A protist (/ˈproʊtɪst/) is any eukaryotic organism (one with cells containing a nucleus) that is not an animal, plant or fungus. The protists do not form a natural group, or clade, since they exclude certain eukaryotes; but, like algae or invertebrates, they are often grouped together for convenience. In some systems of biological classification, such as the popular five-kingdom scheme proposed by Robert Whittaker in 1969, the protists make up a kingdom called Protista, composed of "organisms which are unicellular or unicellular-colonial and which form no tissues".[A]
Besides their relatively simple levels of organization, protists do not necessarily have much in common. When used, the term "protists" is now considered to mean a paraphyletic assemblage of similar-appearing but diverse taxa (biological groups); these taxa do not have an exclusive common ancestor beyond being composed of eukaryotes and have different life cycles, trophic levels, modes of locomotion and cellular structures. In the classification system of Lynn Margulis, the term protist is reserved for microscopic organisms, while the more inclusive term Protoctista is applied to a biological kingdom that includes certain large multicellular eukaryotes, such as kelp, red algae and slime molds. Others use the term protist more broadly, to encompass both microbial eukaryotes and macroscopic organisms that do not fit into the other traditional kingdoms.
In cladistic systems (classifications based on common ancestry), there are no equivalents to the taxa Protista or Protoctista, both terms referring to a paraphyletic group that spans the entire eukaryotic tree of life. In cladistic classification, the contents of Protista are distributed among various supergroups (SAR, such as protozoa and some algae, Archaeplastida, such as land plants and some algae, Excavata, which are a group of unicellular organisms, and Opisthokonta, such as animals and fungi, etc.). "Protista", ''Protoctista'' and "Protozoa" are considered obsolete. However, the term "protist" continues to be used informally as a catch-all term for unicellular eukaryotic microorganisms. For example, the word "protist pathogen" may be used to denote any disease-causing microbe that is not bacteria, virus, viroid, prion, or metazoa.
Supergroups and typical phyla
|Cladistically included but traditionally excluded taxa|
The term protista was first used by Ernst Haeckel in 1866. Protists were traditionally subdivided into several groups based on similarities to the "higher" kingdoms such as:
Some protists, sometimes called ambiregnal protists, have been considered to be both protozoa and algae or fungi (e.g., slime molds and flagellated algae), and names for these have been published under either or both of the ICN and the ICZN. Conflicts, such as these – for example the dual-classification of Euglenids and Dinobryons, which are mixotrophic – is an example of why the kingdom Protista was adopted.
These traditional subdivisions, largely based on superficial commonalities, have been replaced by classifications based on phylogenetics (evolutionary relatedness among organisms). Molecular analyses in modern taxonomy have been used to redistribute former members of this group into diverse and sometimes distantly related phyla. For instance, the water molds are now considered to be closely related to photosynthetic organisms such as Brown algae and Diatoms, the slime molds are grouped mainly under Amoebozoa, and the Amoebozoa itself includes only a subset of "Amoeba" group, and significant number of erstwhile "Amoeboid" genera are distributed among Rhizarians and other Phyla.
However, the older terms are still used as informal names to describe the morphology and ecology of various protists. For example, the term protozoa is used to refer to heterotrophic species of protists that do not form filaments.
Among the pioneers in the study of the protists, which were almost ignored by Linnaeus except for some genera (e.g., Vorticella, Chaos, Volvox, Corallina, Conferva, Ulva, Chara, Fucus) were Leeuwenhoek, O. F. Müller, C. G. Ehrenberg and Félix Dujardin. The first groups used to classify microscopic organism were the Animalcules and the Infusoria. In 1818, the German naturalist Georg August Goldfuss introduced the word Protozoa to refer to organisms such as ciliates and corals. After the cell theory of Schwann and Schleiden (1838–39), this group was modified in 1848 by Carl von Siebold to include only animal-like unicellular organisms, such as foraminifera and amoebae. The formal taxonomic category Protoctista was first proposed in the early 1860s by John Hogg, who argued that the protists should include what he saw as primitive unicellular forms of both plants and animals. He defined the Protoctista as a "fourth kingdom of nature", in addition to the then-traditional kingdoms of plants, animals and minerals. The kingdom of minerals was later removed from taxonomy in 1866 by Ernst Haeckel, leaving plants, animals, and the protists (Protista), defined as a “kingdom of primitive forms”.
