fbpx
Wikipedia

Marine protists

January 01, 1970

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. Life originated as marine 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. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. 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 for all descendants).

Marine protists
Alga (phytoplankton)
Protozoan (zooplankton)
Protists are usually one-celled microorganisms. They include algae (autotrophs which make their own food) and protozoans (heterotrophs which eat the algae for food). In recent years, researchers have discovered many protists are mixotrophs, which can function in both modes.

Most protists are too small to be seen with the naked eye. They are highly diverse organisms currently organised into 18 phyla, but not easy to classify.[1][2] Studies have shown high protist diversity exists in oceans, deep sea-vents and river sediments, suggesting large numbers 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 protists contribute a significant part of the protist biomass.[5] Since protists are eukaryotes (and not prokaryotes) they possess within their cell at least one nucleus, as well as organelles such as mitochondria and Golgi bodies. Many protist species can switch between asexual reproduction and sexual reproduction involving meiosis and fertilization.[6]

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.[7] 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.[8] Some modern authors prefer to exclude multicellular organisms from the traditional definition of a protist, restricting protists to unicellular organisms.[9][10] This more constrained definition excludes all brown, the multicellular red and green algae, and, sometimes, slime molds (slime molds excluded when multicellularity is defined as "complex").[11]

Background edit

"Marine protists are a polyphyletic group of organisms playing major roles in the ecology and biogeochemistry of the oceans, including performing much of Earth's photosynthesis and driving the carbon, nitrogen, and silicon cycles. In addition, marine protists occupy key positions in the tree of life, including as the closest relatives of metazoans [animals]... Unicellular eukaryotes are often lumped as 'protists', a term that is useful despite its taxonomic irrelevance and origin as a definition by exclusion — a protist being any eukaryote that's not a plant, animal, or fungus".[12]

The ocean represents the largest continuous planetary ecosystem, hosting an enormous variety of organisms, which include microscopic biota such as unicellular eukaryotes (protists). Despite their small size, protists play key roles in marine biogeochemical cycles and harbour tremendous evolutionary diversity.[13][14] Notwithstanding their significance for understanding the evolution of life on Earth and their role in marine food webs, as well as driving biogeochemical cycles to maintain habitability, little is known about their cell biology including reproduction, metabolism and signaling.[12] Most of the biological knowledge available is based on comparison of proteins from cultured species to homologs in genetically tractable model taxa.[15][16][17][18] A main impediment to understanding the cell biology of these diverse eukaryotes is that protocols for genetic modification are available for only a small number of species [19][20] that represent neither the most ecologically relevant protists nor the breadth of eukaryotic diversity. Even so, in the decade to 2020, genome [15][16][17] and transcriptome sequencing initiatives [18] have resulted in nearly 120 million unigenes being identified in protists,[21] which is facilitating the development of genetic tools for model species.[22]

 
Phylogenetic and symbiogenetic tree of living organisms, showing a schematic view of the central position occupied by the protista (protists)
 
Schematic view of the eukaryotic tree of life with effigies of main marine protist representatives [22]

Trophic modes edit

Protists can be divided broadly into four groups depending on whether their nutrition is plant-like, animal-like, fungal-like,[23] or a mixture of these.[24]

Protists according to how they get food
Type of protist Description Example Some other examples
Plant-like
Algae
(see below)
 Autotrophic protists that make their own food without needing to consume other organisms, usually by photosynthesis (sometimes by chemosynthesis)   Green algae, Pyramimonas Red and brown algae, diatoms, coccolithophores and some dinoflagellates. Plant-like protists are important components of phytoplankton discussed below.
Animal-like
Protozoans
(see below)
 Heterotrophic protists that get their food consuming other organisms (bacteria, archaea and small algae)   Radiolarian protist as drawn by Haeckel Foraminiferans, and some marine amoebae, ciliates and flagellates.
Fungal-like
Slime moulds
and
slime nets
 Saprotrophic protists that get their food from the remains of organisms that have broken down and decayed   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, 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.[25]
External videos
  How microscopic hunters get their lunch
  Euglenoids: Single-celled shapeshifters
  How do protozoans get around?
  • Ciliate ingesting a diatom
  • Amoeba engulfing a diatom

The fungus-like protist saprobes are specialized to absorb nutrients from nonliving organic matter, such as dead organisms or their wastes. For instance, many types of oomycetes grow on dead animals or algae. Marine saprobic protists have the essential function of returning inorganic nutrients to the water. This process allows for new algal growth, which in turn generates sustenance for other organisms along the food chain. Indeed, without saprobe species, such as protists, fungi, and bacteria, life would cease to exist as all organic carbon became "tied up" in dead organisms.[27][28]

Mixotrophs edit

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

Mixotrophs have no single trophic mode. 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.[29] 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.[30]

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.[31] They can take advantage of different environmental conditions.[32]

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.[33] 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.[34] As a result, Phaeocystis is an important contributor to the marine carbon[35] and sulfur cycles.[36] Phaeocystis species are endosymbionts to acantharian radiolarians.[37][38]

Mixotrophic plankton that combine phototrophy and heterotrophy – table based on Stoecker et al., 2017[39]
General types Description Example Further examples
Bacterioplankton Photoheterotrophic bacterioplankton   Vibrio cholerae Roseobacter spp.
Erythrobacter spp.
Gammaproteobacterial clade OM60
Widespread among bacteria and archaea
Phytoplankton Called constitutive mixotrophs by Mitra et al., 2016.[40] Phytoplankton that eat: photosynthetic protists with inherited plastids and the capacity to ingest prey.   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.[40] 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   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 Dinophysis spp.
Mesodinium rubrum
2. Protists or zooplankton with algal endosymbionts of only one algal species or very closely related algal species   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.

Protist locomotion edit

Main article: Protist 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   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.   Cryptophytes All dinoflagellates and nanoflagellates (choanoflagellates, silicoflagellates, most green algae)[41][42]
(Other protists go through a phase as gametes when they have temporary flagellum – some radiolarians, foraminiferans and Apicomplexa)
Ciliates   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.   Paramecium bursaria
click to see cilia 		Foraminiferans, and some marine amoebae, ciliates and flagellates.
Amoebas
(amoeboids) 		  Pseudopods (Greek for false feet) are lobe-like appendages which amoebas use to anchor to a solid surface and pull themselves forward. They can change their shape by extending and retracting these pseudopods.[43]
 
 Amoeba Found in every major protist lineage. Amoeboid cells occur among the protozoans, but also in the algae and the fungi.[44][45]
Not motile
none
 
 Diatom Coccolithophores, most diatoms, and non‐motile species of Phaeocystis[42] Among protozoans the parasitic Apicomplexa are non‐motile.
 
Difference of beating pattern of flagellum and cilium

Flagella are used in prokaryotes (archaea and bacteria) as well as protists. In addition, both flagella and cilia are widely used in eukaryotic cells (plant and animal) apart from protists.

The regular beat patterns of eukaryotic cilia and flagella generates motion on a cellular level. Examples range from the propulsion of single cells such as the swimming of spermatozoa to the transport of fluid along a stationary layer of cells such as in a respiratory tract. Though eukaryotic flagella and motile cilia are ultrastructurally identical, the beating pattern of the two organelles can be different. In the case of flagella, the motion is often planar and wave-like, whereas the motile cilia often perform a more complicated three-dimensional motion with a power and recovery stroke.

Eukaryotic flagella—those of animal, plant, and protist cells—are complex cellular projections that lash back and forth. Eukaryotic flagella are classed along with eukaryotic motile cilia as undulipodia[46] to emphasize their distinctive wavy appendage role in cellular function or motility. Primary cilia are immotile, and are not undulipodia.

 
Marine flagellates from the genera (left to right)
Cryptaulax, Abollifer, Bodo, Rhynchomonas, Kiitoksia, Allas, and Metromonas [47]
 
Cilia performs powerful forward strokes with a stiffened flagellum followed by relatively slow recovery movement with a relaxed flagellum

Ciliates generally have hundreds to thousands of cilia that are densely packed together in arrays. Like the flagella, the cilia are powered by specialised molecular motors. An efficient forward stroke is made with a stiffened flagellum, followed by an inefficient backward stroke made with a relaxed flagellum. During movement, an individual cilium deforms as it uses the high-friction power strokes and the low-friction recovery strokes. Since there are multiple cilia packed together on an individual organism, they display collective behaviour 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.[48]

External videos
  Paramecium: The White Rat of Ciliates

Marine algae edit

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.[49] 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,[50] mostly multicellular and including many notable seaweeds.[50][51] Brown algae form a class containing about 2,000 recognised species,[52] 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.[53] 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 marine protists 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.[55] 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.[56]

Diatoms edit

 
Diatoms come in many shapes and sizes

Diatoms are photosynthetic unicellular algae populating the oceans and other waters around the globe. They form a (disputed) phylum containing about 100,000 recognised species. Diatoms generate about 20 per cent of all oxygen produced on the planet each year,[26] and take in over 6.7 billion metric tons of silicon each year from the waters in which they live.[57] They produce 25–45% of the total primary production of organic material in the oceans,[58][59][60] owing to their prevalence in open-ocean regions when total phytoplankton biomass is maximal.[61][62]

Diatoms are enclosed in protective silica (glass) shells called frustules. They are classified by the shape of these glass cages in which they live, and which they build as they grow. Each frustule is made from two interlocking parts covered with tiny holes through which the diatom exchanges nutrients and wastes.[63] Dead diatoms drift to the ocean floor where, over millions of years, the remains of their frustules can build up as much as half a mile deep.[64] Diatoms have relatively high sinking speeds compared with other phytoplankton groups, and they account for about 40% of particulate carbon exported to ocean depths.[60][65][62]

  •  
    Diatoms are one of the most common types of phytoplankton
  •  
    Their protective shells (frustles) are made of silicon
  •  
  •  
Diatom shapes
 
 
          Drawings by Haeckel 1904 (click for details)
Diatoms
 
Centric
 
Pennate
Diatoms have a silica shell (frustule) with radial (centric) or bilateral (pennate) symmetry
External videos
  Diatoms: Tiny factories you can see from space
  Diatom 3D interference contrast
 
Structure of a centric diatom frustule[66]

Physically driven seasonal enrichments in surface nutrients favour diatom blooms. Anthropogenic climate change will directly affect these seasonal cycles, changing the timing of blooms and diminishing their biomass, which will reduce primary production and CO2 uptake.[67][62] Remote sensing data suggests there was a global decline of diatoms between 1998 and 2012, particularly in the North Pacific, associated with shallowing of the surface mixed layer and lower nutrient concentrations.[68][62]

  •  
    Silicified frustule of a pennate diatom with two overlapping halves
  •  
    Guinardia delicatula, a diatom responsible for diatom blooms in the North Sea [69]
  •  
    There are over 100,000 species of diatoms accounting for 25–45% of the ocean's primary production
  •  
    Linked diatoms
  •  
    Pennate diatom from an Arctic meltpond, infected with two chytrid-like fungal pathogens. Scale bar = 10 µm.[70]

Coccolithophores edit

Coccolithophores
 
...have plates called coccoliths
 
...extinct fossil
Coccolithophores build calcite skeletons important to the marine carbon cycle[71]

Coccolithophores are minute unicellular photosynthetic protists with two flagella for locomotion. Most of them are protected by calcium carbonate shells covered with ornate circular plates or scales called coccoliths. 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.[72]

 
Fossil of Coccolithus pelagicus, about 10 μm across
 
Diverse coccolithophores from the Maldives [73]
 
The fossil coccolithophore Braarudosphaera bigelowii has an unusual shell with a regular dodecahedral structure about 10 micrometers across.[74]

Dinoflagellates edit

See also: Mixotrophic dinoflagellate and Predatory dinoflagellate
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.[75] 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.[41]

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).[76] 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 Mesodinium rubrum, which sequester their chloroplasts from a specific cryptophyte clade (Geminigera/Plagioselmis/Teleaulax)".[39]

  •  
    Gyrodinium, one of the few naked dinoflagellates which lack armour
  •  
    The dinoflagellate Protoperidinium extrudes a large feeding veil to capture prey
  •  
    Nassellarian radiolarians can be in symbiosis with dinoflagellates
  •  
    The dinoflagellate Dinophysis acuta
 
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[77]
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.[78] 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.[79] There is evidence from DNA analysis that dinoflagellate symbiosis with radiolarians evolved independently from other dinoflagellate symbioses, such as with foraminifera.[80]

Some dinoflagellates are bioluminescent. At night, ocean water can light up internally and sparkle with blue light because of these dinoflagellates.[81][82] 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.[83]

Marine protozoans edit

Protozoans are protists which feed on organic matter such as other microorganisms or organic tissues and debris.[87][88] 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.[89][90] 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 edit

Radiolarian shapes
 
 
          Drawings by Haeckel 1904 (click for details)

Radiolarians are unicellular predatory protists encased in elaborate globular shells, typically between 0.1 and 0.2 millimetres in size, 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.[91]

  •  
    Like diatoms, radiolarians come in many shapes
  •  
    Also like diatoms, radiolarian shells are usually made of silicate
  •  
    However acantharian radiolarians have shells made from strontium sulfate crystals
  •  
    Cutaway schematic diagram of a spherical radiolarian shell
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[92]
External videos
  Radiolarian geometry
  Ernst Haeckel's radiolarian engravings
  •  
    Cladococcus abietinus
  •  
    Cleveiplegma boreale

Foraminiferans edit

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.[93] They are widely researched with well established fossil records which allow scientists to infer a lot about past environments and climates.[91]

Foraminiferans
 
...can have more than one nucleus
 
...and defensive spines
Foraminiferans are important unicellular zooplankton protists, with calcium tests
External videos
  foraminiferans
  Foraminiferal networks and growth

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.[93] Mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters.[95] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.[96]

Amoeba edit

Shelled and naked amoeba
 
Testate amoeba, Cyphoderia sp.
 
Naked amoeba, Chaos sp.
                  Amoeba can be shelled (testate) or naked
  •  
    Naked amoeba showing food vacuoles and ingested diatom
  •  
    Shell or test of a testate amoeba, Arcella sp.
  •  
    Xenogenic testate amoeba covered in diatoms (from Penard's Amoeba Collection)
Amoeba engulfing a diatom
External videos
  Amoebas
  Testate amoebas
  Feeding amoebas

Ciliates edit

Ciliate shapes
 
          Drawings by Haeckel 1904 (click for details)

Marine ciliates are major grazers of the phytoplankton.[97][98]

Phytoplankton primary production supports higher trophic levels and fuels microbial remineralization.[99][100] The dominant pelagic grazers of phytoplankton are typically associated with distinct operating modes of the food web compartments and nutrient cycling. Heterotrophic protist grazers and microzooplankton dominance is usually associated with the microbial loop and regenerated production; while mesozooplankton is associated with a linear food chain and export production.[101][102] Grazing on particulate primary production in the global ocean surface is ~10–15% for mesozooplankton and 59–75% for microzooplankton,[103][104][105][106] with estimates for coastal and estuarine systems usually in the a lower range.[106][98]

Ciliates constitute an important component of the microzooplankton community with preference for small-sized preys, in contrast to mesozooplankton, and many ciliate species are also grazed by mesozooplankton.[107] Thus, ciliates can be an important link between small cells and higher trophic levels.[108] Besides their significant role in carbon transfer, ciliates are also considered high quality food, as a source of proteinaceous compounds with a low C:N ratio in comparison to phytoplankton.[109][110][98]

 
Conjugation of two Coleps sp.
Two similar-looking but sexually distinct partners connected at their front ends exchange genetic material via a plasma bridge.

Although many ciliates are heterotrophs, a number of pelagic species are mixotrophic, combining both phagotrophic and phototrophic nutrition (Stoecker, 1998). The recognition of mixotrophy in the marine plankton food web has challenged the classical understanding of pelagic food webs, as autotrophy and heterotrophy are not necessarily two distinct functional compartments.[111] Classical understanding of ecological interactions among plankton, such as competition for nutrients, indicates that nutrient uptake affinity decreases with organism size,[112] favoring smaller sizes under resource limiting conditions. Mixotrophy is advantageous to organisms under nutrient limited conditions, allowing them to reduce direct competition by grazing on smaller prey and increase direct ingestion of nutrients.[113] Modeling results suggest that mixotrophy favors larger organisms, and therefore enhances trophic transfer efficiency.[113][114] On top of that, mixotrophy appears to be important over both, space and time, in marine systems.[115] stressing the need for ecological field studies to further elucidate the role of mixotrophy.[98]

  • Several taxa of ciliates interacting
  • Blepharisma americanum swimming in a drop of pond water with other microorganisms
External videos
  Peritrich Ciliates
  Conjugating protists

Macroscopic protists edit

Planktonic protists edit

Interactome edit

 
Yellow-brown zooxanthellae, a photosynthetic algae that lives inside hosts like radiolarians and coral
 
Planktonic protist interactome [118]
Bipartite networks, providing an overview of the interactions represented by a manually curated Protist Interaction DAtabase (PIDA).

