Marine protists
Marine protists are defined by their habitat as
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.
In contrast to the cells of prokaryotes, the cells of eukaryotes are highly organised. Plants, animals and fungi are usually
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]
Part of a series of overviews on |
Marine life |
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Background
"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
Trophic modes
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
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Type of protist | Description | Example | Some other examples | ||||
Plant-like | Autotrophic protists that make their own food without needing to consume other organisms, usually by 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
|
amoebae, ciliates and flagellates .
| |||
Fungal-like | slime nets |
Saprotrophic protists that get their food from the remains of organisms that have broken down and decayed
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Marine slime nets form labyrinthine networks of tubes in which amoeba without pseudopods can travel
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Marine lichen | |||
Mixotrophs | Various
(see below) |
osmotrophic protists that get their food from a combination of the above
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Euglena mutabilis, a photosynthetic flagellate | Many marine mixotrops are found among protists, particularly among ciliates and dinoflagellates[5] |
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Fossil diatom frustule from 32 to 40mya
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Radiolarian
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Single-celled alga, Gephyrocapsa oceanica
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Two dinoflagellates
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A single-celledendosymbiotically
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Euglenoid
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This ciliate is digesting cyanobacteria. The cytostome or mouth is at the bottom right.
External videos | |
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How microscopic hunters get their lunch | |
Euglenoids: Single-celled shapeshifters | |
How do protozoans get around? |
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Ciliate ingesting a diatom
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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
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]
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General types | Description | Example | Further examples | ||||
Bacterioplankton | Photoheterotrophic bacterioplankton
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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 | 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 | ||||||
aChloroplast (or plastid) retention = sequestration = enslavement. Some plastid-retaining species also retain other organelles and prey cytoplasm. |
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Tintinnid ciliate Favella
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Euglena mutabilis, a photosynthetic flagellate
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Zoochlorellae (green) living inside the ciliateStichotricha secunda
Protist locomotion
Another way of categorising protists is according to their mode of locomotion. Many unicellular protists, particularly protozoans, are
Protists according to how they move
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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 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 |
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 | |||||
Not motile | none
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Diatom | Coccolithophores, most diatoms, and non‐motile species of Phaeocystis[42] Among protozoans the parasitic Apicomplexa are non‐motile. |
Flagella
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
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.
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]
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Green algal flagellate (Chlamydomonas)
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Paramecium feeding on bacteria
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The ciliateOxytricha trifallaxwith cilia clearly visible
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Amoeba with ingested diatoms
External videos | |
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Paramecium: The White Rat of Ciliates |
Marine algae
Part of a series on |
Plankton |
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Dinoflagellates and diatoms are important components of marine algae and have their own sections below.
Not all algae are microscopic. Green, red and brown algae all have multicellular macroscopic forms that make up the familiar
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Chlamydomonas globosa, a unicellular green alga with twoflagellajust visible at bottom left
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Centric diatom
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Dinoflagellates
Unicellular organisms are usually microscopic, less than one tenth of a millimeter long. There are exceptions.