In 1938, Herbert Copeland resurrected Hogg's label, arguing that Haeckel's term Protista included anucleated microbes such as bacteria, which the term "Protoctista" (literally meaning "first established beings") did not. In contrast, Copeland's term included nucleated eukaryotes such as diatoms, green algae and fungi. This classification was the basis for Whittaker's later definition of Fungi, Animalia, Plantae and Protista as the four kingdoms of life. The kingdom Protista was later modified to separate prokaryotes into the separate kingdom of Monera, leaving the protists as a group of eukaryotic microorganisms. These five kingdoms remained the accepted classification until the development of molecular phylogenetics in the late 20th century, when it became apparent that neither protists nor monera were single groups of related organisms (they were not monophyletic groups).
Systematists today do not treat Protista as a formal taxon, but the term "protist" is still commonly used for convenience in two ways. The most popular contemporary definition is a phylogenetic one, that identifies a paraphyletic group: a protist is any eukaryote that is not an animal, (land) plant, or (true) fungus; this definition excludes many unicellular groups, like the Microsporidia (fungi), many Chytridiomycetes (fungi), and yeasts (fungi), and also a non-unicellular group included in Protista in the past, the Myxozoa (animal). Some systematists judge paraphyletic taxa acceptable, and use Protista in this sense as a formal taxon (as found in some secondary textbooks, for pedagogical purpose).
The other definition describes protists primarily by functional or biological criteria: protists are essentially those eukaryotes that are never multicellular, that either exist as independent cells, or if they occur in colonies, do not show differentiation into tissues (but vegetative cell differentiation may occur restricted to sexual reproduction, alternate vegetative morphology, and quiescent or resistant stages, such as cysts); this definition excludes many brown, multicellular red and green algae, which may have tissues.
The taxonomy of protists is still changing. Newer classifications attempt to present monophyletic groups based on morphological (especially ultrastructural), biochemical (chemotaxonomy) and DNA sequence (molecular research) information. However, there are sometimes discordances between molecular and morphological investigations; these can be categorized as two types: (i) one morphology, multiple lineages (e.g. morphological convergence, cryptic species) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity, multiple life-cycle stages).
Because the protists as a whole are paraphyletic, new systems often split up or abandon the kingdom, instead treating the protist groups as separate lines of eukaryotes. The recent scheme by Adl et al. (2005) does not recognize formal ranks (phylum, class, etc.) and instead treats groups as clades of phylogenetically related organisms. This is intended to make the classification more stable in the long term and easier to update. Some of the main groups of protists, which may be treated as phyla, are listed in the taxobox, upper right. Many are thought to be monophyletic, though there is still uncertainty. For instance, the Excavata are probably not monophyletic and the chromalveolates are probably only monophyletic if the haptophytes and cryptomonads are excluded.
Nutrition can vary according to the type of protist. Most eukaryotic algae are autotrophic, but the pigments were lost in some groups. Other protists are heterotrophic, and may present phagotrophy, osmotrophy, saprotrophy or parasitism. Some are mixotrophic. Some protists that do not have / lost chloroplasts/mitochondria have entered into endosymbiontic relationship with other bacteria/algae to replace the missing functionality. For example, Paramecium bursaria and Paulinella have captured a green alga (Zoochlorella) and a cyanobacterium respectively that act as replacements for chloroplast. Meanwhile, a protist, Mixotricha paradoxa that has lost its mitochondria uses endosymbiontic bacteria as mitochondria and ectosymbiontic hair-like bacteria (Treponema spirochetes) for locomotion.