Interaction between microbial species has played important roles in evolution and speciation.[118] One of the best examples is that the origin of eukaryotes is grounded in the interaction-events of endosymbiosis; giving rise to mitochondria, chloroplasts, and other metabolic capacities in the eukaryotic cell,[119][120][121][122] Microbial interactions guarantee ecosystem function, having crucial roles in, for instance, carbon channeling in photosymbiosis, control of microalgae blooms by parasites, and phytoplankton-associated bacteria influencing the growth and health of their host.[118]

Despite their importance, understanding of microbial interactions in the ocean and other aquatic systems is rudimentary, and the majority of them are still unknown.[13][123][124][125] The earliest surveys of interactions between aquatic microbes date back to the 19th century. In 1851, while on board HMS Rattlesnake in the Pacific Ocean, Thomas Huxley discovered small yellow–green cells inside the conspicuous planktonic radiolarians which he thought were organelles/[126] Later, Karl Brandt established the yellowish cells were symbiotic alga and named them zooxanthella.[127] Since these early studies, hundreds of others have reported microbial interactions by using classic tools, mainly microscopy, but this knowledge has not yet been gathered into one accessible database. In recent years the high throughput sequencing (HTS)[128][129][130] of environmental DNA or RNA has transformed understanding of microbial diversity [131] and evolution,[132] as well as generating hypotheses on microbial interactions based on correlations of estimated microbial abundances over spatiotemporal scales.[133][134][135][136][118]

The diagram on the right is an overview of the interactions between planktonic protists recorded in a manually curated Protist Interaction DAtabase (PIDA). The network is based on 2422 ecological interactions in the PIDA registered from ~500 publications spanning the last 150 years. The nomenclature and taxonomic order of Eukaryota is based on Adl et al. 2019.[137] The nomenclature and taxonomic order of Bacteria is based on Schultz et al. 2017.[138][118]

The nodes are grouped (outer circle) according to eukaryotic supergroups (or Incertae sedis), Bacteria and Archaea. All major protistan lineages were involved in interactions as hosts, symbionts (mutualists and commensalists), parasites, predators, and/or prey. Predation was the most common interaction (39%), followed by symbiosis (29%), parasitism (18%), and unresolved interactions (14%, where it is uncertain whether the interaction is beneficial or antagonistic). Nodes represent eukaryotic and prokaryotic taxa and are colored accordingly. Node size indicates the number of edges/links that are connected to that node. Each node/taxon is assigned a number, which corresponds with the numbers for taxa in B, C and D. Edges represent interactions between two taxa and are colored according to ecological interaction type: predation (orange), symbiosis (green), and parasitism (purple).[118]

The network is undirected, meaning that a node can contain both parasites/symbionts/prey and hosts/predators. To avoid cluttering of the figure, "Self-loops", which represent cases where both interacting organisms belong to the same taxon (e.g., a dinoflagellate eating another dinoflagellate) are not shown as edges/links in this figure, but are considered in the size of nodes. The outermost circle groups taxa in the different eukaryotic ‘supergroups’ or the prokaryotic domains Bacteria and Archaea. Ancryomonadidae is abbreviated An. Telonema is not placed into any of the supergroups, but classified as Incertae sedis (abbreviated I.S. in the figure). In B, B, and D the following abbreviations for supergroups are used: Ar Archaea, Ba Bacteria, Rh Rhizaria, Al Alveolata, St Stramenopiles, Ha Haptista, Cy Cryptista, Ap Archaeplastida, Ex Excavata, Ob Obazoa, Am Amoebozoa, Cu CRuMS, An Ancryomonadidae, Is Incertae sedis.[118]

B: Predator–prey interactions in PIDA. The node numbers correspond to taxa node numbers in a. Abbreviations for supergroups are described above. Background and nodes are colored according to functional role in the interaction: Prey are colored light orange (left part of figure), while predators are depicted in dark orange (right part of figure). The size of each node represents the number of edges connected to that node.[118]

C. Symbiont–host interactions included in PIDA. The node numbers correspond to node numbers in A. Abbreviations for supergroups are described above. Symbionts are to the left, colored light green, and their hosts are to the right in dark green. The size of each node represents the number of edges connected to that node.[118]

D: Parasite–host interactions included in PIDA. The node numbers correspond to node numbers in A. Abbreviations for supergroups are described above. Parasite taxa are depicted in light purple (left), hosts in dark purple (right).[118]

It was found that protist predators seem to be "multivorous" while parasite–host and symbiont–host interactions appear to have moderate degrees of specialization. The SAR supergroup (i.e., Stramenopiles, Alveolata, and Rhizaria) heavily dominated PIDA, and comparisons against a global-ocean molecular survey (Tara expedition) indicated that several SAR lineages, which are abundant and diverse in the marine realm, were underrepresented among the recorded interactions.[118]

Protist shells edit

Main article: Protist shell
See also: Protists in the fossil record

Many protists have protective shells or tests,[139] usually made from calcium carbonate (chalk) or silica (glass). Protists are mostly single-celled and microscopic. Their shells are often tough mineralised forms that resist degradation, and can survive the death of the protist as a microfossil. Although protists are very small, they are ubiquitous. Their numbers are such that their shells play a huge part in the formation of ocean sediments, and in the global cycling of elements and nutrients.

Diatom shells are called frustules and are made from silica. These glass structures have accumulated for over 100 million years leaving rich deposits of nano and microstructured silicon oxide in the form of diatomaceous earth around the globe. The evolutionary causes for the generation of nano and microstructured silica by photosynthetic algae are not yet clear. However, in 2018 it was shown that reflection of ultraviolet light by nanostructured silica protects the DNA in the algal cells, and this may be an evolutionary cause for the formation of the glass cages.[140][141]

Coccolithophores are protected by a shell constructed from ornate circular plates or scales called coccoliths. The coccoliths are made from calcium carbonate or chalk. The term coccolithophore derives from the Greek for a seed carrying stone, referring to their small size and the coccolith stones they carry.[72]

Diatoms
 
Diatoms, major components of marine plankton, have glass skeletons called frustules. "The microscopic structures of diatoms help them manipulate light, leading to hopes they could be used in new technologies for light detection, computing or robotics.[142]
 
SEM images of pores in diatom frustules[140]
Coccolithophores
 
Coccolithophores are armoured with chalk plates or stones called coccoliths. The images above show the size comparison between the relatively large coccolithophore Scyphosphaera apsteinii and the relatively small but ubiquitous Emiliania huxleyi[143]
Benefits of having shells
 
Benefits in coccolithophore calcification [144] – see text below
Costs of having shells
 
Energetic costs in coccolithophore calcification [144]

There are benefits for protists that carry protective shells. The diagram on the left above shows some benefits coccolithophore get from carrying coccoliths. In the diagram, (A) represents accelerated photosynthesis including carbon concentrating mechanisms (CCM) and enhanced light uptake via scattering of scarce photons for deep-dwelling species. (B) represents protection from photodamage including sunshade protection from ultraviolet light (UV) and photosynthetic active radiation (PAR) and energy dissipation under high-light conditions. (C) represents armour protection includes protection against viral/bacterial infections and grazing by selective and nonselective grazers.[144]

There are also costs for protists that carry protective shells. The diagram on the right above shows some of the energetic costs coccolithophore incur from carrying coccoliths. In the diagram, the energetic costs are reported in percentage of total photosynthetic budget. (A) represents transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates Ca2+ and HCO3− (black arrows) and the removal of the end product H+ from the cell (gray arrow). The transport of Ca2+ through the cytoplasm to the coccolith vesicle (CV) is the dominant cost associated with calcification. (B) represents metabolic processes include the synthesis of coccolith-associated polysaccharides (CAPs – gray rectangles) by the Golgi complex (white rectangles) that regulate the nucleation and geometry of CaCO3 crystals. The completed coccolith (gray plate) is a complex structure of intricately arranged CAPs and CaCO3 crystals. (C) Mechanical and structural processes account for the secretion of the completed coccoliths that are transported from their original position adjacent to the nucleus to the cell periphery, where they are transferred to the surface of the cell.[144]

