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Eukaryote
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"Eukaryotic cell" redirects here. For the journal, see Eukaryotic Cell (journal).
Eukaryotes
Temporal range: Rhyacian - Present 2100–0Ma
Had'n
Archean
Proterozoic
Pha.
Eukaryotes and some examples of their diversity
Scientific classification
Domain: Eukaryota
(Chatton, 1925) Whittaker & Margulis,1978
Supergroups[1] and kingdoms
Archaeplastida
Kingdom Plantae – Plants
SAR (Stramenopiles + Alveolata + Rhizaria)
Excavata
Amoebozoa
Opisthokonta
Kingdom Animalia – Animals
Kingdom Fungi
Eukaryotic organisms that cannot be classified under the kingdoms Plantae, Animalia or Fungi are sometimes grouped in the kingdom Protista.
A eukaryote (/juːˈkæri.oʊt/ or /juːˈkæriət/) is any organism whose cells have a cell nucleus and other organelles enclosed within membranes. Eukaryotes belong to the domain Eukaryota or Eukarya, and can be unicellular or multicellular organisms. The defining feature that sets eukaryotic cells apart from prokaryotic cells (Bacteria and Archaea) is that they have membrane-bound organelles, especially the nucleus, which contains the genetic material enclosed by the nuclear membrane.[2][3][4] The presence of a nucleus gives eukaryotes their name, which comes from the Greek εὖ (eu, "well" or "true") and κάρυον (karyon, "nut" or "kernel").[5] Eukaryotic cells also contain other membrane-bound organelles such as mitochondria and the Golgi apparatus. In addition, plants and algae contain chloroplasts. Unlike unicellular archaea and bacteria, eukaryotes may also be multicellular and include organisms consisting of many kinds of tissue and cell types.
Eukaryotes can reproduce both asexually through mitosis and sexually through meiosis and gamete fusion. In mitosis, one cell divides to produce two genetically identical cells. In meiosis, DNA replication is followed by two rounds of cell division to produce four haploid daughter cells. These act as sex cells (gametes). Each gamete has just one set of chromosomes, each a unique mix of the corresponding pair of parental chromosomes resulting from genetic recombination during meiosis.
The domain Eukaryota appears to be monophyletic, and makes up one of the domains of life in the three-domain system. The two other domains, Bacteria and Archaea, are prokaryotes[6] and have none of the above features. Eukaryotes represent a tiny minority of all living things.[7] However, due to their generally much larger size, their collective worldwide biomass is estimated to be about equal to that of prokaryotes.[7] Eukaryotes evolved approximately 1.6–2.1 billion years ago, during the Proterozoic eon.
Contents
[hide]
1 History
2 Cell features
2.1 Internal membrane
2.2 Mitochondria and plastids
2.3 Cytoskeletal structures
2.4 Cell wall
3 Differences among eukaryotic cells
3.1 Animal cell
3.2 Plant cell
3.3 Fungal cell
3.4 Other eukaryotic cells
4 Reproduction
5 Classification
5.1 Phylogeny
5.1.1 Five supergroups
5.1.2 Cavalier-Smith's tree
6 Origin of eukaryotes
6.1 Fossils
6.2 Relationship to Archaea
6.3 Endomembrane system and mitochondria
6.4 Hypotheses
6.4.1 Autogenous models
6.4.2 Chimeric models
7 See also
8 References
9 External links
History[edit]
Konstantin Mereschkowski proposed a symbiotic origin for cells with nuclei.