Many protists are flagellate, for example, and filter feeding can take place where flagellates find prey. Other protists can engulf bacteria and other food particles, by extending their cell membrane around them to form a food vacuole and digesting them internally in a process termed phagocytosis.
|Nutritional type||Source of energy||Source of carbon||Examples|
|Photoautotrophs||Sunlight||Organic compounds or carbon fixation||Most algae|
|Chemoheterotrophs||Organic compounds||Organic compounds||Apicomplexa, Trypanosomes or Amoebae|
For most important cellular structures and functions of animal and plants, it can be found a heritage among protists.
Some species, for example Plasmodium falciparum, have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually. However, it is unclear how frequently sexual reproduction causes genetic exchange between different strains of Plasmodium in nature and most populations of parasitic protists may be clonal lines that rarely exchange genes with other members of their species.
Eukaryotes emerged in evolution more than 1.5 billion years ago. The earliest eukaryotes were likely protists. Although sexual reproduction is widespread among extant eukaryotes, it seemed unlikely until recently, that sex could be a primordial and fundamental characteristic of eukaryotes. A principal reason for this view was that sex appeared to be lacking in certain pathogenic protists whose ancestors branched off early from the eukaryotic family tree. However, several of these protists are now known to be capable of, or to recently have had the capability for, meiosis and hence sexual reproduction. For example, the common intestinal parasite Giardia lamblia was once considered to be a descendant of a protist lineage that predated the emergence of meiosis and sex. However, G. lamblia was recently found to have a core set of genes that function in meiosis and that are widely present among sexual eukaryotes. These results suggested that G. lamblia is capable of meiosis and thus sexual reproduction. Furthermore, direct evidence for meiotic recombination, indicative of sex, was also found in G. lamblia.
Trichomonas vaginalis, a parasitic protist, is not known to undergo meiosis, but when Malik et al. tested for 29 genes that function in meiosis, they found 27 to be present, including 8 of 9 genes specific to meiosis in model eukaryotes. These findings suggest that T. vaginalis may be capable of meiosis. Since 21 of the 29 meiotic genes were also present in G. lamblia, it appears that most of these meiotic genes were likely present in a common ancestor of T. vaginalis and G. lamblia. These two species are descendants of protist lineages that are highly divergent among eukaryotes, leading Malik et al. to suggest that these meiotic genes were likely present in a common ancestor of all eukaryotes.
Based on a phylogenetic analysis, Dacks and Roger proposed that facultative sex was present in the common ancestor of all eukaryotes.
This view was further supported by a study of amoebae by Lahr et al. Amoeba have generally been regarded as asexual protists. However, these authors describe evidence that most amoeboid lineages are anciently sexual, and that the majority of asexual groups likely arose recently and independently. Early researchers (e.g., Calkins) have interpreted phenomena related to chromidia (chromatin granules free in the cytoplasm) in amoeboid organisms as sexual reproduction.
Protists generally reproduce asexually under favorable environmental conditions, but tend to reproduce sexually under stressful conditions, such as starvation or heat shock. Oxidative stress, which is associated with the production of reactive oxygen species leading to DNA damage, also appears to be an important factor in the induction of sex in protists.
Some commonly found Protist pathogens such as Toxoplasma gondii are capable of infecting and undergoing asexual reproduction in a wide variety of animals – which act as secondary or intermediate host – but can undergo sexual reproduction only in the primary or definitive host (for example: felids such as domestic cats in this case).
Free-living Protists occupy almost any environment that contains liquid water. Many protists, such as algae, are photosynthetic and are vital primary producers in ecosystems, particularly in the ocean as part of the plankton. Protists make up a large portion of the biomass in both marine and terrestrial environments.