See also edit

References edit

  1. ^ Cavalier-Smith T (December 1993). "Kingdom protozoa and its 18 phyla". Microbiological Reviews. 57 (4): 953–94. doi:10.1128/mmbr.57.4.953-994.1993. PMC 372943. PMID 8302218.
  2. ^ Corliss JO (1992). "Should there be a separate code of nomenclature for the protists?". BioSystems. 28 (1–3): 1–14. doi:10.1016/0303-2647(92)90003-H. PMID 1292654.
  3. ^ Slapeta J, Moreira D, López-García P (2005). "The extent of protist diversity: insights from molecular ecology of freshwater eukaryotes". Proceedings of the Royal Society B: Biological Sciences. 272 (1576): 2073–81. doi:10.1098/rspb.2005.3195. PMC 1559898. PMID 16191619.
  4. ^ Moreira D, López-García P (2002). "The molecular ecology of microbial eukaryotes unveils a hidden world" (PDF). Trends in Microbiology. 10 (1): 31–8. doi:10.1016/S0966-842X(01)02257-0. PMID 11755083.
  5. ^ a b c Leles, S.G.; Mitra, A.; Flynn, K.J.; Stoecker, D.K.; Hansen, P.J.; Calbet, A.; McManus, G.B.; Sanders, R.W.; Caron, D.A.; Not, F.; Hallegraeff, G.M. (2017). "Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance". Proceedings of the Royal Society B: Biological Sciences. 284 (1860): 20170664. doi:10.1098/rspb.2017.0664. PMC 5563798. PMID 28768886.
  6. ^ Characteristics of Protists In: Rye, Connie; Avissar, Yael; Choi, Jung Ho; DeSaix, Jean; Jurukovski, Vladimir; Wise, Robert R. (2013). Biology. Houston, Texas. ISBN 978-1-938168-09-3. OCLC 896421272.{{cite book}}: CS1 maint: location missing publisher (link)   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  7. ^ Devreotes P (1989). "Dictyostelium discoideum: a model system for cell-cell interactions in development". Science. 245 (4922): 1054–8. Bibcode:1989Sci...245.1054D. doi:10.1126/science.2672337. PMID 2672337.
  8. ^ Neil A C, Reece J B, Simon E J (2004) Essential biology with physiology Pearson/Benjamin Cummings, Page 291. ISBN 9780805375039
  9. ^ O'Malley MA, Simpson AG, Roger AJ (2012). "The other eukaryotes in light of evolutionary protistology". Biology & Philosophy. 28 (2): 299–330. doi:10.1007/s10539-012-9354-y. S2CID 85406712.
  10. ^ Adl SM, Simpson AG, Farmer MA, Andersen RA, Anderson OR, Barta JR, Bowser SS, Brugerolle G, Fensome RA, Fredericq S, James TY, Karpov S, Kugrens P, Krug J, Lane CE, Lewis LA, Lodge J, Lynn DH, Mann DG, McCourt RM, Mendoza L, Moestrup O, Mozley-Standridge SE, Nerad TA, Shearer CA, Smirnov AV, Spiegel FW, Taylor MF (2005). "The new higher level classification of eukaryotes with emphasis on the taxonomy of protists". The Journal of Eukaryotic Microbiology. 52 (5): 399–451. doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873. S2CID 8060916.
  11. ^ Margulis L, Chapman MJ (19 March 2009). Kingdoms and Domains: An Illustrated Guide to the Phyla of Life on Earth. Academic Press. ISBN 9780080920146.
  12. ^ a b Collier, Jackie L.; Rest, Joshua S. (2019). "Swimming, gliding, and rolling toward the mainstream: Cell biology of marine protists". Molecular Biology of the Cell. 30 (11): 1245–1248. doi:10.1091/mbc.E18-11-0724. PMC 6724603. PMID 31084566.
  13. ^ a b Worden, A. Z.; Follows, M. J.; Giovannoni, S. J.; Wilken, S.; Zimmerman, A. E.; Keeling, P. J. (2015). "Rethinking the marine carbon cycle: Factoring in the multifarious lifestyles of microbes". Science. 347 (6223). doi:10.1126/science.1257594. PMID 25678667. S2CID 206560125.
  14. ^ De Vargas, C.; et al. (2015). "Eukaryotic plankton diversity in the sunlit ocean". Science. 348 (6237). doi:10.1126/science.1261605. hdl:10261/117736. PMID 25999516. S2CID 12853481.
  15. ^ a b Curtis, Bruce A.; et al. (2012). "Algal genomes reveal evolutionary mosaicism and the fate of nucleomorphs". Nature. 492 (7427): 59–65. Bibcode:2012Natur.492...59C. doi:10.1038/nature11681. PMID 23201678. S2CID 4380094.
  16. ^ a b Armbrust, E. V.; et al. (2004). "The Genome of the Diatom Thalassiosira Pseudonana: Ecology, Evolution, and Metabolism". Science. 306 (5693): 79–86. Bibcode:2004Sci...306...79A. doi:10.1126/science.1101156. PMID 15459382. S2CID 8593895.
  17. ^ a b Read, Betsy A.; et al. (2013). "Pan genome of the phytoplankton Emiliania underpins its global distribution". Nature. 499 (7457): 209–213. Bibcode:2013Natur.499..209.. doi:10.1038/nature12221. hdl:1854/LU-4120924. PMID 23760476. S2CID 4428297.
  18. ^ a b Keeling, Patrick J.; et al. (2014). "The Marine Microbial Eukaryote Transcriptome Sequencing Project (MMETSP): Illuminating the Functional Diversity of Eukaryotic Life in the Oceans through Transcriptome Sequencing". PLOS Biology. 12 (6): e1001889. doi:10.1371/journal.pbio.1001889. PMC 4068987. PMID 24959919.
  19. ^ Nymark, Marianne; Sharma, Amit Kumar; Sparstad, Torfinn; Bones, Atle M.; Winge, Per (2016). "A CRISPR/Cas9 system adapted for gene editing in marine algae". Scientific Reports. 6: 24951. Bibcode:2016NatSR...624951N. doi:10.1038/srep24951. PMC 4842962. PMID 27108533.
  20. ^ Hopes, Amanda; Nekrasov, Vladimir; Kamoun, Sophien; Mock, Thomas (2016). "Editing of the urease gene by CRISPR-Cas in the diatom Thalassiosira pseudonana". Plant Methods. 12: 49. doi:10.1186/s13007-016-0148-0. PMC 5121945. PMID 27904648.
  21. ^ Carradec, Quentin; et al. (2018). "A global ocean atlas of eukaryotic genes". Nature Communications. 9 (1): 373. Bibcode:2018NatCo...9..373C. doi:10.1038/s41467-017-02342-1. PMC 5785536. PMID 29371626.
  22. ^ a b Faktorová, Drahomíra; et al. (2020). "Genetic tool development in marine protists: Emerging model organisms for experimental cell biology". Nature Methods. 17 (5): 481–494. doi:10.1038/s41592-020-0796-x. PMC 7200600. PMID 32251396.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  23. ^ Whittaker, R.H.; Margulis, L. (1978). "Protist classification and the kingdoms of organisms". Biosystems. 10 (1–2): 3–18. doi:10.1016/0303-2647(78)90023-0. PMID 418827.
  24. ^ Faure, E; Not, F; Benoiston, AS; Labadie, K; Bittner, L; Ayata, SD (2019). "Mixotrophic protists display contrasted biogeographies in the global ocean". ISME Journal. 13 (4): 1072–1083. doi:10.1038/s41396-018-0340-5. PMC 6461780. PMID 30643201.
  25. ^ Budd, Graham E; Jensen, Sören (2017). "The origin of the animals and a 'Savannah' hypothesis for early bilaterian evolution". Biological Reviews. 92 (1): 446–473. doi:10.1111/brv.12239. PMID 26588818.
  26. ^ a b The Air You're Breathing? A Diatom Made That
  27. ^ Clark M A, Douglas M and Choi J (2018) Biology 2e, 23.4 "Ecology of Protists", OpenStax, Houston, Texas.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  28. ^ Vallet, Marine; Baumeister, Tim U. H.; Kaftan, Filip; Grabe, Veit; Buaya, Anthony; Thines, Marco; Svatoš, Aleš; Pohnert, Georg (2019). "The oomycete Lagenisma coscinodisci hijacks host alkaloid synthesis during infection of a marine diatom". Nature Communications. 10 (1): 4938. Bibcode:2019NatCo..10.4938V. doi:10.1038/s41467-019-12908-w. PMC 6821873. PMID 31666506.
  29. ^ Beware the mixotrophs - they can destroy entire ecosystems 'in a matter of hours'
  30. ^ Microscopic body snatchers infest our oceans - Phys.org
  31. ^ Eiler A (December 2006). "Evidence for the Ubiquity of Mixotrophic Bacteria in the Upper Ocean: Implications and Consequences". Appl Environ Microbiol. 72 (12): 7431–7. Bibcode:2006ApEnM..72.7431E. doi:10.1128/AEM.01559-06. PMC 1694265. PMID 17028233.
  32. ^ Katechakis A, Stibor H (July 2006). "The mixotroph Ochromonas tuberculata may invade and suppress specialist phago- and phototroph plankton communities depending on nutrient conditions". Oecologia. 148 (4): 692–701. Bibcode:2006Oecol.148..692K. doi:10.1007/s00442-006-0413-4. PMID 16568278. S2CID 22837754.
  33. ^ Schoemann, Véronique; Becquevort, Sylvie; Stefels, Jacqueline; Rousseau, Véronique; Lancelot, Christiane (1 January 2005). "Phaeocystis blooms in the global ocean and their controlling mechanisms: a review". Journal of Sea Research. Iron Resources and Oceanic Nutrients - Advancement of Global Environmental Simulations. 53 (1–2): 43–66. Bibcode:2005JSR....53...43S. CiteSeerX 10.1.1.319.9563. doi:10.1016/j.seares.2004.01.008.
  34. ^ "Welcome to the Phaeocystis antarctica genome sequencing project homepage".
  35. ^ DiTullio, G. R.; Grebmeier, J. M.; Arrigo, K. R.; Lizotte, M. P.; Robinson, D. H.; Leventer, A.; Barry, J. P.; VanWoert, M. L.; Dunbar, R. B. (2000). "Rapid and early export of Phaeocystis antarctica blooms in the Ross Sea, Antarctica". Nature. 404 (6778): 595–598. Bibcode:2000Natur.404..595D. doi:10.1038/35007061. PMID 10766240. S2CID 4409009.
  36. ^ J, Stefels; L, Dijkhuizen; WWC, Gieskes (20 July 1995). "DMSP-lyase activity in a spring phytoplankton bloom off the Dutch coast, related to Phaeocystis sp. abundance" (PDF). Marine Ecology Progress Series. 123: 235–243. Bibcode:1995MEPS..123..235S. doi:10.3354/meps123235.
  37. ^ Decelle, Johan; Simó, Rafel; Galí, Martí; Vargas, Colomban de; Colin, Sébastien; Desdevises, Yves; Bittner, Lucie; Probert, Ian; Not, Fabrice (30 October 2012). "An original mode of symbiosis in open ocean plankton". Proceedings of the National Academy of Sciences. 109 (44): 18000–18005. Bibcode:2012PNAS..10918000D. doi:10.1073/pnas.1212303109. ISSN 0027-8424. PMC 3497740. PMID 23071304.
  38. ^ Mars Brisbin, Margaret; Grossmann, Mary M.; Mesrop, Lisa Y.; Mitarai, Satoshi (2018). "Intra-host Symbiont Diversity and Extended Symbiont Maintenance in Photosymbiotic Acantharea (Clade F)". Frontiers in Microbiology. 9: 1998. doi:10.3389/fmicb.2018.01998. ISSN 1664-302X. PMC 6120437. PMID 30210473.
  39. ^ a b Stoecker, D.K.; Hansen, P.J.; Caron, D.A.; Mitra, A. (2017). "Mixotrophy in the marine plankton". Annual Review of Marine Science. 9: 311–335. Bibcode:2017ARMS....9..311S. doi:10.1146/annurev-marine-010816-060617. PMID 27483121.
  40. ^ a b Mitra, A; Flynn, KJ; Tillmann, U; Raven, J; Caron, D; et al. (2016). "Defining planktonic protist functional groups on mechanisms for energy and nutrient acquisition; incorporation of diverse mixotrophic strategies". Protist. 167 (2): 106–20. doi:10.1016/j.protis.2016.01.003. hdl:10261/131722. PMID 26927496.
  41. ^ a b Dawson, Scott C; Paredez, Alexander R (2013). "Alternative cytoskeletal landscapes: cytoskeletal novelty and evolution in basal excavate protists". Current Opinion in Cell Biology. 25 (1): 134–141. doi:10.1016/j.ceb.2012.11.005. PMC 4927265. PMID 23312067.
  42. ^ a b Atkinson, A.; Polimene, L.; Fileman, E.S.; Widdicombe, C.E.; McEvoy, A.J.; Smyth, T.J.; Djeghri, N.; Sailley, S.F.; Cornwell, L.E. (2018). ""Comment. What drives plankton seasonality in a stratifying shelf sea? Some competing and complementary theories"]" (PDF). Limnology and Oceanography. 63 (6): 2877–2884. Bibcode:2018LimOc..63.2877A. doi:10.1002/lno.11036. S2CID 91380765.
  43. ^ Singleton, Paul (2006). Dictionary of Microbiology and Molecular Biology, 3rd Edition, revised. Chichester, UK: John Wiley & Sons. pp. 32. ISBN 978-0-470-03545-0.
  44. ^ David J. Patterson. "Amoebae: Protists Which Move and Feed Using Pseudopodia". Tree of Life web project.
  45. ^ . The University of Edinburgh. Archived from the original on 10 June 2009.
  46. ^ A Dictionary of Biology, 2004, accessed 2011-01-01.
  47. ^ Patterson, David J. (2000) "Flagellates: Heterotrophic Protists With Flagella" Tree of Life.
  48. ^ Lauga, Eric; Thomas R Powers (25 August 2009). "The hydrodynamics of swimming microorganisms". Reports on Progress in Physics. 72 (9): 096601. arXiv:0812.2887. Bibcode:2009RPPh...72i6601L. doi:10.1088/0034-4885/72/9/096601. S2CID 3932471.
  49. ^ Guiry MD (October 2012). "How many species of algae are there?". Journal of Phycology. 48 (5): 1057–63. doi:10.1111/j.1529-8817.2012.01222.x. PMID 27011267. S2CID 30911529.
  50. ^ a b Guiry, M.D.; Guiry, G.M. (2016). "Algaebase". www.algaebase.org. Retrieved 20 November 2016.
  51. ^ D. Thomas (2002). Seaweeds. Life Series. Natural History Museum, London. ISBN 978-0-565-09175-0.
  52. ^ Hoek, Christiaan; den Hoeck, Hoeck Van; Mann, David; Jahns, H.M. (1995). Algae : an introduction to phycology. Cambridge University Press. p. 166. ISBN 9780521316873. OCLC 443576944.
  53. ^ Starckx, Senne (31 October 2012) A place in the sun - Algae is the crop of the future, according to researchers in Geel 7 November 2017 at the Wayback Machine Flanders Today, Retrieved 8 December 2012
  54. ^ Duval, B.; Margulis, L. (1995). "The microbial community of Ophrydium versatile colonies: endosymbionts, residents, and tenants". Symbiosis. 18: 181–210. PMID 11539474.
  55. ^ Mandoli, DF (1998). "Elaboration of Body Plan and Phase Change during Development of Acetabularia: How Is the Complex Architecture of a Giant Unicell Built?". Annual Review of Plant Physiology and Plant Molecular Biology. 49: 173–198. doi:10.1146/annurev.arplant.49.1.173. PMID 15012232. S2CID 6241264.
  56. ^ Pierre Madl; Maricela Yip (2004). . BUFUS-Info. 19 (31). Archived from the original on 8 October 2022. Retrieved 12 May 2020.
  57. ^ Treguer, P.; Nelson, D. M.; Van Bennekom, A. J.; Demaster, D. J.; Leynaert, A.; Queguiner, B. (1995). "The Silica Balance in the World Ocean: A Reestimate". Science. 268 (5209): 375–9. Bibcode:1995Sci...268..375T. doi:10.1126/science.268.5209.375. PMID 17746543. S2CID 5672525.
  58. ^ Nelson, David M.; Tréguer, Paul; Brzezinski, Mark A.; Leynaert, Aude; Quéguiner, Bernard (1995). "Production and dissolution of biogenic silica in the ocean: Revised global estimates, comparison with regional data and relationship to biogenic sedimentation". Global Biogeochemical Cycles. 9 (3): 359–372. Bibcode:1995GBioC...9..359N. doi:10.1029/95GB01070.
  59. ^ Malviya, Shruti; Scalco, Eleonora; Audic, Stéphane; Vincent, Flora; Veluchamy, Alaguraj; Poulain, Julie; Wincker, Patrick; Iudicone, Daniele; De Vargas, Colomban; Bittner, Lucie; Zingone, Adriana; Bowler, Chris (2016). "Insights into global diatom distribution and diversity in the world's ocean". Proceedings of the National Academy of Sciences. 113 (11): E1516–E1525. Bibcode:2016PNAS..113E1516M. doi:10.1073/pnas.1509523113. PMC 4801293. PMID 26929361. S2CID 22035749.
  60. ^ a b Tréguer, Paul; Bowler, Chris; Moriceau, Brivaela; Dutkiewicz, Stephanie; Gehlen, Marion; Aumont, Olivier; Bittner, Lucie; Dugdale, Richard; Finkel, Zoe; Iudicone, Daniele; Jahn, Oliver; Guidi, Lionel; Lasbleiz, Marine; Leblanc, Karine; Levy, Marina; Pondaven, Philippe (2018). "Influence of diatom diversity on the ocean biological carbon pump" (PDF). Nature Geoscience. 11 (1): 27–37. Bibcode:2018NatGe..11...27T. doi:10.1038/s41561-017-0028-x. S2CID 134885922.
  61. ^ Mahadevan, Amala; d'Asaro, Eric; Lee, Craig; Perry, Mary Jane (2012). "Eddy-Driven Stratification Initiates North Atlantic Spring Phytoplankton Blooms". Science. 337 (6090): 54–58. Bibcode:2012Sci...337...54M. doi:10.1126/science.1218740. PMID 22767922. S2CID 42312402.
  62. ^ a b c d Cavicchioli, Ricardo; Ripple, William J.; Timmis, Kenneth N.; Azam, Farooq; Bakken, Lars R.; Baylis, Matthew; Behrenfeld, Michael J.; Boetius, Antje; Boyd, Philip W.; Classen, Aimée T.; Crowther, Thomas W.; Danovaro, Roberto; Foreman, Christine M.; Huisman, Jef; Hutchins, David A.; Jansson, Janet K.; Karl, David M.; Koskella, Britt; Mark Welch, David B.; Martiny, Jennifer B. H.; Moran, Mary Ann; Orphan, Victoria J.; Reay, David S.; Remais, Justin V.; Rich, Virginia I.; Singh, Brajesh K.; Stein, Lisa Y.; Stewart, Frank J.; Sullivan, Matthew B.; et al. (2019). "Scientists' warning to humanity: Microorganisms and climate change". Nature Reviews Microbiology. 17 (9): 569–586. doi:10.1038/s41579-019-0222-5. PMC 7136171. PMID 31213707.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  63. ^ Wassilieff, Maggy (2006) "Plankton - Plant plankton", Te Ara - the Encyclopedia of New Zealand. Accessed: 2 November 2019.
  64. ^ "King's College London - Lake Megachad". www.kcl.ac.uk. Retrieved 5 May 2018.
  65. ^ Boyd, Philip W.; Claustre, Hervé; Levy, Marina; Siegel, David A.; Weber, Thomas (2019). "Multi-faceted particle pumps drive carbon sequestration in the ocean" (PDF). Nature. 568 (7752): 327–335. Bibcode:2019Natur.568..327B. doi:10.1038/s41586-019-1098-2. PMID 30996317. S2CID 119513489.
  66. ^ Zhang, D.; Wang, Y.; Cai, J.; Pan, J.; Jiang, X.; Jiang, Y. (2012). "Bio-manufacturing technology based on diatom micro- and nanostructure". Chinese Science Bulletin. 57 (30): 3836–3849. Bibcode:2012ChSBu..57.3836Z. doi:10.1007/s11434-012-5410-x.
  67. ^ Behrenfeld, Michael J.; Doney, Scott C.; Lima, Ivan; Boss, Emmanuel S.; Siegel, David A. (2013). "Annual cycles of ecological disturbance and recovery underlying the subarctic Atlantic spring plankton bloom". Global Biogeochemical Cycles. 27 (2): 526–540. Bibcode:2013GBioC..27..526B. doi:10.1002/gbc.20050. hdl:1912/6250.
  68. ^ Rousseaux, Cecile S.; Gregg, Watson W. (2015). "Recent decadal trends in global phytoplankton composition". Global Biogeochemical Cycles. 29 (10): 1674–1688. Bibcode:2015GBioC..29.1674R. doi:10.1002/2015GB005139.
  69. ^ Arsenieff, L.; Simon, N.; Rigaut-Jalabert, F.; Le Gall, F.; Chaffron, S.; Corre, E.; Com, E.; Bigeard, E.; Baudoux, A.C. (2018). "First Viruses Infecting the Marine Diatom Guinardia delicatula". Frontiers in Microbiology. 9: 3235. doi:10.3389/fmicb.2018.03235. PMC 6334475. PMID 30687251.