In 1905 and 1910, the Russian biologist Konstantin Mereschkowsky (1855–1921) argued three things about the origin of nucleated cells. Firstly, plastids were reduced cyanobacteria in a symbiosis with a non-photosynthetic (heterotrophic) host. Secondly, the host had earlier in evolution formed by symbiosis between an amoeba-like host and a bacteria-like ("micrococcal") cell that formed the nucleus. Thirdly, plants inherited photosynthesis from cyanobacteria.[8]
The concept of the eukaryote has been attributed to the French biologist Edouard Chatton (1883-1947). The terms prokaryote and eukaryote were more definitively reintroduced by the Canadian microbiologist Roger Stanier and the Dutch-American microbiologist C. B. van Niel in 1962. In his 1938 work Titres et Travaux Scientifiques, Chatton had proposed the two terms, calling the bacteria prokaryotes and organisms with nuclei in their cells eukaryotes. However he mentioned this in only one paragraph, and the idea was effectively ignored until Chatton's statement was rediscovered by Stanier and van Niel.[9]
In 1967, Lynn Margulis provided microbiological evidence for endosymbiosis as the origin of chloroplasts and mitochondria in eukaryotic cells in her paper, On the origin of mitosing cells.[10] In the 1970s, Carl Woese explored microbial phylogenetics, studying variations in 16S ribosomal RNA. This helped to uncover the origin of the eukaryotes and the symbiogenesis of two important eukaryote organelles, mitochondria and chloroplasts. In 1977, Woese and George Fox introduced a "third form of life", which they called the Archaebacteria; in 1990, Woese, Otto Kandler and Mark L. Wheeler renamed this the Archaea.[9]
In 1979, G. W. Gould and G. J. Dring suggested that the eukaryotic cell's nucleus came from the ability of Gram-positive bacteria to form endospores. In 1987 and later papers, Thomas Cavalier-Smith proposed instead that the membranes of the nucleus and endoplasmic reticulum first formed by infolding a prokaryote's plasma membrane. In the 1990s, several other biologists proposed endosymbiotic origins for the nucleus, effectively reviving Mereschkowsky's theory.[8]
Cell features[edit]
Eukaryotic cells are typically much larger than those of prokaryotes having a volume of around 10,000 times greater than the prokaryotic cell.[11] They have a variety of internal membrane-bound structures, called organelles, and a cytoskeleton composed of microtubules, microfilaments, and intermediate filaments, which play an important role in defining the cell's organization and shape. Eukaryotic DNA is divided into several linear bundles called chromosomes, which are separated by a microtubular spindle during nuclear division.
Internal membrane[edit]
The endomembrane system and its components
Eukaryote cells include a variety of membrane-bound structures, collectively referred to as the endomembrane system.[12] Simple compartments, called vesicles and vacuoles, can form by budding off other membranes. Many cells ingest food and other materials through a process of endocytosis, where the outer membrane invaginates and then pinches off to form a vesicle.[13] It is probable that most other membrane-bound organelles are ultimately derived from such vesicles. Alternatively some products produced by the cell can leave in a vesicle through exocytosis.
The nucleus is surrounded by a double membrane (commonly referred to as a nuclear membrane or nuclear envelope), with pores that allow material to move in and out.[14] Various tube- and sheet-like extensions of the nuclear membrane form the endoplasmic reticulum, which is involved in protein transport and maturation. It includes the rough endoplasmic reticulum where ribosomes are attached to synthesize proteins, which enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth endoplasmic reticulum.[15] In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles (cisternae), the Golgi apparatus.[16]
Vesicles may be specialized for various purposes. For instance, lysosomes contain digestive enzymes that break down most biomolecules in the cytoplasm.[17] Peroxisomes are used to break down peroxide, which is otherwise toxic. Many protozoans have contractile vacuoles, which collect and expel excess water, and extrusomes, which expel material used to deflect predators or capture prey. In higher plants, most of a cell's volume is taken up by a central vacuole, which mostly contains water and primarily maintains its osmotic pressure.