Some protists are significant parasites of animals (e.g.; five species of the parasitic genus Plasmodium cause malaria in humans and many others cause similar diseases in other vertebrates), plants (the oomycete Phytophthora infestans causes late blight in potatoes) or even of other protists. Protist pathogens share many metabolic pathways with their eukaryotic hosts. This makes therapeutic target development extremely difficult – a drug that harms a protist parasite is also likely to harm its animal/plant host. A more thorough understanding of protist biology may allow these diseases to be treated more efficiently. For example, the apicoplast (a nonphotosynthetic chloroplast but essential to carry out important functions other than photosynthesis) present in apicomplexans provides an attractive target for treating diseases caused by dangerous pathogens such as plasmodium.
Researchers from the Agricultural Research Service are taking advantage of protists as pathogens to control red imported fire ant (Solenopsis invicta) populations in Argentina. Spore-producing protists such as Kneallhazia solenopsae (recognized as a sister clade or the closest relative to the fungus kingdom now) can reduce red fire ant populations by 53–100%. Researchers have also been able to infect phorid fly parasitoids of the ant with the protist without harming the flies. This turns the flies into a vector that can spread the pathogenic protist between red fire ant colonies.
Many protists have neither hard parts nor resistant spores, and their fossils are extremely rare or unknown. Examples of such groups include the apicomplexans, most ciliates, some green algae (the Klebsormidiales), choanoflagellates, oomycetes, brown algae, yellow-green algae, Excavata (e.g., euglenids). Some of these have been found preserved in amber (fossilized tree resin) or under unusual conditions (e.g., Paleoleishmania, a kinetoplastid).
Others are relatively common in the fossil record, as the diatoms, golden algae, haptophytes (coccoliths), silicoflagellates, tintinnids (ciliates), dinoflagellates, green algae, red algae, heliozoans, radiolarians, foraminiferans, ebriids and testate amoebae (euglyphids, arcellaceans). Some are even used as paleoecological indicators to reconstruct ancient environments.
More probable eukaryote fossils begin to appear at about 1.8 billion years ago, the acritarchs, spherical fossils of likely algal protists. Another possible representative of early fossil eukaryotes are the Gabonionta.
Ancoracysta twista is a eukaryotic microorganism. It is a predatory protist that appears to be sister to the Haptista.Arcellinida
Arcellinid testate amoebae or Arcellinida, Arcellacean or lobose testate amoebae are single-celled protists partially enclosed in a simple test (shell).
Arcellinid testate amoebae are commonly found in soils, leaf litter, peat bogs and near/in fresh water. They use their pseudopodia, a temporary cell extension, for moving and taking in food. Like most amoebae, they are generally believed to reproduce asexually via binary fission. However a recent review suggests that sexual recombination may be the rule rather than the exception in amoeboid protists in general, including the Arcellinid testate amoebae.Cafeteria (bicosoecid)
Cafeteria is a genus of marine bicosoecid described in 1988 by Tom Fenchel and D. J. Patterson. It was created after the discovery of a new species Cafeteria roenbergensis, a tiny (5–10 µm) eukaryotic organism that is eaten by protozoa and small invertebrates. The name is meant to indicate the importance of the genus in the food web.