  70. ^ Kilias, Estelle S.; Junges, Leandro; Šupraha, Luka; Leonard, Guy; Metfies, Katja; Richards, Thomas A. (2020). "Chytrid fungi distribution and co-occurrence with diatoms correlate with sea ice melt in the Arctic Ocean". Communications Biology. 3 (1): 183. doi:10.1038/s42003-020-0891-7. PMC 7174370. PMID 32317738. S2CID 216033140.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  71. ^ Rost, B. and Riebesell, U. (2004) "Coccolithophores and the biological pump: responses to environmental changes". In: Coccolithophores: From Molecular Processes to Global Impact, pages 99–125, Springer. ISBN 9783662062784.
  72. ^ a b Wassilieff, Maggy (2006) "A coccolithophore", Te Ara - the Encyclopedia of New Zealand. Accessed: 2 November 2019.
  73. ^ Young, J.R.; Pratiwi, S.; Su, X. (2017). "Data report: Surface seawater plankton sampling for coccolithophores undertaken during IODP Expedition 359". Proceedings of the International Ocean Discovery Program. doi:10.14379/iodp.proc.359.111.2017. {{cite journal}}: Cite journal requires |journal= (help)
  74. ^ Hagino, K., Onuma, R., Kawachi, M. and Horiguchi, T. (2013) "Discovery of an endosymbiotic nitrogen-fixing cyanobacterium UCYN-A in Braarudosphaera bigelowii (Prymnesiophyceae)". PLoS One, 8(12): e81749. doi:10.1371/journal.pone.0081749.
  75. ^ Gómez F (2012). "A checklist and classification of living dinoflagellates (Dinoflagellata, Alveolata)". CICIMAR Oceánides. 27 (1): 65–140. doi:10.37543/oceanides.v27i1.111.
  76. ^ Stoecker DK (1999). "Mixotrophy among Dinoflagellates". The Journal of Eukaryotic Microbiology. 46 (4): 397–401. doi:10.1111/j.1550-7408.1999.tb04619.x. S2CID 83885629.
  77. ^ Suggested Explanation for Glowing Seas--Including Currently Glowing California Seas National Science Foundation, 18 October 2011.
  78. ^ Boltovskoy, Demetrio; Anderson, O. Roger; Correa, Nancy M. (2017). "Radiolaria and Phaeodaria". Handbook of the Protists. Springer, Cham. pp. 731–763. doi:10.1007/978-3-319-28149-0_19. ISBN 9783319281476.
  79. ^ Anderson, O. R. (1983). Radiolaria. Springer Science & Business Media.
  80. ^ Gast, R. J.; Caron, D. A. (1 November 1996). "Molecular phylogeny of symbiotic dinoflagellates from planktonic foraminifera and radiolaria". Molecular Biology and Evolution. 13 (9): 1192–1197. doi:10.1093/oxfordjournals.molbev.a025684. ISSN 0737-4038. PMID 8896371.
  81. ^ Castro P, Huber ME (2010). Marine Biology (8th ed.). McGraw Hill. pp. 95. ISBN 978-0071113021.
  82. ^ Hastings JW (1996). "Chemistries and colors of bioluminescent reactions: a review". Gene. 173 (1 Spec No): 5–11. doi:10.1016/0378-1119(95)00676-1. PMID 8707056.
  83. ^ Haddock SH, Moline MA, Case JF (2009). "Bioluminescence in the sea". Annual Review of Marine Science. 2: 443–93. Bibcode:2010ARMS....2..443H. doi:10.1146/annurev-marine-120308-081028. PMID 21141672. S2CID 3872860.
  84. ^ . The Merck Veterinary Manual. Archived from the original on 3 March 2016. Retrieved 4 November 2019.
  85. ^ Brand, Larry E.; Campbell, Lisa; Bresnan, Eileen (2012). "Karenia: The biology and ecology of a toxic genus". Harmful Algae. 14: 156–178. doi:10.1016/j.hal.2011.10.020. PMC 9891709. PMID 36733478.
  86. ^ Buskey, E.J. (1995). "Growth and bioluminescence of Noctiluca scintillans on varying algal diets". Journal of Plankton Research. 17 (1): 29–40. doi:10.1093/plankt/17.1.29.
  87. ^ Panno, Joseph (14 May 2014). The Cell: Evolution of the First Organism. Infobase Publishing. ISBN 9780816067367.
  88. ^ Bertrand, Jean-Claude; Caumette, Pierre; Lebaron, Philippe; Matheron, Robert; Normand, Philippe; Sime-Ngando, Télesphore (26 January 2015). Environmental Microbiology: Fundamentals and Applications: Microbial Ecology. Springer. ISBN 9789401791182.
  89. ^ Madigan, Michael T. (2012). Brock Biology of Microorganisms. Benjamin Cummings. ISBN 9780321649638.
  90. ^ Yaeger, Robert G. (1996). Protozoa: Structure, Classification, Growth, and Development. NCBI. ISBN 9780963117212. PMID 21413323. Retrieved 23 March 2018.
  91. ^ a b Wassilieff, Maggy (2006) "Plankton - Animal plankton", Te Ara - the Encyclopedia of New Zealand. Accessed: 2 November 2019.
  92. ^ Varea, C.; Aragon, J.L.; Barrio, R.A. (1999). "Turing patterns on a sphere". Physical Review E. 60 (4): 4588–92. Bibcode:1999PhRvE..60.4588V. doi:10.1103/PhysRevE.60.4588. PMID 11970318.
  93. ^ a b Hemleben, C.; Anderson, O.R.; Spindler, M. (1989). Modern Planktonic Foraminifera. Springer-Verlag. ISBN 978-3-540-96815-3.
  94. ^ Foraminifera: History of Study, University College London. Retrieved: 18 November 2019.
  95. ^ Advances in Microbial Ecology, Volum 11
  96. ^ Bernhard, J. M.; Bowser, S.M. (1999). "Benthic Foraminifera of dysoxic sediments: chloroplast sequestration and functional morphology". Earth-Science Reviews. 46 (1): 149–165. Bibcode:1999ESRv...46..149B. doi:10.1016/S0012-8252(99)00017-3.
  97. ^ Calbet, Albert; Landry, Michael R. (2004). "Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems". Limnology and Oceanography. 49 (1): 51–57. Bibcode:2004LimOc..49...51C. doi:10.4319/lo.2004.49.1.0051. hdl:10261/134985. S2CID 22995996.
  98. ^ a b c d Haraguchi, Lumi; Jakobsen, Hans H.; Lundholm, Nina; Carstensen, Jacob (2018). "Phytoplankton Community Dynamic: A Driver for Ciliate Trophic Strategies". Frontiers in Marine Science. 5. doi:10.3389/fmars.2018.00272. S2CID 51925344.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  99. ^ Azam, F.; Fenchel, T.; Field, JG; Gray, JS; Meyer-Reil, LA; Thingstad, F. (1983). "The Ecological Role of Water-Column Microbes in the Sea". Marine Ecology Progress Series. 10: 257–263. Bibcode:1983MEPS...10..257A. doi:10.3354/meps010257.
  100. ^ Sherr, Evelyn; Sherr, Barry (1988). "Role of microbes in pelagic food webs: A revised concept". Limnology and Oceanography. 33 (5): 1225–1227. Bibcode:1988LimOc..33.1225S. doi:10.4319/lo.1988.33.5.1225.
  101. ^ Fenchel, T. (1988). "Marine Plankton Food Chains". Annual Review of Ecology and Systematics. 19: 19–38. doi:10.1146/annurev.es.19.110188.000315.
  102. ^ Buitenhuis, Erik; Le Quéré, Corinne; Aumont, Olivier; Beaugrand, Grégory; Bunker, Adrian; Hirst, Andrew; Ikeda, Tsutomu; O'Brien, Todd; Piontkovski, Sergey; Straile, Dietmar (2006). "Biogeochemical fluxes through mesozooplankton". Global Biogeochemical Cycles. 20 (2): n/a. Bibcode:2006GBioC..20.2003B. doi:10.1029/2005GB002511. hdl:2115/13694.
  103. ^ Behrenfeld, Michael J.; Falkowski, Paul G. (1997). "Photosynthetic rates derived from satellite-based chlorophyll concentration". Limnology and Oceanography. 42 (1): 1–20. Bibcode:1997LimOc..42....1B. doi:10.4319/lo.1997.42.1.0001. S2CID 15857675.
  104. ^ Calbet, Albert (2001). "Mesozooplankton grazing effect on primary production: A global comparative analysis in marine ecosystems". Limnology and Oceanography. 46 (7): 1824–1830. Bibcode:2001LimOc..46.1824C. doi:10.4319/lo.2001.46.7.1824. hdl:10261/49263. S2CID 85461746.
  105. ^ Landry, Michael R.; Calbet, Albert (2004). "Microzooplankton production in the oceans". ICES Journal of Marine Science. 61 (4): 501–507. doi:10.1016/j.icesjms.2004.03.011.
  106. ^ a b Buitenhuis, Erik T.; Rivkin, Richard B.; Sailley, Sévrine; Le Quéré, Corinne (2010). "Biogeochemical fluxes through microzooplankton". Global Biogeochemical Cycles. 24 (4): n/a. Bibcode:2010GBioC..24.4015B. doi:10.1029/2009GB003601. S2CID 131413083.
  107. ^ Hansen, Per Juel; Bjørnsen, Peter Koefoed; Hansen, Benni Winding (2000). "Zooplankton grazing and growth: Scaling within the 2-2,000-µm body size range". Limnology and Oceanography. 45 (8): 1891. Bibcode:2000LimOc..45.1891H. doi:10.4319/lo.2000.45.8.1891.
  108. ^ Nielsen, Torkel Gissel; Kicrboe, Thomas (1994). "Regulation of zooplankton biomass and production in a temperate, coastal ecosystem. 2. Ciliates". Limnology and Oceanography. 39 (3): 508–519. Bibcode:1994LimOc..39..508N. doi:10.4319/lo.1994.39.3.0508.
  109. ^ Stoecker, Diane K.; Capuzzo, Judith Mcdowell (1990). "Predation on Protozoa: its importance to zooplankton". Journal of Plankton Research. 12 (5): 891–908. doi:10.1093/plankt/12.5.891.
  110. ^ Gifford, Dian J. (1991). "The Protozoan-Metazoan Trophic Link in Pelagic Ecosystems". The Journal of Protozoology. 38: 81–86. doi:10.1111/j.1550-7408.1991.tb04806.x.
  111. ^ Flynn, Kevin J.; Stoecker, Diane K.; Mitra, Aditee; Raven, John A.; Glibert, Patricia M.; Hansen, Per Juel; Granéli, Edna; Burkholder, Joann M. (2013). "Misuse of the phytoplankton–zooplankton dichotomy: The need to assign organisms as mixotrophs within plankton functional types". Journal of Plankton Research. 35: 3–11. doi:10.1093/plankt/fbs062.
  112. ^ Edwards, Kyle F.; Thomas, Mridul K.; Klausmeier, Christopher A.; Litchman, Elena (2012). "Allometric scaling and taxonomic variation in nutrient utilization traits and maximum growth rate of phytoplankton". Limnology and Oceanography. 57 (2): 554–566. Bibcode:2012LimOc..57..554E. doi:10.4319/lo.2012.57.2.0554. S2CID 13376583.
  113. ^ a b Mitra, A.; Flynn, K. J.; Burkholder, J. M.; Berge, T.; Calbet, A.; Raven, J. A.; Granéli, E.; Glibert, P. M.; Hansen, P. J.; Stoecker, D. K.; Thingstad, F.; Tillmann, U.; Våge, S.; Wilken, S.; Zubkov, M. V. (2014). "The role of mixotrophic protists in the biological carbon pump". Biogeosciences. 11 (4): 995–1005. Bibcode:2014BGeo...11..995M. doi:10.5194/bg-11-995-2014. hdl:10261/93693.
  114. ^ Ward, Ben A.; Follows, Michael J. (2016). "Marine mixotrophy increases trophic transfer efficiency, mean organism size, and vertical carbon flux". Proceedings of the National Academy of Sciences. 113 (11): 2958–2963. Bibcode:2016PNAS..113.2958W. doi:10.1073/pnas.1517118113. PMC 4801304. PMID 26831076.
  115. ^ Leles, S. G.; Mitra, A.; Flynn, K. J.; Stoecker, D. K.; Hansen, P. J.; Calbet, A.; McManus, G. B.; Sanders, R. W.; Caron, D. A.; Not, F.; Hallegraeff, G. M.; Pitta, P.; Raven, J. A.; Johnson, M. D.; Glibert, P. M.; Våge, S. (2017). "Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance". Proceedings of the Royal Society B: Biological Sciences. 284 (1860). doi:10.1098/rspb.2017.0664. PMC 5563798. PMID 28768886.
  116. ^ Matz, Mikhail V.; Tamara M. Frank; N. Justin Marshall; Edith A. Widder; Sonke Johnsen (9 December 2008). "Giant Deep-Sea Protist Produces Bilaterian-like Traces" (PDF). Current Biology. 18 (23). Elsevier Ltd: 1849–1854. doi:10.1016/j.cub.2008.10.028. PMID 19026540. S2CID 8819675.
  117. ^ Gooday, A. J.; Aranda da Silva, A.; Pawlowski, J. (1 December 2011). "Xenophyophores (Rhizaria, Foraminifera) from the Nazaré Canyon (Portuguese margin, NE Atlantic)". Deep-Sea Research Part II: Topical Studies in Oceanography. The Geology, Geochemistry, and Biology of Submarine Canyons West of Portugal. 58 (23–24): 2401–2419. Bibcode:2011DSRII..58.2401G. doi:10.1016/j.dsr2.2011.04.005.
  118. ^ a b c d e f g h i j k Bjorbækmo, Marit F. Markussen; Evenstad, Andreas; Røsæg, Line Lieblein; Krabberød, Anders K.; Logares, Ramiro (2020). "The planktonic protist interactome: where do we stand after a century of research?". The ISME Journal. 14 (2): 544–559. doi:10.1038/s41396-019-0542-5. PMC 6976576. PMID 31685936.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  119. ^ Margulis, Lynn; Fester, René (1991). Symbiosis as a Source of Evolutionary Innovation: Speciation and Morphogenesis. MIT Press. ISBN 9780262132695.
  120. ^ López-García, Purificación; Eme, Laura; Moreira, David (2017). "Symbiosis in eukaryotic evolution". Journal of Theoretical Biology. 434: 20–33. Bibcode:2017JThBi.434...20L. doi:10.1016/j.jtbi.2017.02.031. PMC 5638015. PMID 28254477.
  121. ^ Archibald, John M. (2015). "Endosymbiosis and Eukaryotic Cell Evolution". Current Biology. 25 (19): R911–R921. doi:10.1016/j.cub.2015.07.055. PMID 26439354. S2CID 16089231.
  122. ^ Cavalier-Smith, Thomas (2013). "Symbiogenesis: Mechanisms, Evolutionary Consequences, and Systematic Implications". Annual Review of Ecology, Evolution, and Systematics. 44: 145–172. doi:10.1146/annurev-ecolsys-110411-160320.
  123. ^ Mahé, Frédéric; De Vargas, Colomban; Bass, David; Czech, Lucas; Stamatakis, Alexandros; Lara, Enrique; Singer, David; Mayor, Jordan; Bunge, John; Sernaker, Sarah; Siemensmeyer, Tobias; Trautmann, Isabelle; Romac, Sarah; Berney, Cédric; Kozlov, Alexey; Mitchell, Edward A. D.; Seppey, Christophe V. W.; Egge, Elianne; Lentendu, Guillaume; Wirth, Rainer; Trueba, Gabriel; Dunthorn, Micah (2017). "Parasites dominate hyperdiverse soil protist communities in Neotropical rainforests". Nature Ecology & Evolution. 1 (4): 91. doi:10.1038/s41559-017-0091. PMID 28812652. S2CID 2631960.
  124. ^ Biard, Tristan; Stemmann, Lars; Picheral, Marc; Mayot, Nicolas; Vandromme, Pieter; Hauss, Helena; Gorsky, Gabriel; Guidi, Lionel; Kiko, Rainer; Not, Fabrice (2016). "In situ imaging reveals the biomass of giant protists in the global ocean" (PDF). Nature. 532 (7600): 504–507. Bibcode:2016Natur.532..504B. doi:10.1038/nature17652. PMID 27096373. S2CID 205248710.
  125. ^ Finlay, B.J.; Esteban, G.F. (1998). "Freshwater protozoa: Biodiversity and ecological function". Biodiversity and Conservation. 7 (9): 1163–1186. doi:10.1023/A:1008879616066. S2CID 10702795.
  126. ^ Huxley, Thomas H. (1851). "XXXIV.—Zoological notes and observations made on board H.M.S. Rattlesnake". Annals and Magazine of Natural History. 8 (48): 433–442. doi:10.1080/03745486109495002.
  127. ^ Brandt K. (1881) "Uber das Zusammenleben von Thieren und Algen". Verh Physiol Ges, 1: 524–527.
  128. ^ Logares, Ramiro; Haverkamp, Thomas H.A.; Kumar, Surendra; Lanzén, Anders; Nederbragt, Alexander J.; Quince, Christopher; Kauserud, Håvard (2012). "Environmental microbiology through the lens of high-throughput DNA sequencing: Synopsis of current platforms and bioinformatics approaches". Journal of Microbiological Methods. 91 (1): 106–113. doi:10.1016/j.mimet.2012.07.017. PMID 22849829.
  129. ^ Sogin, M. L.; Morrison, H. G.; Huber, J. A.; Welch, D. M.; Huse, S. M.; Neal, P. R.; Arrieta, J. M.; Herndl, G. J. (2006). "Microbial diversity in the deep sea and the underexplored "rare biosphere"". Proceedings of the National Academy of Sciences. 103 (32): 12115–12120. Bibcode:2006PNAS..10312115S. doi:10.1073/pnas.0605127103. PMC 1524930. PMID 16880384.
  130. ^ Goodwin, Sara; McPherson, John D.; McCombie, W. Richard (2016). "Coming of age: Ten years of next-generation sequencing technologies". Nature Reviews Genetics. 17 (6): 333–351. doi:10.1038/nrg.2016.49. PMC 10373632. PMID 27184599. S2CID 8295541.
  131. ^ Pedrós-Alió C, Acinas SG, Logares R, Massana R. Marine microbial diversity as seen by high throughput sequencing. In: Gasol, Josep M.; Kirchman, David L. (27 March 2018). Microbial Ecology of the Oceans. John Wiley & Sons. ISBN 9781119107187., pp. 47–87.
  132. ^ Spang, Anja; Saw, Jimmy H.; Jørgensen, Steffen L.; Zaremba-Niedzwiedzka, Katarzyna; Martijn, Joran; Lind, Anders E.; Van Eijk, Roel; Schleper, Christa; Guy, Lionel; Ettema, Thijs J. G. (2015). "Complex archaea that bridge the gap between prokaryotes and eukaryotes". Nature. 521 (7551): 173–179. Bibcode:2015Natur.521..173S. doi:10.1038/nature14447. PMC 4444528. PMID 25945739.
  133. ^ Faust, Karoline; Lahti, Leo; Gonze, Didier; De Vos, Willem M.; Raes, Jeroen (2015). "Metagenomics meets time series analysis: Unraveling microbial community dynamics". Current Opinion in Microbiology. 25: 56–66. doi:10.1016/j.mib.2015.04.004. PMID 26005845.
  134. ^ Faust, Karoline; Raes, Jeroen (2012). "Microbial interactions: From networks to models". Nature Reviews Microbiology. 10 (8): 538–550. doi:10.1038/nrmicro2832. PMID 22796884. S2CID 22872711.
  135. ^ Lima-Mendez, G.; et al. (2015). "Determinants of community structure in the global plankton interactome". Science. 348 (6237). doi:10.1126/science.1262073. hdl:10261/117702. PMID 25999517. S2CID 10326640.
  136. ^ Layeghifard, Mehdi; Hwang, David M.; Guttman, David S. (2017). "Disentangling Interactions in the Microbiome: A Network Perspective". Trends in Microbiology. 25 (3): 217–228. doi:10.1016/j.tim.2016.11.008. PMC 7172547. PMID 27916383.
  137. ^ Adl, Sina M.; et al. (2019). "Revisions to the classification, nomenclature, and diversity of eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. PMC 6492006. PMID 30257078.
  138. ^ Schulz, Frederik; Eloe-Fadrosh, Emiley A.; Bowers, Robert M.; Jarett, Jessica; Nielsen, Torben; Ivanova, Natalia N.; Kyrpides, Nikos C.; Woyke, Tanja (2017). "Towards a balanced view of the bacterial tree of life". Microbiome. 5 (1): 140. doi:10.1186/s40168-017-0360-9. PMC 5644168. PMID 29041958.
  139. ^ "Groups of Protists | Boundless Biology". courses.lumenlearning.com. Retrieved 16 February 2021.
  140. ^ a b Aguirre, L.E., Ouyang, L., Elfwing, A., Hedblom, M., Wulff, A. and Inganäs, O. (2018) "Diatom frustules protect DNA from ultraviolet light". Scientific reports, 8(1): 1–6. doi:10.1038/s41598-018-21810-2.   Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  141. ^ De Tommasi, E., Congestri, R., Dardano, P., De Luca, A.C., Managò, S., Rea, I. and De Stefano, M. (2018) "UV-shielding and wavelength conversion by centric diatom nanopatterned frustules". Scientific Repor