Diatoms
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
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Diatoms are one of the most common types of phytoplankton
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Their protective shells (frustles) are made of silicon
External videos | |
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Diatoms: Tiny factories you can see from space | |
Diatom 3D interference contrast |
Physically driven seasonal enrichments in surface nutrients favour
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Silicified frustule of a pennate diatom with two overlapping halves
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Guinardia delicatula, a diatom responsible fordiatom blooms in the North Sea [69]
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There are over 100,000 species of diatoms accounting for 25–45% of the ocean's primary production
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Linked diatoms
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Pennate diatom from an Arcticmeltpond, infected with two chytrid-like fungal pathogens. Scale bar = 10 µm.[70]
Coccolithophores
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]
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Algae bloomof Emiliania huxleyi off the southern coast of England
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Dinoflagellates
By trophic orientation dinoflagellates are all over the place. Some dinoflagellates are known to be
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]
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Gyrodinium, one of the few naked dinoflagellates which lack armour
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The dinoflagellate Protoperidinium extrudes a large feeding veil to capture prey
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Nassellarianradiolarians can be in symbiosis with dinoflagellates
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The dinoflagellate Dinophysis acuta
Dinoflagellates often live in
Some dinoflagellates are
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Tripos muelleri is recognisable by its U-shaped horns
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velvet disease in fish[84]
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Karenia brevis produces red tides highly toxic to humans[85]
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Red tide
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Noctiluca scintillans, a bioluminescent dinoflagellate[86]
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Ornithocercus heteroporus - prominent lists on display
Marine protozoans
Marine protozoans include
Radiolarians
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Like diatoms, radiolarians come in many shapes
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Also like diatoms, radiolarian shells are usually made of silicate
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Howeveracantharian radiolarians have shells made from strontium sulfatecrystals
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Cutaway schematic diagram of a spherical radiolarian shell
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Radiolarian geometry | |
Ernst Haeckel's radiolarian engravings |
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Cladococcus abietinus
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Cleveiplegma boreale
Foraminiferans
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]
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foraminiferans | |
Foraminiferal networks and growth |
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section showing chambers of a spiral foram
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Live Ammonia tepida streaming granular ectoplasm for catching food
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Group of planktonic forams
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Fossil nummulitid forams of various sizes from the Eocene
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TheEgyptian 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
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Naked amoeba showing food vacuoles and ingested diatom
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Shell or test of atestate amoeba, Arcellasp.
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Xenogenic testate amoeba covered in diatoms (from Penard's Amoeba Collection)
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Amoebas | |
Testate amoebas | |
Feeding amoebas |
Ciliates
Marine ciliates are major grazers of the phytoplankton.[97][98]
Phytoplankton primary production supports higher
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]
Although many ciliates are heterotrophs, a number of pelagic species are
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Tintinnopsis campanula
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The marine ciliate Strombidium rassoulzadegani
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Holophyra ovum
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Several taxa of ciliates interacting
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Blepharisma americanum swimming in a drop of pond water with other microorganisms
External videos | |
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Peritrich Ciliates | |
Conjugating protists |
Macroscopic protists
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The single-celledgiant amoeba has up to 1000 nucleiand reaches lengths of 5 mm
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testate amoeba which makes mud trails. Its diameter is up to 3.8 cm.[116]
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foraminiferan with an appearance and lifestyle that mimics a sponge, grows to 5 cm long.
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Thexenophyophore, another single-celled foraminiferan, lives in abyssal zones. It has a giant shell up to 20 cm across.[117]
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Giant kelp, a brown algae, is not a true plant, yet it is multicellular and can grow to 50m
Planktonic protists
Interactome
Interaction between microbial species has played important roles in evolution and speciation.
Despite their importance, understanding of microbial interactions in the ocean and other aquatic systems is rudimentary, and the majority of them are still unknown.
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.
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
Many protists have protective shells or
Diatom shells are called
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]
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
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
See also
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Further references
- 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. .
- Flynn, Kevin J.; Mitra, Aditee; Anestis, Konstantinos; Anschütz, Anna A.; Calbet, Albert; Ferreira, Guilherme Duarte; Gypens, Nathalie; Hansen, Per J.; John, Uwe; Martin, Jon Lapeyra; Mansour, Joost S.; Maselli, Maira; Medić, Nikola; Norlin, Andreas; Not, Fabrice; Pitta, Paraskevi; Romano, Filomena; Saiz, Enric; Schneider, Lisa K.; Stolte, Willem; Traboni, Claudia (2019). "Mixotrophic protists and a new paradigm for marine ecology: Where does plankton research go now?". Journal of Plankton Research. 41 (4): 375–391. hdl:10261/192145.
- Leles, Suzana Gonçalves; Polimene, Luca; Bruggeman, Jorn; Blackford, Jeremy; Ciavatta, Stefano; Mitra, Aditee; Flynn, Kevin John (2018). "Modelling mixotrophic functional diversity and implications for ecosystem function". Journal of Plankton Research. 40 (6): 627–642. .
- Keeling, Patrick J.; Campo, Javier del (2017). "Marine Protists Are Not Just Big Bacteria". Current Biology. 27 (11). Elsevier BV: R541–R549. S2CID 207052528.