, sometimes called the phosphorescence of the sea, occurs as brief (0.1 sec) blue flashes or sparks when individual scintillons are stimulated, usually by mechanical disturbances from, for example, a boat or a swimmer or surf.[58]

Protozoans

Protozoans are protists which feed on organic matter such as other microorganisms or organic tissues and debris.[62][63] Historically, the protozoa were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae.[64][65] Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to identify single-celled organisms that can move independently and feed by heterotrophy.

Marine protozoans include zooflagellates, foraminiferans, radiolarians and some dinoflagellates.

Radiolarians

Radiolarian shapes
          Drawings by Haeckel 1904 (click for details)

Radiolarians are unicellular predatory protists encased in elaborate globular shells usually made of silica and pierced with holes. Their name comes from the Latin for "radius". They catch prey by extending parts of their body through the holes. As with the silica frustules of diatoms, radiolarian shells can sink to the ocean floor when radiolarians die and become preserved as part of the ocean sediment. These remains, as microfossils, provide valuable information about past oceanic conditions.[66]

Turing and radiolarian morphology
Shell of a spherical radiolarian
Shell micrographs
Computer simulations of Turing patterns on a sphere
closely replicate some radiolarian shell patterns[67]

Foraminiferans

Foraminiferan shapes
          Drawings by Haeckel 1904 (click for details)

Like radiolarians, foraminiferans (forams for short) are single-celled predatory protists, also protected with shells that have holes in them. Their name comes from the Latin for "hole bearers". Their shells, often called tests, are chambered (forams add more chambers as they grow). The shells are usually made of calcite, but are sometimes made of agglutinated sediment particles or chiton, and (rarely) of silica. Most forams are benthic, but about 40 species are planktic.[68] They are widely researched with well established fossil records which allow scientists to infer a lot about past environments and climates.[66]

Foraminiferans
...can have more than one nucleus
...and defensive spines
Foraminiferans are important unicellular zooplankton protists, with calcium tests

A number of forams are mixotrophic (see below). These have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates.[68] Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters.[70] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.[71]

Amoeba

Shelled and naked amoeba
                  Amoeba can be shelled (testate) or naked
Amoeba engulfing a diatom

Ciliates

Ciliate shapes
          Drawings by Haeckel 1904 (click for details)

Macroscopic protists

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