January 01, 1970
marine, protists, defined, their, habitat, protists, that, live, marine, environments, that, saltwater, seas, oceans, brackish, water, coastal, estuaries, life, originated, marine, single, celled, prokaryotes, bacteria, archaea, later, evolved, into, more, com. 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 Life originated as marine 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 Protists are the eukaryotes that cannot be classified as plants fungi or animals They are mostly single celled and microscopic 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 for all descendants Marine protistsAlga phytoplankton Protozoan zooplankton Protists are usually one celled microorganisms They include algae autotrophs which make their own food and protozoans heterotrophs which eat the algae for food In recent years researchers have discovered many protists are mixotrophs which can function in both modes Most protists are too small to be seen with the naked eye They are highly diverse organisms currently organised into 18 phyla but not easy to classify 1 2 Studies have shown high protist diversity exists in oceans deep sea vents and river sediments suggesting large numbers 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 protists contribute a significant part of the protist biomass 5 Since protists are eukaryotes and not prokaryotes they possess within their cell at least one nucleus as well as organelles such as mitochondria and Golgi bodies Many protist species can switch between asexual reproduction and sexual reproduction involving meiosis and fertilization 6 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 7 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 8 Some modern authors prefer to exclude multicellular organisms from the traditional definition of a protist restricting protists to unicellular organisms 9 10 This more constrained definition excludes all brown the multicellular red and green algae and sometimes slime molds slime molds excluded when multicellularity is defined as complex 11 Contents 1 Background 2 Trophic modes 3 Mixotrophs 4 Protist locomotion 5 Marine algae 5 1 Diatoms 5 2 Coccolithophores 5 3 Dinoflagellates 6 Marine protozoans 6 1 Radiolarians 6 2 Foraminiferans 6 3 Amoeba 6 4 Ciliates 7 Macroscopic protists 8 Planktonic protists 8 1 Interactome 9 Protist shells 10 See also 11 References 12 Further referencesBackground edit Marine protists are a polyphyletic group of organisms playing major roles in the ecology and biogeochemistry of the oceans including performing much of Earth s photosynthesis and driving the carbon nitrogen and silicon cycles In addition marine protists occupy key positions in the tree of life including as the closest relatives of metazoans animals Unicellular eukaryotes are often lumped as protists a term that is useful despite its taxonomic irrelevance and origin as a definition by exclusion a protist being any eukaryote that s not a plant animal or fungus 12 The ocean represents the largest continuous planetary ecosystem hosting an enormous variety of organisms which include microscopic biota such as unicellular eukaryotes protists Despite their small size protists play key roles in marine biogeochemical cycles and harbour tremendous evolutionary diversity 13 14 Notwithstanding their significance for understanding the evolution of life on Earth and their role in marine food webs as well as driving biogeochemical cycles to maintain habitability little is known about their cell biology including reproduction metabolism and signaling 12 Most of the biological knowledge available is based on comparison of proteins from cultured species to homologs in genetically tractable model taxa 15 16 17 18 A main impediment to understanding the cell biology of these diverse eukaryotes is that protocols for genetic modification are available for only a small number of species 19 20 that represent neither the most ecologically relevant protists nor the breadth of eukaryotic diversity Even so in the decade to 2020 genome 15 16 17 and transcriptome sequencing initiatives 18 have resulted in nearly 120 million unigenes being identified in protists 21 which is facilitating the development of genetic tools for model species 22 nbsp Phylogenetic and symbiogenetic tree of living organisms showing a schematic view of the central position occupied by the protista protists nbsp Schematic view of the eukaryotic tree of life with effigies of main marine protist representatives 22 Trophic modes editProtists can be divided broadly into four groups depending on whether their nutrition is plant like animal like fungal like 23 or a mixture of these 24 Protists according to how they get food Type of protist Description Example Some other examples Plant like Algae see below Autotrophic protists that make their own food without needing to consume other organisms usually by photosynthesis sometimes by chemosynthesis nbsp Green algae Pyramimonas Red and brown algae diatoms coccolithophores and some dinoflagellates Plant like protists are important components of phytoplankton discussed below Animal like Protozoans see below Heterotrophic protists that get their food consuming other organisms bacteria archaea and small algae nbsp Radiolarian protist as drawn by Haeckel Foraminiferans and some marine amoebae ciliates and flagellates Fungal like Slime mouldsandslime nets Saprotrophic protists that get their food from the remains of organisms that have broken down and decayed nbsp 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 nbsp Euglena mutabilis a photosynthetic flagellate Many marine mixotrops are found among protists particularly among ciliates and dinoflagellates 5 nbsp micrograph nbsp cell schematicChoanoflagellates unicellular collared flagellate protists are thought to be the closest living relatives of the animals 25 Single celled and microscopic protists nbsp Diatoms are a major algae group generating about 20 of world oxygen production 26 nbsp Fossil diatom frustule from 32 to 40 mya nbsp Radiolarian nbsp Single celled alga Gephyrocapsa oceanica nbsp Two dinoflagellates nbsp A single celled ciliate with green zoochlorellae living inside endosymbiotically nbsp Euglenoid nbsp This ciliate is digesting cyanobacteria The cytostome or mouth is at the bottom right External videos nbsp How microscopic hunters get their lunch nbsp Euglenoids Single celled shapeshifters nbsp How do protozoans get around source source source source source source Ciliate ingesting a diatom source source source source source source Amoeba engulfing a diatom The fungus like protist saprobes are specialized to absorb nutrients from nonliving organic matter such as dead organisms or their wastes For instance many types of oomycetes grow on dead animals or algae Marine saprobic protists have the essential function of returning inorganic nutrients to the water This process allows for new algal growth which in turn generates sustenance for other organisms along the food chain Indeed without saprobe species such as protists fungi and bacteria life would cease to exist as all organic carbon became tied up in dead organisms 27 28 Mixotrophs editMixotrophic radiolarians nbsp Acantharian radiolarian hosts Phaeocystis symbionts nbsp White Phaeocystis algal foam washing up on a beach Mixotrophs have no single trophic mode 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 29 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 30 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 31 They can take advantage of different environmental conditions 32 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 33 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 34 As a result Phaeocystis is an important contributor to the marine carbon 35 and sulfur cycles 36 Phaeocystis species are endosymbionts to acantharian radiolarians 37 38 Mixotrophic plankton that combine phototrophy and heterotrophy table based on Stoecker et al 2017 39 General types Description Example Further examples Bacterioplankton Photoheterotrophic bacterioplankton nbsp Vibrio cholerae Roseobacter spp Erythrobacter spp Gammaproteobacterial clade OM60Widespread among bacteria and archaea Phytoplankton Called constitutive mixotrophs by Mitra et al 2016 40 Phytoplankton that eat photosynthetic protists with inherited plastids and the capacity to ingest prey nbsp Ochromonas species Ochromonas spp Prymnesium parvumDinoflagellate examples Fragilidium subglobosum Heterocapsa triquetra Karlodinium veneficum Neoceratium furca Prorocentrum minimum Zooplankton Called nonconstitutive mixotrophs by Mitra et al 2016 40 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 nbsp 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 nbsp Dinophysis acuminata Dinophysis spp Mesodinium rubrum 2 Protists or zooplankton with algal endosymbionts of only one algal species or very closely related algal species nbsp Noctiluca scintillans Metazooplankton with algal endosymbiontsMost 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 Mixoplankton nbsp Tintinnid ciliate Favella nbsp Euglena mutabilis a photosynthetic flagellate nbsp Zoochlorellae green living inside the ciliate Stichotricha secundaProtist locomotion editMain article Protist 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 nbsp 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 nbsp Cryptophytes All dinoflagellates and nanoflagellates choanoflagellates silicoflagellates most green algae 41 42 Other protists go through a phase as gametes when they have temporary flagellum some radiolarians foraminiferans and Apicomplexa Ciliates nbsp 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 nbsp Paramecium bursariaclick to see cilia Foraminiferans and some marine amoebae ciliates and flagellates Amoebas amoeboids nbsp Pseudopods Greek for false feet are lobe like appendages which amoebas use to anchor to a solid surface and pull themselves forward They can change their shape by extending and retracting these pseudopods 43 nbsp Amoeba Found in every major protist lineage Amoeboid cells occur among the protozoans but also in the algae and the fungi 44 45 Not motile none nbsp Diatom Coccolithophores most diatoms and non motile species of Phaeocystis 42 Among protozoans the parasitic Apicomplexa are non motile nbsp Difference of beating pattern of flagellum and cilium Flagella are used in prokaryotes archaea and bacteria as well as protists In addition both flagella and cilia are widely used in eukaryotic cells plant and animal apart from protists The regular beat patterns of eukaryotic cilia and flagella generates motion on a cellular level Examples range from the propulsion of single cells such as the swimming of spermatozoa to the transport of fluid along a stationary layer of cells such as in a respiratory tract Though eukaryotic flagella and motile cilia are ultrastructurally identical the beating pattern of the two organelles can be different In the case of flagella the motion is often planar and wave like whereas the motile cilia often perform a more complicated three dimensional motion with a power and recovery stroke Eukaryotic flagella those of animal plant and protist cells are complex cellular projections that lash back and forth Eukaryotic flagella are classed along with eukaryotic motile cilia as undulipodia 46 to emphasize their distinctive wavy appendage role in cellular function or motility Primary cilia are immotile and are not undulipodia nbsp Marine flagellates from the genera left to right Cryptaulax Abollifer Bodo Rhynchomonas Kiitoksia Allas and Metromonas 47 nbsp Cilia performs powerful forward strokes with a stiffened flagellum followed by relatively slow recovery movement with a relaxed flagellum Ciliates generally have hundreds to thousands of cilia that are densely packed together in arrays Like the flagella the cilia are powered by specialised molecular motors An efficient forward stroke is made with a stiffened flagellum followed by an inefficient backward stroke made with a relaxed flagellum During movement an individual cilium deforms as it uses the high friction power strokes and the low friction recovery strokes Since there are multiple cilia packed together on an individual organism they display collective behaviour 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 48 Flagellate ciliates and amoeba nbsp Green algal flagellate Chlamydomonas nbsp Paramecium feeding on bacteria nbsp The ciliate Oxytricha trifallax with cilia clearly visible nbsp Amoeba with ingested diatoms External videos nbsp Paramecium The White Rat of CiliatesMarine algae editAlgae 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 49 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 50 mostly multicellular and including many notable seaweeds 50 51 Brown algae form a class containing about 2 000 recognised species 52 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 53 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 nbsp Chlamydomonas globosa a unicellular green alga with two flagella just visible at bottom left nbsp Chlorella vulgaris a common green microalgae in endosymbiosis with a ciliate 54 nbsp Centric diatom nbsp Dinoflagellates 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 marine protists 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 55 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 56 Diatoms edit nbsp Diatoms come in many shapes and sizes Diatoms are photosynthetic unicellular algae populating the oceans and other waters around the globe They form a disputed phylum containing about 100 000 recognised species Diatoms generate about 20 per cent of all oxygen produced on the planet each year 26 and take in over 6 7 billion metric tons of silicon each year from the waters in which they live 57 They produce 25 45 of the total primary production of organic material in the oceans 58 59 60 owing to their prevalence in open ocean regions when total phytoplankton biomass is maximal 61 62 Diatoms are enclosed in protective silica glass shells called frustules They are classified by the shape of these glass cages in which they live and which they build as they grow Each frustule is made from two interlocking parts covered with tiny holes through which the diatom exchanges nutrients and wastes 63 Dead diatoms drift to the ocean floor where over millions of years the remains of their frustules can build up as much as half a mile deep 64 Diatoms have relatively high sinking speeds compared with other phytoplankton groups and they account for about 40 of particulate carbon exported to ocean depths 60 65 62 nbsp Diatoms are one of the most common types of phytoplankton nbsp Their protective shells frustles are made of silicon nbsp nbsp Diatom shapes nbsp nbsp Drawings by Haeckel 1904 click for details Diatoms nbsp Centric nbsp PennateDiatoms have a silica shell frustule with radial centric or bilateral pennate symmetry External videos nbsp Diatoms Tiny factories you can see from space nbsp Diatom 3D interference contrast nbsp Structure of a centric diatom frustule 66 Physically driven seasonal enrichments in surface nutrients favour diatom blooms Anthropogenic climate change will directly affect these seasonal cycles changing the timing of blooms and diminishing their biomass which will reduce primary production and CO2 uptake 67 62 Remote sensing data suggests there was a global decline of diatoms between 1998 and 2012 particularly in the North Pacific associated with shallowing of the surface mixed layer and lower nutrient concentrations 68 62 nbsp Silicified frustule of a pennate diatom with two overlapping halves nbsp Guinardia delicatula a diatom responsible for diatom blooms in the North Sea 69 nbsp There are over 100 000 species of diatoms accounting for 25 45 of the ocean s primary production nbsp Linked diatoms nbsp Pennate diatom from an Arctic meltpond infected with two chytrid like fungal pathogens Scale bar 10 µm 70 Coccolithophores edit Coccolithophores nbsp have plates called coccoliths nbsp extinct fossilCoccolithophores build calcite skeletons important to the marine carbon cycle 71 Coccolithophores are minute unicellular photosynthetic protists with two flagella for locomotion Most of them are protected by calcium carbonate shells covered with ornate circular