Mitochondria and plastids[edit]
Simplified structure of a mitochondrion
Mitochondria are organelles found in nearly all eukaryotes that provide energy to the cell by converting ingested sugars into ATP.[18] They have two surrounding membranes (each a phospholipid bi-layer), the inner of which is folded into invaginations called cristae, where aerobic respiration takes place. Mitochondria contain their own DNA. They are now generally held to have developed from endosymbiotic prokaryotes, probably proteobacteria. Protozoa and microbes that lack mitochondria, such as the amoebozoan Pelomyxa and metamonads such as Giardia and Trichomonas, have usually been found to contain mitochondrion-derived organelles, such as hydrogenosomes and mitosomes, and thus probably lost the mitochondria secondarily. They obtain energy by enzymatic action on nutrients absorbed from the environment. The metamonad Monocercomonoides has also acquired, by lateral gene transfer, a cytosolic sulphur mobilisation system which provides the clusters of iron and sulfur required for protein synthesis. The normal mitochondrial iron-sulphur cluster pathway has been lost secondarily.[19][20]
Plants and various groups of algae also have plastids. Plastids have their own DNA and are developed from endosymbionts, in this case cyanobacteria. They usually take the form of chloroplasts, which like cyanobacteria contain chlorophyll and produce organic compounds (such as glucose) through photosynthesis. Others are involved in storing food. Although plastids probably had a single origin, not all plastid-containing groups are closely related. Instead, some eukaryotes have obtained them from others through secondary endosymbiosis or ingestion.[21]
Endosymbiotic origins have also been proposed for the nucleus, and for eukaryotic flagella.[22]
Cytoskeletal structures[edit]
Main article: Cytoskeleton
Longitudinal section through the flagellum of Chlamydomonas reinhardtii
Many eukaryotes have long slender motile cytoplasmic projections, called flagella, or similar structures called cilia. Flagella and cilia are sometimes referred to as undulipodia,[23] and are variously involved in movement, feeding, and sensation. They are composed mainly of tubulin. These are entirely distinct from prokaryotic flagellae. They are supported by a bundle of microtubules arising from a centriole, characteristically arranged as nine doublets surrounding two singlets. Flagella also may have hairs, or mastigonemes, and scales connecting membranes and internal rods. Their interior is continuous with the cell's cytoplasm.
Microfilamental structures composed of actin and actin binding proteins, e.g., α-actinin, fimbrin, filamin are present in submembraneous cortical layers and bundles, as well. Motor proteins of microtubules, e.g., dynein or kinesin and actin, e.g., myosins provide dynamic character of the network.
Centrioles are often present even in cells and groups that do not have flagella, but conifers and flowering plants have neither. They generally occur in groups that give rise to various microtubular roots. These form a primary component of the cytoskeletal structure, and are often assembled over the course of several cell divisions, with one flagellum retained from the parent and the other derived from it. Centrioles produce the spindle during nuclear division.[24]
The significance of cytoskeletal structures is underlined in the determination of shape of the cells, as well as their being essential components of migratory responses like chemotaxis and chemokinesis. Some protists have various other microtubule-supported organelles. These include the radiolaria and heliozoa, which produce axopodia used in flotation or to capture prey, and the haptophytes, which have a peculiar flagellum-like organelle called the haptonema.
Cell wall[edit]
Main article: Cell wall
The cells of plants, fungi, and most chromalveolates have a cell wall, a layer outside the cell membrane, providing the cell with structural support, protection, and a filtering mechanism. The cell wall also prevents over-expansion when water enters the cell.[25]
The major polysaccharides making up the primary cell wall of land plants are cellulose, hemicellulose, and pectin. The cellulose microfibrils are linked via hemicellulosic tethers to form the cellulose-hemicellulose network, which is embedded in the pectin matrix. The most common hemicellulose in the primary cell wall is xyloglucan.[26]
Differences among eukaryotic cells[edit]
There are many different types of eukaryotic cells, though animals and plants are the most familiar eukaryotes, and thus provide an excellent starting point for understanding eukaryotic structure. Fungi and many protists have some substantial differences, however.
Animal cell[edit]
Structure of a typical animal cell
Riproduci file multimediale
3D Simulation of animal cell
Structure of a typical plant cell
All animals are eukaryotic. Animal cells are distinct from those of other eukaryotes, most notably plants, as they lack cell walls and chloroplasts and have smaller vacuoles. Due to the lack of a cell wall, animal cells can adopt a variety of shapes. A phagocytic cell can even engulf other structures.
There are many other types of cell. For instance, there are approximately 210 distinct cell types in the adult human body.