At least three species are recognised:
Cafeteria marsupialis Larsen & Patterson, 1990
Cafeteria minuta (Griessmann, 1913) Larsen & Patterson, 1990
Cafeteria roenbergensis Fenchel & D.J.Patterson, 1988Cercozoa
The Cercozoa are a group of single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, being defined by molecular phylogenies of rRNA and actin or polyubiquitin.Collodictyonidae
Collodictyonidae (also Diphylleidae) is a group of aquatic, unicellular eukaryotic organisms with two to four terminal flagella. They feed by phagocytosis, ingesting other unicellular organisms like algae and bacteria. The most remarkable fact of this clade is its uncertain position in the tree of life. A recent phylogenomic analysis places it either as sister group of the excavate protist Malawimonas or as sister group of the Bikonta clade, although both positions received considerably low branch support values. This means that Collodictyonidae is somewhere near the root of the Eukaryote tree, probably close to the unikont–bikont bifurcation.Endohelea
Endohelea is a proposed clade of eukaryotes that are related to Archaeplastida and the SAR supergroup.Garnia (Apicomplexa)
Garnia is a genus of parasitic alveolates belonging to the phylum Apicomplexia.Hatena arenicola
Hatena arenicola is a species of single-celled eukaryotes discovered in 2000, and first reported in 2005, It was discovered by Japanese biologists Noriko Okamoto and Isao Inouye at the
University of Tsukuba, and they gave the scientific description and formal name in 2006. The species is a flagellate, and can resemble a plant at one stage of its life, in which it carries a photosynthesizing alga inside itself, or an animal, acting as predator in another stage of its life. Researchers believe that this organism is in the process of secondary endosymbiosis, in which one organism is incorporated into another, resulting in a completely new life form.Labyrnavirus
Labyrnavirus is a monotypic genus of viruses in the order Picornavirales. Protist and labyrinthulomycetes serve as natural hosts, notably Aurantiochytrium. There is only one species in this genus: the type species Aurantiochytrium single-stranded RNA virus 01.Microbial cyst
A microbial cyst is a resting or dormant stage of a microorganism, usually a bacterium or a protist or rarely an invertebrate animal, that helps the organism to survive in unfavorable environmental conditions. It can be thought of as a state of suspended animation in which the metabolic processes of the cell are slowed down and the cell ceases all activities like feeding and locomotion. Encystment also helps the microbe to disperse easily, from one host to another or to a more favorable environment. When the encysted microbe reaches an environment favorable to its growth and survival, the cyst wall breaks down by a process known as excystation.
Unfavorable environmental conditions such as lack of nutrients or oxygen, extreme temperatures, lack of moisture and presence of toxic chemicals, which are not conducive for the growth of the microbe trigger the formation of a cyst.Microheliella maris
Microheliella maris is a protist species first described in 2015. It has a variety of unusual morphological characteristics which make its broader classification difficult. These include a centrosome with two concentric granular shells and axopodia much simpler structure than in visually similar protists.Noctiluca scintillans
Noctiluca scintillans, commonly known as the sea sparkle, and also published as Noctiluca miliaris, is a free-living, marine-dwelling species of dinoflagellate that exhibits bioluminescence when disturbed (popularly known as mareel). Its bioluminescence is produced throughout the cytoplasm of this single-celled protist, by a luciferin-luciferase reaction in thousands of spherically shaped organelles, called scintillons.Nomenclature codes
Nomenclature codes or codes of nomenclature are the various rulebooks that govern biological taxonomic nomenclature, each in their own broad field of organisms. To an end-user who only deals with names of species, with some awareness that species are assignable to families, it may not be noticeable that there is more than one code, but beyond this basic level these are rather different in the way they work.
The successful introduction of two-part names for species by Linnaeus was the start for an ever-expanding system of nomenclature. With all naturalists worldwide adopting this approach to thinking up names there arose several schools of thought about the details. It became ever more apparent that a detailed body of rules was necessary to govern scientific names. From the mid-nineteenth century onwards there were several initiatives to arrive at worldwide-accepted sets of rules. Presently nomenclature codes govern the naming of:
Algae, Fungi and Plants – International Code of Nomenclature for algae, fungi, and plants (ICN), which in July 2011 replaced the International Code of Botanical Nomenclature (ICBN) and the earlier International Rules of Botanical Nomenclature.