plates or scales called coccoliths 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 72 nbsp Fossil of Coccolithus pelagicus about 10 mm across nbsp Diverse coccolithophores from the Maldives 73 nbsp The fossil coccolithophore Braarudosphaera bigelowii has an unusual shell with a regular dodecahedral structure about 10 micrometers across 74 nbsp The coccolithophore Emiliania huxleyi nbsp Algae bloom of Emiliania huxleyi off the southern coast of England nbsp Coccolithophores named after the BBC documentary seriesThe Blue Planet Dinoflagellates edit See also Mixotrophic dinoflagellate and Predatory dinoflagellate Dinoflagellate shapes nbsp Unarmored dinoflagellates Kofoid 1921 nbsp 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 75 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 41 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 76 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 Mesodinium rubrum which sequester their chloroplasts from a specific cryptophyte clade Geminigera Plagioselmis Teleaulax 39 nbsp Gyrodinium one of the few naked dinoflagellates which lack armour nbsp The dinoflagellate Protoperidinium extrudes a large feeding veil to capture prey nbsp Nassellarian radiolarians can be in symbiosis with dinoflagellates nbsp The dinoflagellate Dinophysis acuta nbsp A surf wave at night sparkles with blue light due to the presence of a bioluminescent dinoflagellate such as Lingulodinium polyedrum nbsp Suggested explanation for glowing seas 77 Dinoflagellates nbsp Armoured nbsp UnarmouredTraditionally 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 78 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 79 There is evidence from DNA analysis that dinoflagellate symbiosis with radiolarians evolved independently from other dinoflagellate symbioses such as with foraminifera 80 Some dinoflagellates are bioluminescent At night ocean water can light up internally and sparkle with blue light because of these dinoflagellates 81 82 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 83 nbsp Tripos muelleri is recognisable by its U shaped horns nbsp Oodinium a genus of parasitic dinoflagellates causes velvet disease in fish 84 nbsp Karenia brevis produces red tides highly toxic to humans 85 nbsp Red tide nbsp Noctiluca scintillans a bioluminescent dinoflagellate 86 nbsp Ornithocercus heteroporus prominent lists on displayMarine protozoans editProtozoans are protists which feed on organic matter such as other microorganisms or organic tissues and debris 87 88 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 89 90 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 edit Radiolarian shapes nbsp nbsp Drawings by Haeckel 1904 click for details Radiolarians are unicellular predatory protists encased in elaborate globular shells typically between 0 1 and 0 2 millimetres in size 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 91 nbsp Like diatoms radiolarians come in many shapes nbsp Also like diatoms radiolarian shells are usually made of silicate nbsp However acantharian radiolarians have shells made from strontium sulfate crystals nbsp Cutaway schematic diagram of a spherical radiolarian shell Turing and radiolarian morphology nbsp Shell of a spherical radiolarian nbsp Shell micrographsComputer simulations of Turing patterns on a sphereclosely replicate some radiolarian shell patterns 92 External videos nbsp Radiolarian geometry nbsp Ernst Haeckel s radiolarian engravings nbsp Cladococcus abietinus nbsp Cleveiplegma boreale Foraminiferans edit Foraminiferan shapes nbsp nbsp 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 93 They are widely researched with well established fossil records which allow scientists to infer a lot about past environments and climates 91 Foraminiferans nbsp can have more than one nucleus nbsp and defensive spinesForaminiferans are important unicellular zooplankton protists with calcium tests External videos nbsp foraminiferans nbsp Foraminiferal networks and growth nbsp section showing chambers of a spiral foram nbsp Live Ammonia tepida streaming granular ectoplasm for catching food nbsp Group of planktonic forams nbsp Fossil nummulitid forams of various sizes from the Eocene nbsp The Egyptian pyramids were constructed from limestone that contained nummulites 94 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 93 Mixotrophic foraminifers are particularly common in nutrient poor oceanic waters 95 Some forams are kleptoplastic retaining chloroplasts from ingested algae to conduct photosynthesis 96 Amoeba edit Shelled and naked amoeba nbsp Testate amoeba Cyphoderia sp nbsp Naked amoeba Chaos sp Amoeba can be shelled testate or naked nbsp Naked amoeba showing food vacuoles and ingested diatom nbsp Shell or test of a testate amoeba Arcella sp nbsp Xenogenic testate amoeba covered in diatoms from Penard s Amoeba Collection source source source source source source Amoeba engulfing a diatom External videos nbsp Amoebas nbsp Testate amoebas nbsp Feeding amoebas Ciliates edit Ciliate shapes nbsp Drawings by Haeckel 1904 click for details Marine ciliates are major grazers of the phytoplankton 97 98 Phytoplankton primary production supports higher trophic levels and fuels microbial remineralization 99 100 The dominant pelagic grazers of phytoplankton are typically associated with distinct operating modes of the food web compartments and nutrient cycling Heterotrophic protist grazers and microzooplankton dominance is usually associated with the microbial loop and regenerated production while mesozooplankton is associated with a linear food chain and export production 101 102 Grazing on particulate primary production in the global ocean surface is 10 15 for mesozooplankton and 59 75 for microzooplankton 103 104 105 106 with estimates for coastal and estuarine systems usually in the a lower range 106 98 Ciliates constitute an important component of the microzooplankton community with preference for small sized preys in contrast to mesozooplankton and many ciliate species are also grazed by mesozooplankton 107 Thus ciliates can be an important link between small cells and higher trophic levels 108 Besides their significant role in carbon transfer ciliates are also considered high quality food as a source of proteinaceous compounds with a low C N ratio in comparison to phytoplankton 109 110 98 nbsp Conjugation of two Coleps sp Two similar looking but sexually distinct partners connected at their front ends exchange genetic material via a plasma bridge Although many ciliates are heterotrophs a number of pelagic species are mixotrophic combining both phagotrophic and phototrophic nutrition Stoecker 1998 The recognition of mixotrophy in the marine plankton food web has challenged the classical understanding of pelagic food webs as autotrophy and heterotrophy are not necessarily two distinct functional compartments 111 Classical understanding of ecological interactions among plankton such as competition for nutrients indicates that nutrient uptake affinity decreases with organism size 112 favoring smaller sizes under resource limiting conditions Mixotrophy is advantageous to organisms under nutrient limited conditions allowing them to reduce direct competition by grazing on smaller prey and increase direct ingestion of nutrients 113 Modeling results suggest that mixotrophy favors larger organisms and therefore enhances trophic transfer efficiency 113 114 On top of that mixotrophy appears to be important over both space and time in marine systems 115 stressing the need for ecological field studies to further elucidate the role of mixotrophy 98 nbsp Tintinnopsis campanula nbsp Oxytricha chlorelligera nbsp Stylonychia putrina nbsp The marine ciliate Strombidium rassoulzadegani nbsp Holophyra ovum nbsp Blepharisma japonicum source source source source source source source Several taxa of ciliates interacting source source source source source source source source Blepharisma americanum swimming in a drop of pond water with other microorganisms External videos nbsp Peritrich Ciliates nbsp Conjugating protistsMacroscopic protists editMacroscopic protists see also unicellular macroalgae nbsp The single celled giant amoeba has up to 1000 nuclei and reaches lengths of 5 mm nbsp Gromia sphaerica is a large spherical testate amoeba which makes mud trails Its diameter is up to 3 8 cm 116 nbsp Spiculosiphon oceana a unicellular foraminiferan with an appearance and lifestyle that mimics a sponge grows to 5 cm long nbsp The xenophyophore another single celled foraminiferan lives in abyssal zones It has a giant shell up to 20 cm across 117 nbsp Giant kelp a brown algae is not a true plant yet it is multicellular and can grow to 50mPlanktonic protists editInteractome edit nbsp Yellow brown zooxanthellae a photosynthetic algae that lives inside hosts like radiolarians and coral nbsp Planktonic protist interactome 118 Bipartite networks providing an overview of the interactions represented by a manually curated Protist Interaction DAtabase PIDA Interaction between microbial species has played important roles in evolution and speciation 118 One of the best examples is that the origin of eukaryotes is grounded in the interaction events of endosymbiosis giving rise to mitochondria chloroplasts and other metabolic capacities in the eukaryotic cell 119 120 121 122 Microbial interactions guarantee ecosystem function having crucial roles in for instance carbon channeling in photosymbiosis control of microalgae blooms by parasites and phytoplankton associated bacteria influencing the growth and health of their host 118 Despite their importance understanding of microbial interactions in the ocean and other aquatic systems is rudimentary and the majority of them are still unknown 13 123 124 125 The earliest surveys of interactions between aquatic microbes date back to the 19th century In 1851 while on board HMS Rattlesnake in the Pacific Ocean Thomas Huxley discovered small yellow green cells inside the conspicuous planktonic radiolarians which he thought were organelles 126 Later Karl Brandt established the yellowish cells were symbiotic alga and named them zooxanthella 127 Since these early studies hundreds of others have reported microbial interactions by using classic tools mainly microscopy but this knowledge has not yet been gathered into one accessible database In recent years the high throughput sequencing HTS 128 129 130 of environmental DNA or RNA has transformed understanding of microbial diversity 131 and evolution 132 as well as generating hypotheses on microbial interactions based on correlations of estimated microbial abundances over spatiotemporal scales 133 134 135 136 118 The diagram on the right is an overview of the interactions between planktonic protists recorded in a manually curated Protist Interaction DAtabase PIDA The network is based on 2422 ecological interactions in the PIDA registered from 500 publications spanning the last 150 years The nomenclature and taxonomic order of Eukaryota is based on Adl et al 2019 137 The nomenclature and taxonomic order of Bacteria is based on Schultz et al 2017 138 118 The nodes are grouped outer circle according to eukaryotic supergroups or Incertae sedis Bacteria and Archaea All major protistan lineages were involved in interactions as hosts symbionts mutualists and commensalists parasites predators and or prey Predation was the most common interaction 39 followed by symbiosis 29 parasitism 18 and unresolved interactions 14 where it is uncertain whether the interaction is beneficial or antagonistic Nodes represent eukaryotic and prokaryotic taxa and are colored accordingly Node size indicates the number of edges links that are connected to that node Each node taxon is assigned a number which corresponds with the numbers for taxa in B C and D Edges represent interactions between two taxa and are colored according to ecological interaction type predation orange symbiosis green and parasitism purple 118 The network is undirected meaning that a node can contain both parasites symbionts prey and hosts predators To avoid cluttering of the figure Self loops which represent cases where both interacting organisms belong to the same taxon e g a dinoflagellate eating another dinoflagellate are not shown as edges links in this figure but are considered in the size of nodes The outermost circle groups taxa in the different eukaryotic supergroups or the prokaryotic domains Bacteria and Archaea Ancryomonadidae is abbreviated An Telonema is not placed into any of the supergroups but classified as Incertae sedis abbreviated I S in the figure In B B and D the following abbreviations for supergroups are used Ar Archaea Ba Bacteria Rh Rhizaria Al Alveolata St Stramenopiles Ha Haptista Cy Cryptista Ap Archaeplastida Ex Excavata Ob Obazoa Am Amoebozoa Cu CRuMS An Ancryomonadidae Is Incertae sedis 118 B Predator prey interactions in PIDA The node numbers correspond to taxa node numbers in a Abbreviations for supergroups are described above Background and nodes are colored according to functional role in the interaction Prey are colored light orange left part of figure while predators are depicted in dark orange right part of figure The size of each node represents the number of edges connected to that node 118 C Symbiont host interactions included in PIDA The node numbers correspond to node numbers in A Abbreviations for supergroups are described above Symbionts are to the left colored light green and their hosts are to the right in dark green The size of each node represents the number of edges connected to that node 118 D Parasite host interactions included in PIDA The node numbers correspond to node numbers in A Abbreviations for supergroups are described above Parasite taxa are depicted in light purple left hosts in dark purple right 118 It was found that protist predators seem to be multivorous while parasite host and symbiont host interactions appear to have moderate degrees of specialization The SAR supergroup i e Stramenopiles Alveolata and Rhizaria heavily dominated PIDA and comparisons against a global ocean molecular survey Tara expedition indicated that several SAR lineages which are abundant and diverse in the marine realm were underrepresented among the recorded interactions 118 Protist shells editMain article Protist shell See also Protists in the fossil record Many protists have protective shells or tests 139 usually made from calcium carbonate chalk or silica glass Protists are mostly single celled and microscopic Their shells are often tough mineralised forms that resist degradation and can survive the death of the protist as a microfossil Although protists are very small they are ubiquitous Their numbers are such that their shells play a huge part in the formation of ocean sediments and in the global cycling of elements and nutrients Diatom shells are called frustules and are made from silica These glass structures have accumulated for over 100 million years leaving rich deposits of nano and microstructured silicon oxide in the form of diatomaceous earth around the globe The evolutionary causes for the generation of nano and microstructured silica by photosynthetic algae are not yet clear However in 2018 it was shown that reflection of ultraviolet light by nanostructured silica protects the DNA in the algal cells and this may be an evolutionary cause for the formation of the glass cages 140 141 Coccolithophores are protected by a shell constructed from ornate circular plates or scales called coccoliths The coccoliths are made from calcium carbonate or chalk The term coccolithophore derives from the Greek for a seed carrying stone referring to their small size and the coccolith stones they carry 72 Diatoms nbsp Diatoms major components of marine plankton have glass skeletons called frustules The microscopic structures of diatoms help them manipulate light leading to hopes they could be used in new technologies for light detection computing or robotics 142 nbsp SEM images of pores in diatom frustules 140 Coccolithophores nbsp Coccolithophores are armoured with chalk plates or stones called coccoliths The images above show the size comparison between the relatively large coccolithophore Scyphosphaera apsteinii and the relatively small but ubiquitous Emiliania huxleyi 143 Benefits of having shells nbsp Benefits in coccolithophore calcification 144 see text below Costs of having shells nbsp Energetic costs in coccolithophore calcification 144 There are benefits for protists that carry protective shells The diagram on the left above shows some benefits coccolithophore get from carrying coccoliths In the diagram A represents accelerated photosynthesis including carbon concentrating mechanisms CCM and enhanced light uptake via scattering of scarce photons for deep dwelling species B represents protection from photodamage including sunshade protection from ultraviolet light UV and photosynthetic active radiation PAR and energy dissipation under high light conditions C represents armour protection includes protection against viral bacterial infections and grazing by selective and nonselective grazers 144 There are also costs for protists that carry protective shells The diagram on the right above shows some of the energetic costs coccolithophore incur from carrying coccoliths In the diagram the energetic costs are reported in percentage of total photosynthetic budget A represents transport processes include the transport into the cell from the surrounding seawater of primary calcification substrates Ca2 and HCO3 black arrows and the removal of the end product H from the cell gray arrow The transport of Ca2 through the cytoplasm to the coccolith vesicle CV is the dominant cost associated with calcification B represents metabolic processes include the synthesis of coccolith associated polysaccharides CAPs gray rectangles by the Golgi complex white rectangles that regulate the nucleation and geometry of CaCO3 crystals The completed coccolith gray plate is a complex structure of intricately arranged CAPs and CaCO3 crystals C Mechanical and structural processes account for the secretion of the completed coccoliths that are transported from their original position adjacent to the nucleus to the cell periphery where they are transferred to the surface of the cell 144 See also editCavalier Smith s system of classificationReferences edit Cavalier Smith T December 1993 Kingdom protozoa and its 18 phyla Microbiological Reviews 57 4 953 94 doi 10 1128 mmbr 57 4 953 994 1993 PMC 372943 PMID 8302218 Corliss JO 1992 Should there be a separate code of nomenclature for the protists BioSystems 28 1 3 1 14 doi 10 1016 0303 2647 92 90003 H PMID 1292654 Slapeta J Moreira D Lopez Garcia P 2005 The extent of protist diversity insights from molecular ecology of freshwater eukaryotes Proceedings of the Royal Society B Biological Sciences 272 1576 2073 81 doi 10 1098 rspb 2005 3195 PMC 1559898 PMID 16191619 Moreira D Lopez Garcia P 2002 The molecular ecology of microbial eukaryotes unveils a hidden world PDF Trends in Microbiology 10 1 31 8 doi 10 1016 S0966 842X 01 02257 0 PMID 11755083 a b c Leles S G Mitra A Flynn K J Stoecker D K Hansen P J Calbet A McManus G B Sanders R W Caron D A Not F Hallegraeff G M 2017 Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance Proceedings of the Royal Society B Biological Sciences 284 1860 20170664 doi 10 1098 rspb 2017 0664 PMC 5563798 PMID 28768886 Characteristics of Protists In Rye Connie Avissar Yael Choi Jung Ho DeSaix Jean Jurukovski Vladimir Wise Robert R 2013 Biology Houston Texas ISBN 978 1 938168 09 3 OCLC 896421272 a href Template Cite book html title Template Cite book cite book a CS1 maint location missing publisher link nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Devreotes P 1989 Dictyostelium discoideum a model system for cell cell interactions in development Science 245 4922 1054 8 Bibcode 1989Sci 245 1054D doi 10 1126 science 2672337 PMID 2672337 Neil A C Reece J B Simon E J 2004 Essential biology with physiology Pearson Benjamin Cummings Page 291 ISBN 9780805375039 O Malley MA Simpson AG Roger AJ 2012 The other eukaryotes in light of evolutionary protistology Biology amp Philosophy 28 2 299 330 doi 10 1007 s10539 012 9354 y S2CID 85406712 Adl SM Simpson AG Farmer MA Andersen RA Anderson OR Barta JR Bowser SS Brugerolle G Fensome RA Fredericq S James TY Karpov S Kugrens P Krug J Lane CE Lewis LA Lodge J Lynn DH Mann DG McCourt RM Mendoza L Moestrup O Mozley Standridge SE Nerad TA Shearer CA Smirnov AV Spiegel FW Taylor MF 2005 The new higher level classification of eukaryotes with emphasis on the taxonomy of protists The Journal of Eukaryotic Microbiology 52 5 399 451 doi 10 1111 j 1550 7408 2005 00053 x PMID 16248873 S2CID 8060916 Margulis L Chapman MJ 19 March 2009 Kingdoms and Domains An Illustrated Guide to the Phyla of Life on Earth Academic Press ISBN 9780080920146 a b Collier Jackie L Rest Joshua S 2019 Swimming gliding and rolling toward the mainstream Cell biology of marine protists Molecular Biology of the Cell 30 11 1245 1248 doi 10 1091 mbc E18 11 0724 PMC 6724603 PMID 31084566 a b Worden A Z Follows M J Giovannoni S J Wilken S Zimmerman A E Keeling P J 2015 Rethinking the marine carbon cycle Factoring in the multifarious lifestyles of microbes Science 347 6223 doi 10 1126 science 1257594 PMID 25678667 S2CID 206560125 De Vargas C et al 2015 Eukaryotic plankton diversity in the sunlit ocean Science 348 6237 doi 10 1126 science 1261605 hdl 10261 117736 PMID 25999516 S2CID 12853481 a b Curtis Bruce A et al 2012 Algal genomes reveal evolutionary mosaicism and the fate of nucleomorphs Nature 492 7427 59 65 Bibcode 2012Natur 492 59C doi 10 1038 nature11681 PMID 23201678 S2CID 4380094 a b Armbrust E V et al 2004 The Genome of the Diatom Thalassiosira Pseudonana Ecology Evolution and Metabolism Science 306 5693 79 86 Bibcode 2004Sci 306 79A doi 10 1126 science 1101156 PMID 15459382 S2CID 8593895 a b Read Betsy A et al 2013 Pan genome of the phytoplankton Emiliania underpins its global distribution Nature 499 7457 209 213 Bibcode 2013Natur 499 209 doi 10 1038 nature12221 hdl 1854 LU 4120924 PMID 23760476 S2CID 4428297 a b Keeling Patrick J et al 2014 The Marine Microbial Eukaryote Transcriptome Sequencing Project MMETSP Illuminating the Functional Diversity of Eukaryotic Life in the Oceans through Transcriptome Sequencing PLOS Biology 12 6 e1001889 doi 10 1371 journal pbio 1001889 PMC 4068987 PMID 24959919 Nymark Marianne Sharma Amit Kumar Sparstad Torfinn Bones Atle M Winge Per 2016 A CRISPR Cas9 system adapted for gene editing in marine algae Scientific Reports 6 24951 Bibcode 2016NatSR 624951N doi 10 1038 srep24951 PMC 4842962 PMID 27108533 Hopes Amanda Nekrasov Vladimir Kamoun Sophien Mock Thomas 2016 Editing of the urease gene by CRISPR Cas in the diatom Thalassiosira pseudonana Plant Methods 12 49 doi 10 1186 s13007 016 0148 0 PMC 5121945 PMID 27904648 Carradec Quentin et al 2018 A global ocean atlas of eukaryotic genes Nature Communications 9 1 373 Bibcode 2018NatCo 9 373C doi 10 1038 s41467 017 02342 1 PMC 5785536 PMID 29371626 a b Faktorova Drahomira et al 2020 Genetic tool development in marine protists Emerging model organisms for experimental cell biology Nature Methods 17 5 481 494 doi 10 1038 s41592 020 0796 x PMC 7200600 PMID 32251396 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Whittaker R H Margulis L 1978 Protist classification and the kingdoms of organisms Biosystems 10 1 2 3 18 doi 10 1016 0303 2647 78 90023 0 PMID 418827 Faure E Not F Benoiston AS Labadie K Bittner L Ayata SD 2019 Mixotrophic protists display contrasted biogeographies in the global ocean ISME Journal 13 4 1072 1083 doi 10 1038 s41396 018 0340 5 PMC 6461780 PMID 30643201 Budd Graham E Jensen Soren 2017 The origin of the animals and a Savannah hypothesis for early bilaterian evolution Biological Reviews 92 1 446 473 doi 10 1111 brv 12239 PMID 26588818 a b The Air You re Breathing A Diatom Made That Clark M A Douglas M and Choi J 2018 Biology 2e 23 4 Ecology of Protists OpenStax Houston Texas nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Vallet Marine Baumeister Tim U H Kaftan Filip Grabe Veit Buaya Anthony Thines Marco Svatos Ales Pohnert Georg 2019 The oomycete Lagenisma coscinodisci hijacks host alkaloid synthesis during infection of a marine diatom Nature Communications 10 1 4938 Bibcode 2019NatCo 10 4938V doi 10 1038 s41467 019 12908 w PMC 6821873 PMID 31666506 Beware the mixotrophs they can destroy entire ecosystems in a matter of hours Microscopic body snatchers infest our oceans Phys org Eiler A December 2006 Evidence for the Ubiquity of Mixotrophic Bacteria in the Upper Ocean Implications and Consequences Appl Environ Microbiol 72 12 7431 7 Bibcode 2006ApEnM 72 7431E doi 10 1128 AEM 01559 06 PMC 1694265 PMID 17028233 Katechakis A Stibor H July 2006 The mixotroph Ochromonas tuberculata may invade and suppress specialist phago and phototroph plankton communities depending on nutrient conditions Oecologia 148 4 692 701 Bibcode 2006Oecol 148 692K doi 10 1007 s00442 006 0413 4 PMID 16568278 S2CID 22837754 Schoemann Veronique Becquevort Sylvie Stefels Jacqueline Rousseau Veronique Lancelot Christiane 1 January 2005 Phaeocystis blooms in the global ocean and their controlling mechanisms a review Journal of Sea Research Iron Resources and Oceanic Nutrients Advancement of Global Environmental Simulations 53 1 2 43 66 Bibcode 2005JSR 53 43S CiteSeerX 10 1 1 319 9563 doi 10 1016 j seares 2004 01 008 Welcome to the Phaeocystis antarctica genome sequencing project homepage DiTullio G R Grebmeier J M Arrigo K R Lizotte M P Robinson D H Leventer A Barry J P VanWoert M L Dunbar R B 2000 Rapid and early export of Phaeocystis antarctica blooms in the Ross Sea Antarctica Nature 404 6778 595 598 Bibcode 2000Natur 404 595D doi 10 1038 35007061 PMID 10766240 S2CID 4409009 J Stefels L Dijkhuizen WWC Gieskes 20 July 1995 DMSP lyase activity in a spring phytoplankton bloom off the Dutch coast related to Phaeocystis sp abundance PDF Marine Ecology Progress Series 123 235 243 Bibcode 1995MEPS 123 235S doi 10 3354 meps123235 Decelle Johan Simo Rafel Gali Marti Vargas Colomban de Colin Sebastien Desdevises Yves Bittner Lucie Probert Ian Not Fabrice 30 October 2012 An original mode of symbiosis in open ocean plankton Proceedings of the National Academy of Sciences 109 44 18000 18005 Bibcode 2012PNAS 10918000D doi 10 1073 pnas 1212303109 ISSN 0027 8424 PMC 3497740 PMID 23071304 Mars Brisbin Margaret Grossmann Mary M Mesrop Lisa Y Mitarai Satoshi 2018 Intra host Symbiont Diversity and Extended Symbiont Maintenance in Photosymbiotic Acantharea Clade F Frontiers in Microbiology 9 1998 doi 10 3389 fmicb 2018 01998 ISSN 1664 302X PMC 6120437 PMID 30210473 a b Stoecker D K Hansen P J Caron D A Mitra A 2017 Mixotrophy in the marine plankton Annual Review of Marine Science 9 311 335 Bibcode 2017ARMS 9 311S doi 10 1146 annurev marine 010816 060617 PMID 27483121 a b Mitra A Flynn KJ Tillmann U Raven J Caron D et al 2016 Defining planktonic protist functional groups on mechanisms for energy and nutrient acquisition incorporation of diverse mixotrophic strategies Protist 167 2 106 20 doi 10 1016 j protis 2016 01 003 hdl 10261 131722 PMID 26927496 a b Dawson Scott C Paredez Alexander R 2013 Alternative cytoskeletal landscapes cytoskeletal novelty and evolution in basal excavate protists Current Opinion in Cell Biology 25 1 134 141 doi 10 1016 j ceb 2012 11 005 PMC 4927265 PMID 23312067 a b Atkinson A Polimene L Fileman E S Widdicombe C E McEvoy A J Smyth T J Djeghri N Sailley S F Cornwell L E 2018 Comment What drives plankton seasonality in a stratifying shelf sea Some competing and complementary theories PDF Limnology and Oceanography 63 6 2877 2884 Bibcode 2018LimOc 63 2877A doi 10 1002 lno 11036 S2CID 91380765 Singleton Paul 2006 Dictionary of Microbiology and Molecular Biology 3rd Edition revised Chichester UK John Wiley amp Sons pp 32 ISBN 978 0 470 03545 0 David J Patterson Amoebae Protists Which Move and Feed Using Pseudopodia Tree of Life web project The Amoebae The University of Edinburgh Archived from the original on 10 June 2009 A Dictionary of Biology 2004 accessed 2011 01 01 Patterson David J 2000 Flagellates Heterotrophic Protists With Flagella Tree of Life Lauga Eric Thomas R Powers 25 August 2009 The hydrodynamics of swimming microorganisms Reports on Progress in Physics 72 9 096601 arXiv 0812 2887 Bibcode 2009RPPh 72i6601L doi 10 1088 0034 4885 72 9 096601 S2CID 3932471 Guiry MD October 2012 How many species of algae are there Journal of Phycology 48 5 1057 63 doi 10 1111 j 1529 8817 2012 01222 x PMID 27011267 S2CID 30911529 a b Guiry M D Guiry G M 2016 Algaebase www algaebase org Retrieved 20 November 2016 D Thomas 2002 Seaweeds Life Series Natural History Museum London ISBN 978 0 565 09175 0 Hoek Christiaan den Hoeck Hoeck Van Mann David Jahns H M 1995 Algae an introduction to phycology Cambridge University Press p 166 ISBN 9780521316873 OCLC 443576944 Starckx Senne 31 October 2012 A place in the sun Algae is the crop of the future according to researchers in Geel Archived 7 November 2017 at the Wayback Machine Flanders Today Retrieved 8 December 2012 Duval B Margulis L 1995 The microbial community of Ophrydium versatile colonies endosymbionts residents and tenants Symbiosis 18 181 210 PMID 11539474 Mandoli DF 1998 Elaboration of Body Plan and Phase Change during Development of Acetabularia How Is the Complex Architecture of a Giant Unicell Built Annual Review of Plant Physiology and Plant Molecular Biology 49 173 198 doi 10 1146 annurev arplant 49 1 173 PMID 15012232 S2CID 6241264 Pierre Madl Maricela Yip 2004 Literature Review of Caulerpa taxifolia BUFUS Info 19 31 Archived from the original on 8 October 2022 Retrieved 12 May 2020 Treguer P Nelson D M Van Bennekom A J Demaster D J Leynaert A Queguiner B 1995 The Silica Balance in the World Ocean A Reestimate Science 268 5209 375 9 Bibcode 1995Sci 268 375T doi 10 1126 science 268 5209 375 PMID 17746543 S2CID 5672525 Nelson David M Treguer Paul Brzezinski Mark A Leynaert Aude Queguiner Bernard 1995 Production and dissolution of biogenic silica in the ocean Revised global estimates comparison with regional data and relationship to biogenic sedimentation Global Biogeochemical Cycles 9 3 359 372 Bibcode 1995GBioC 9 359N doi 10 1029 95GB01070 Malviya Shruti Scalco Eleonora Audic Stephane Vincent Flora Veluchamy Alaguraj Poulain Julie Wincker Patrick Iudicone Daniele De Vargas Colomban Bittner Lucie Zingone Adriana Bowler Chris 2016 Insights into global diatom distribution and diversity in the world s ocean Proceedings of the National Academy of Sciences 113 11 E1516 E1525 Bibcode 2016PNAS 113E1516M doi 10 1073 pnas 1509523113 PMC 4801293 PMID 26929361 S2CID 22035749 a b Treguer Paul Bowler Chris Moriceau Brivaela Dutkiewicz Stephanie Gehlen Marion Aumont Olivier Bittner Lucie Dugdale Richard Finkel Zoe Iudicone Daniele Jahn Oliver Guidi Lionel Lasbleiz Marine Leblanc Karine Levy Marina Pondaven Philippe 2018 Influence of diatom diversity on the ocean biological carbon pump PDF Nature Geoscience 11 1 27 37 Bibcode 2018NatGe 11 27T doi 10 1038 s41561 017 0028 x S2CID 134885922 Mahadevan Amala d Asaro Eric Lee Craig Perry Mary Jane 2012 Eddy Driven Stratification Initiates North Atlantic Spring Phytoplankton Blooms Science 337 6090 54 58 Bibcode 2012Sci 337 54M doi 10 1126 science 1218740 PMID 22767922 S2CID 42312402 a b c d Cavicchioli Ricardo Ripple William J Timmis Kenneth N Azam Farooq Bakken Lars R Baylis Matthew Behrenfeld Michael J Boetius Antje Boyd Philip W Classen Aimee T Crowther Thomas W Danovaro Roberto Foreman Christine M Huisman Jef Hutchins David A Jansson Janet K Karl David M Koskella Britt Mark Welch David B Martiny Jennifer B H Moran Mary Ann Orphan Victoria J Reay David S Remais Justin V Rich Virginia I Singh Brajesh K Stein Lisa Y Stewart Frank J Sullivan Matthew B et al 2019 Scientists warning to humanity Microorganisms and climate change Nature Reviews Microbiology 17 9 569 586 doi 10 1038 s41579 019 0222 5 PMC 7136171 PMID 31213707 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Wassilieff Maggy 2006 Plankton Plant plankton Te Ara the Encyclopedia of New Zealand Accessed 2 November 2019 King s College London Lake Megachad www kcl ac uk Retrieved 5 May 2018 Boyd Philip W Claustre Herve Levy Marina Siegel David A Weber Thomas 2019 Multi faceted particle pumps drive carbon sequestration in the ocean PDF Nature 568 7752 327 335 Bibcode 2019Natur 568 327B doi 10 1038 s41586 019 1098 2 PMID 30996317 