Animals – International Code of Zoological Nomenclature (ICZN)
Bacteria – International Code of Nomenclature of Prokaryotes (ICNP), which in 2008 replaced the International Code of Nomenclature of Bacteria (ICNB)
Cultivated plants – International Code of Nomenclature for Cultivated Plants (ICNCP)
Plant associations – International Code of Phytosociological Nomenclature (ICPN)
Viruses – The International Code of Virus Classification and Nomenclature (ICVCN); see also virus classificationPalpitomonas bilix
Palpitomonas bilix is a protist species first described in 2010.Perkinsozoa
Perkinsozoa is a proposed phylum of intracellular parasites in the superphylum Alveolata, which was suggested to account for the genus Perkinsus and other protist species that do not fit into existing Alveolata phyla.Perkinsus marinus
Perkinsus marinus is a species of alveolates belonging to the phylum Perkinsozoa. It is similar to a dinoflagellate. It is known as a prevalent pathogen of oysters, causing massive mortality in oyster populations. The disease it causes is known as dermo or perkinsosis, and is characterized by the degradation of oyster tissues. The genome of this species has been sequenced.The species originally was named Dermocystidium marinum.Predatory dinoflagellate
Predatory dinoflagellates are predatory heterotrophic or mixotrophic alveolates that derive some or most of their nutrients from digesting other organisms. About one half of dinoflagellates lack photosynthetic pigments and specialize in consuming other eukaryotic cells, and even photosynthetic forms are often predatory.Organisms that derive their nutrition in this manner include Oxyrrhis marina, which feeds phagocytically on phytoplankton, Polykrikos kofoidii, which feeds on several species of red-tide and/or toxic dinoflagellates, Ceratium furca, which is primarily photosynthetic but also capable of ingesting other protists such as ciliates, Cochlodinium polykrikoides, which feeds on phytoplankton, Gambierdiscus toxicus, which feeds on algae and produces a toxin that causes ciguatera fish poisoning when ingested, and Pfiesteria and related species such as Luciella masanensis, which feed on diverse prey including fish skin and human blood cells. Predatory dinoflagellates can kill their prey by releasing toxins or phagocytize small prey directly.Some predatory algae have evolved extreme survival strategies. For example, Oxyrrhis marina can turn cannibalistic on its own species when no suitable non-self prey is available, and Pfiesteria and related species have been discovered to kill and feed on fish, and since have been (mistakenly) referred to as carnivorous "algae" by the media.Protist (journal)
Protist is a peer-reviewed scientific journal focusing on protists. It was founded as Archiv für Protistenkunde by editor Fritz Shaudinn in 1902, and originally published by Gustav Fischer and later Jena. The journal is now published by Elsevier, and is currently edited by Michael Melkonian (Botanisches Institut, Universität zu Köln). The journal changed its name to Protist in 1998.Protozoan infection
Protozoan infections are parasitic diseases caused by organisms formerly classified in the Kingdom Protozoa. They include organisms classified in Amoebozoa, Excavata, and Chromalveolata.
Examples include Entamoeba histolytica, Plasmodium (some of which cause malaria), and Giardia lamblia. Trypanosoma brucei, transmitted by the tsetse fly and the cause of African sleeping sickness, is another example.The species traditionally collectively termed "protozoa" are not closely related to each other, and have only superficial similarities (eukaryotic, unicellular, motile, though with exceptions). The terms "protozoa" (and protist) are usually discouraged in the modern biosciences. However, this terminology is still encountered in medicine. This is partially because of the conservative character of medical classification, and partially due to the necessity of making identifications of organisms based upon appearances and not upon DNA.
Protozoan infections in animals may be caused by organisms in the sub-class Coccidia (disease: Coccidiosis) and species in the genus Besnoitia (disease: Besnoitiosis).
Several pathogenic protozoans appear to be capable of sexual processes involving meiosis (or at least a modified form of meiosis). Included among these protozoans are Plasmodium falciparum (malaria), Toxoplasma gondii (toxoplasmosis), Leishmania species (leishmaniases), Trypanosoma brucei (African sleeping sickness), Trypanosoma cruzi (Chagas disease) and Giardia intestinalis (giardiasis).
Extant Life phyla/divisions by domain
|Microscopic discoveries 1|