S2CID 119513489 Zhang D Wang Y Cai J Pan J Jiang X Jiang Y 2012 Bio manufacturing technology based on diatom micro and nanostructure Chinese Science Bulletin 57 30 3836 3849 Bibcode 2012ChSBu 57 3836Z doi 10 1007 s11434 012 5410 x Behrenfeld Michael J Doney Scott C Lima Ivan Boss Emmanuel S Siegel David A 2013 Annual cycles of ecological disturbance and recovery underlying the subarctic Atlantic spring plankton bloom Global Biogeochemical Cycles 27 2 526 540 Bibcode 2013GBioC 27 526B doi 10 1002 gbc 20050 hdl 1912 6250 Rousseaux Cecile S Gregg Watson W 2015 Recent decadal trends in global phytoplankton composition Global Biogeochemical Cycles 29 10 1674 1688 Bibcode 2015GBioC 29 1674R doi 10 1002 2015GB005139 Arsenieff L Simon N Rigaut Jalabert F Le Gall F Chaffron S Corre E Com E Bigeard E Baudoux A C 2018 First Viruses Infecting the Marine Diatom Guinardia delicatula Frontiers in Microbiology 9 3235 doi 10 3389 fmicb 2018 03235 PMC 6334475 PMID 30687251 Kilias Estelle S Junges Leandro Supraha Luka Leonard Guy Metfies Katja Richards Thomas A 2020 Chytrid fungi distribution and co occurrence with diatoms correlate with sea ice melt in the Arctic Ocean Communications Biology 3 1 183 doi 10 1038 s42003 020 0891 7 PMC 7174370 PMID 32317738 S2CID 216033140 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Rost B and Riebesell U 2004 Coccolithophores and the biological pump responses to environmental changes In Coccolithophores From Molecular Processes to Global Impact pages 99 125 Springer ISBN 9783662062784 a b Wassilieff Maggy 2006 A coccolithophore Te Ara the Encyclopedia of New Zealand Accessed 2 November 2019 Young J R Pratiwi S Su X 2017 Data report Surface seawater plankton sampling for coccolithophores undertaken during IODP Expedition 359 Proceedings of the International Ocean Discovery Program doi 10 14379 iodp proc 359 111 2017 a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Hagino K Onuma R Kawachi M and Horiguchi T 2013 Discovery of an endosymbiotic nitrogen fixing cyanobacterium UCYN A in Braarudosphaera bigelowii Prymnesiophyceae PLoS One 8 12 e81749 doi 10 1371 journal pone 0081749 Gomez F 2012 A checklist and classification of living dinoflagellates Dinoflagellata Alveolata CICIMAR Oceanides 27 1 65 140 doi 10 37543 oceanides v27i1 111 Stoecker DK 1999 Mixotrophy among Dinoflagellates The Journal of Eukaryotic Microbiology 46 4 397 401 doi 10 1111 j 1550 7408 1999 tb04619 x S2CID 83885629 Suggested Explanation for Glowing Seas Including Currently Glowing California Seas National Science Foundation 18 October 2011 Boltovskoy Demetrio Anderson O Roger Correa Nancy M 2017 Radiolaria and Phaeodaria Handbook of the Protists Springer Cham pp 731 763 doi 10 1007 978 3 319 28149 0 19 ISBN 9783319281476 Anderson O R 1983 Radiolaria Springer Science amp Business Media Gast R J Caron D A 1 November 1996 Molecular phylogeny of symbiotic dinoflagellates from planktonic foraminifera and radiolaria Molecular Biology and Evolution 13 9 1192 1197 doi 10 1093 oxfordjournals molbev a025684 ISSN 0737 4038 PMID 8896371 Castro P Huber ME 2010 Marine Biology 8th ed McGraw Hill pp 95 ISBN 978 0071113021 Hastings JW 1996 Chemistries and colors of bioluminescent reactions a review Gene 173 1 Spec No 5 11 doi 10 1016 0378 1119 95 00676 1 PMID 8707056 Haddock SH Moline MA Case JF 2009 Bioluminescence in the sea Annual Review of Marine Science 2 443 93 Bibcode 2010ARMS 2 443H doi 10 1146 annurev marine 120308 081028 PMID 21141672 S2CID 3872860 Protozoa Infecting Gills and Skin The Merck Veterinary Manual Archived from the original on 3 March 2016 Retrieved 4 November 2019 Brand Larry E Campbell Lisa Bresnan Eileen 2012 Karenia The biology and ecology of a toxic genus Harmful Algae 14 156 178 doi 10 1016 j hal 2011 10 020 PMC 9891709 PMID 36733478 Buskey E J 1995 Growth and bioluminescence of Noctiluca scintillans on varying algal diets Journal of Plankton Research 17 1 29 40 doi 10 1093 plankt 17 1 29 Panno Joseph 14 May 2014 The Cell Evolution of the First Organism Infobase Publishing ISBN 9780816067367 Bertrand Jean Claude Caumette Pierre Lebaron Philippe Matheron Robert Normand Philippe Sime Ngando Telesphore 26 January 2015 Environmental Microbiology Fundamentals and Applications Microbial Ecology Springer ISBN 9789401791182 Madigan Michael T 2012 Brock Biology of Microorganisms Benjamin Cummings ISBN 9780321649638 Yaeger Robert G 1996 Protozoa Structure Classification Growth and Development NCBI ISBN 9780963117212 PMID 21413323 Retrieved 23 March 2018 a b Wassilieff Maggy 2006 Plankton Animal plankton Te Ara the Encyclopedia of New Zealand Accessed 2 November 2019 Varea C Aragon J L Barrio R A 1999 Turing patterns on a sphere Physical Review E 60 4 4588 92 Bibcode 1999PhRvE 60 4588V doi 10 1103 PhysRevE 60 4588 PMID 11970318 a b Hemleben C Anderson O R Spindler M 1989 Modern Planktonic Foraminifera Springer Verlag ISBN 978 3 540 96815 3 Foraminifera History of Study University College London Retrieved 18 November 2019 Advances in Microbial Ecology Volum 11 Bernhard J M Bowser S M 1999 Benthic Foraminifera of dysoxic sediments chloroplast sequestration and functional morphology Earth Science Reviews 46 1 149 165 Bibcode 1999ESRv 46 149B doi 10 1016 S0012 8252 99 00017 3 Calbet Albert Landry Michael R 2004 Phytoplankton growth microzooplankton grazing and carbon cycling in marine systems Limnology and Oceanography 49 1 51 57 Bibcode 2004LimOc 49 51C doi 10 4319 lo 2004 49 1 0051 hdl 10261 134985 S2CID 22995996 a b c d Haraguchi Lumi Jakobsen Hans H Lundholm Nina Carstensen Jacob 2018 Phytoplankton Community Dynamic A Driver for Ciliate Trophic Strategies Frontiers in Marine Science 5 doi 10 3389 fmars 2018 00272 S2CID 51925344 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Azam F Fenchel T Field JG Gray JS Meyer Reil LA Thingstad F 1983 The Ecological Role of Water Column Microbes in the Sea Marine Ecology Progress Series 10 257 263 Bibcode 1983MEPS 10 257A doi 10 3354 meps010257 Sherr Evelyn Sherr Barry 1988 Role of microbes in pelagic food webs A revised concept Limnology and Oceanography 33 5 1225 1227 Bibcode 1988LimOc 33 1225S doi 10 4319 lo 1988 33 5 1225 Fenchel T 1988 Marine Plankton Food Chains Annual Review of Ecology and Systematics 19 19 38 doi 10 1146 annurev es 19 110188 000315 Buitenhuis Erik Le Quere Corinne Aumont Olivier Beaugrand Gregory Bunker Adrian Hirst Andrew Ikeda Tsutomu O Brien Todd Piontkovski Sergey Straile Dietmar 2006 Biogeochemical fluxes through mesozooplankton Global Biogeochemical Cycles 20 2 n a Bibcode 2006GBioC 20 2003B doi 10 1029 2005GB002511 hdl 2115 13694 Behrenfeld Michael J Falkowski Paul G 1997 Photosynthetic rates derived from satellite based chlorophyll concentration Limnology and Oceanography 42 1 1 20 Bibcode 1997LimOc 42 1B doi 10 4319 lo 1997 42 1 0001 S2CID 15857675 Calbet Albert 2001 Mesozooplankton grazing effect on primary production A global comparative analysis in marine ecosystems Limnology and Oceanography 46 7 1824 1830 Bibcode 2001LimOc 46 1824C doi 10 4319 lo 2001 46 7 1824 hdl 10261 49263 S2CID 85461746 Landry Michael R Calbet Albert 2004 Microzooplankton production in the oceans ICES Journal of Marine Science 61 4 501 507 doi 10 1016 j icesjms 2004 03 011 a b Buitenhuis Erik T Rivkin Richard B Sailley Sevrine Le Quere Corinne 2010 Biogeochemical fluxes through microzooplankton Global Biogeochemical Cycles 24 4 n a Bibcode 2010GBioC 24 4015B doi 10 1029 2009GB003601 S2CID 131413083 Hansen Per Juel Bjornsen Peter Koefoed Hansen Benni Winding 2000 Zooplankton grazing and growth Scaling within the 2 2 000 µm body size range Limnology and Oceanography 45 8 1891 Bibcode 2000LimOc 45 1891H doi 10 4319 lo 2000 45 8 1891 Nielsen Torkel Gissel Kicrboe Thomas 1994 Regulation of zooplankton biomass and production in a temperate coastal ecosystem 2 Ciliates Limnology and Oceanography 39 3 508 519 Bibcode 1994LimOc 39 508N doi 10 4319 lo 1994 39 3 0508 Stoecker Diane K Capuzzo Judith Mcdowell 1990 Predation on Protozoa its importance to zooplankton Journal of Plankton Research 12 5 891 908 doi 10 1093 plankt 12 5 891 Gifford Dian J 1991 The Protozoan Metazoan Trophic Link in Pelagic Ecosystems The Journal of Protozoology 38 81 86 doi 10 1111 j 1550 7408 1991 tb04806 x Flynn Kevin J Stoecker Diane K Mitra Aditee Raven John A Glibert Patricia M Hansen Per Juel Graneli Edna Burkholder Joann M 2013 Misuse of the phytoplankton zooplankton dichotomy The need to assign organisms as mixotrophs within plankton functional types Journal of Plankton Research 35 3 11 doi 10 1093 plankt fbs062 Edwards Kyle F Thomas Mridul K Klausmeier Christopher A Litchman Elena 2012 Allometric scaling and taxonomic variation in nutrient utilization traits and maximum growth rate of phytoplankton Limnology and Oceanography 57 2 554 566 Bibcode 2012LimOc 57 554E doi 10 4319 lo 2012 57 2 0554 S2CID 13376583 a b Mitra A Flynn K J Burkholder J M Berge T Calbet A Raven J A Graneli E Glibert P M Hansen P J Stoecker D K Thingstad F Tillmann U Vage S Wilken S Zubkov M V 2014 The role of mixotrophic protists in the biological carbon pump Biogeosciences 11 4 995 1005 Bibcode 2014BGeo 11 995M doi 10 5194 bg 11 995 2014 hdl 10261 93693 Ward Ben A Follows Michael J 2016 Marine mixotrophy increases trophic transfer efficiency mean organism size and vertical carbon flux Proceedings of the National Academy of Sciences 113 11 2958 2963 Bibcode 2016PNAS 113 2958W doi 10 1073 pnas 1517118113 PMC 4801304 PMID 26831076 Leles S G Mitra A Flynn K J Stoecker D K Hansen P J Calbet A McManus G B Sanders R W Caron D A Not F Hallegraeff G M Pitta P Raven J A Johnson M D Glibert P M Vage S 2017 Oceanic protists with different forms of acquired phototrophy display contrasting biogeographies and abundance Proceedings of the Royal Society B Biological Sciences 284 1860 doi 10 1098 rspb 2017 0664 PMC 5563798 PMID 28768886 Matz Mikhail V Tamara M Frank N Justin Marshall Edith A Widder Sonke Johnsen 9 December 2008 Giant Deep Sea Protist Produces Bilaterian like Traces PDF Current Biology 18 23 Elsevier Ltd 1849 1854 doi 10 1016 j cub 2008 10 028 PMID 19026540 S2CID 8819675 Gooday A J Aranda da Silva A Pawlowski J 1 December 2011 Xenophyophores Rhizaria Foraminifera from the Nazare Canyon Portuguese margin NE Atlantic Deep Sea Research Part II Topical Studies in Oceanography The Geology Geochemistry and Biology of Submarine Canyons West of Portugal 58 23 24 2401 2419 Bibcode 2011DSRII 58 2401G doi 10 1016 j dsr2 2011 04 005 a b c d e f g h i j k Bjorbaekmo Marit F Markussen Evenstad Andreas Rosaeg Line Lieblein Krabberod Anders K Logares Ramiro 2020 The planktonic protist interactome where do we stand after a century of research The ISME Journal 14 2 544 559 doi 10 1038 s41396 019 0542 5 PMC 6976576 PMID 31685936 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License Margulis Lynn Fester Rene 1991 Symbiosis as a Source of Evolutionary Innovation Speciation and Morphogenesis MIT Press ISBN 9780262132695 Lopez Garcia Purificacion Eme Laura Moreira David 2017 Symbiosis in eukaryotic evolution Journal of Theoretical Biology 434 20 33 Bibcode 2017JThBi 434 20L doi 10 1016 j jtbi 2017 02 031 PMC 5638015 PMID 28254477 Archibald John M 2015 Endosymbiosis and Eukaryotic Cell Evolution Current Biology 25 19 R911 R921 doi 10 1016 j cub 2015 07 055 PMID 26439354 S2CID 16089231 Cavalier Smith Thomas 2013 Symbiogenesis Mechanisms Evolutionary Consequences and Systematic Implications Annual Review of Ecology Evolution and Systematics 44 145 172 doi 10 1146 annurev ecolsys 110411 160320 Mahe Frederic De Vargas Colomban Bass David Czech Lucas Stamatakis Alexandros Lara Enrique Singer David Mayor Jordan Bunge John Sernaker Sarah Siemensmeyer Tobias Trautmann Isabelle Romac Sarah Berney Cedric Kozlov Alexey Mitchell Edward A D Seppey Christophe V W Egge Elianne Lentendu Guillaume Wirth Rainer Trueba Gabriel Dunthorn Micah 2017 Parasites dominate hyperdiverse soil protist communities in Neotropical rainforests Nature Ecology amp Evolution 1 4 91 doi 10 1038 s41559 017 0091 PMID 28812652 S2CID 2631960 Biard Tristan Stemmann Lars Picheral Marc Mayot Nicolas Vandromme Pieter Hauss Helena Gorsky Gabriel Guidi Lionel Kiko Rainer Not Fabrice 2016 In situ imaging reveals the biomass of giant protists in the global ocean PDF Nature 532 7600 504 507 Bibcode 2016Natur 532 504B doi 10 1038 nature17652 PMID 27096373 S2CID 205248710 Finlay B J Esteban G F 1998 Freshwater protozoa Biodiversity and ecological function Biodiversity and Conservation 7 9 1163 1186 doi 10 1023 A 1008879616066 S2CID 10702795 Huxley Thomas H 1851 XXXIV Zoological notes and observations made on board H M S Rattlesnake Annals and Magazine of Natural History 8 48 433 442 doi 10 1080 03745486109495002 Brandt K 1881 Uber das Zusammenleben von Thieren und Algen Verh Physiol Ges 1 524 527 Logares Ramiro Haverkamp Thomas H A Kumar Surendra Lanzen Anders Nederbragt Alexander J Quince Christopher Kauserud Havard 2012 Environmental microbiology through the lens of high throughput DNA sequencing Synopsis of current platforms and bioinformatics approaches Journal of Microbiological Methods 91 1 106 113 doi 10 1016 j mimet 2012 07 017 PMID 22849829 Sogin M L Morrison H G Huber J A Welch D M Huse S M Neal P R Arrieta J M Herndl G J 2006 Microbial diversity in the deep sea and the underexplored rare biosphere Proceedings of the National Academy of Sciences 103 32 12115 12120 Bibcode 2006PNAS 10312115S doi 10 1073 pnas 0605127103 PMC 1524930 PMID 16880384 Goodwin Sara McPherson John D McCombie W Richard 2016 Coming of age Ten years of next generation sequencing technologies Nature Reviews Genetics 17 6 333 351 doi 10 1038 nrg 2016 49 PMC 10373632 PMID 27184599 S2CID 8295541 Pedros Alio C Acinas SG Logares R Massana R Marine microbial diversity as seen by high throughput sequencing In Gasol Josep M Kirchman David L 27 March 2018 Microbial Ecology of the Oceans John Wiley amp Sons ISBN 9781119107187 pp 47 87 Spang Anja Saw Jimmy H Jorgensen Steffen L Zaremba Niedzwiedzka Katarzyna Martijn Joran Lind Anders E Van Eijk Roel Schleper Christa Guy Lionel Ettema Thijs J G 2015 Complex archaea that bridge the gap between prokaryotes and eukaryotes Nature 521 7551 173 179 Bibcode 2015Natur 521 173S doi 10 1038 nature14447 PMC 4444528 PMID 25945739 Faust Karoline Lahti Leo Gonze Didier De Vos Willem M Raes Jeroen 2015 Metagenomics meets time series analysis Unraveling microbial community dynamics Current Opinion in Microbiology 25 56 66 doi 10 1016 j mib 2015 04 004 PMID 26005845 Faust Karoline Raes Jeroen 2012 Microbial interactions From networks to models Nature Reviews Microbiology 10 8 538 550 doi 10 1038 nrmicro2832 PMID 22796884 S2CID 22872711 Lima Mendez G et al 2015 Determinants of community structure in the global plankton interactome Science 348 6237 doi 10 1126 science 1262073 hdl 10261 117702 PMID 25999517 S2CID 10326640 Layeghifard Mehdi Hwang David M Guttman David S 2017 Disentangling Interactions in the Microbiome A Network Perspective Trends in Microbiology 25 3 217 228 doi 10 1016 j tim 2016 11 008 PMC 7172547 PMID 27916383 Adl Sina M et al 2019 Revisions to the classification nomenclature and diversity of eukaryotes Journal of Eukaryotic Microbiology 66 1 4 119 doi 10 1111 jeu 12691 PMC 6492006 PMID 30257078 Schulz Frederik Eloe Fadrosh Emiley A Bowers Robert M Jarett Jessica Nielsen Torben Ivanova Natalia N Kyrpides Nikos C Woyke Tanja 2017 Towards a balanced view of the bacterial tree of life Microbiome 5 1 140 doi 10 1186 s40168 017 0360 9 PMC 5644168 PMID 29041958 Groups of Protists Boundless Biology courses lumenlearning com Retrieved 16 February 2021 a b Aguirre L E Ouyang L Elfwing A Hedblom M Wulff A and Inganas O 2018 Diatom frustules protect DNA from ultraviolet light Scientific reports 8 1 1 6 doi 10 1038 s41598 018 21810 2 nbsp Modified text was copied from this source which is available under a Creative Commons Attribution 4 0 International License De Tommasi E Congestri R Dardano P De Luca A C Manago S Rea I and De Stefano M 2018 UV shielding and wavelength conversion by centric diatom nanopatterned frustules Scientific Repor, wikipedia, wiki, book, books, library,

article

, read, download, free, free download, mp3, video, mp4, 3gp, jpg, jpeg, gif, png, picture, music, song, movie, book, game, games.