Plant cell[edit]
Main article: Plant cell
Plant cells are quite different from the cells of the other eukaryotic organisms. Their distinctive features are:
A large central vacuole (enclosed by a membrane, the tonoplast), which maintains the cell's turgor and controls movement of molecules between the cytosol and sap[27]
A primary cell wall containing cellulose, hemicellulose and pectin, deposited by the protoplast on the outside of the cell membrane; this contrasts with the cell walls of fungi, which contain chitin, and the cell envelopes of prokaryotes, in which peptidoglycans are the main structural molecules
The plasmodesmata, linking pores in the cell wall that allow each plant cell to communicate with other adjacent cells;[28] this is different from the functionally analogous system of gap junctions between animal cells.
Plastids, especially chloroplasts that contain chlorophyll, the pigment that gives plants their green color and allows them to perform photosynthesis
Bryophytes and seedless vascular plants lack flagellae and centrioles except in the sperm cells.[29] Sperm of cycads and Ginkgo are large, complex cells that swim with hundreds to thousands of flagellae.[30]
Conifers (Pinophyta) and flowering plants (Angiospermae) lack the flagellae and centrioles that are present in animal cells.
Fungal cell[edit]
Fungal Hyphae Cells
1- Hyphal wall 2- Septum 3- Mitochondrion 4- Vacuole 5- Ergosterol crystal 6- Ribosome 7- Nucleus 8- Endoplasmic reticulum 9- Lipid body 10- Plasma membrane 11- Spitzenkörper 12- Golgi apparatus
The cells of fungi are most similar to animal cells, with the following exceptions:[31]
A cell wall that contains chitin
Less definition between cells; the hyphae of higher fungi have porous partitions called septa, which allow the passage of cytoplasm, organelles, and, sometimes, nuclei. Primitive fungi have few or no septa, so each organism is essentially a giant multinucleate supercell; these fungi are described as coenocytic.
Only the most primitive fungi, chytrids, have flagella.
Other eukaryotic cells[edit]
Eukaryotes are a very diverse group, and their cell structures are equally diverse. Many have cell walls; many do not. Many have chloroplasts, derived from primary, secondary, or even tertiary endosymbiosis; and many do not. Some groups have unique structures, such as the cyanelles (unusual chloroplasts) of the glaucophytes,[32] the haptonema of the haptophytes, or the ejectosomes of the cryptomonads. Other structures, such as pseudopodia, are found in various eukaryote groups in different forms, such as the lobose amoebozoans or the reticulose foraminiferans.[33]
Reproduction[edit]
This diagram illustrates the twofold cost of sex. If each individual were to contribute to the same number of offspring (two), (a) the sexual population remains the same size each generation, where the (b) asexual population doubles in size each generation.
Cell division generally takes place asexually by mitosis, a process that allows each daughter nucleus to receive one copy of each chromosome. In most eukaryotes, there is also a process of sexual reproduction, typically involving an alternation between haploid generations, wherein only one copy of each chromosome is present, and diploid generations, wherein two copies of each chromosome are present, occurring through meiosis. There is considerable variation in this pattern.
Eukaryotes have a smaller surface area to volume ratio than prokaryotes, and thus have lower metabolic rates and longer generation times.[34] In some multicellular organisms, cells specialized for metabolism have enlarged surface areas, such as intestinal villi.
The evolution of sexual reproduction may be a primordial and fundamental characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and Roger proposed that facultative sex was present in the common ancestor of all eukaryotes.[35] A core set of genes that function in meiosis is present in both Trichomonas vaginalis and Giardia intestinalis, two organisms previously thought to be asexual.[36][37] Since these two species are descendants of lineages that diverged early from the eukaryotic evolutionary tree, it was inferred that core meiotic genes, and hence sex, were likely present in a common ancestor of all eukaryotes.[36][37] Eukaryotic species once thought to be asexual, such as parasitic protozoa of the genus Leishmania, have been shown to have a sexual cycle.[38] Also, evidence now indicates that amoebae, previously regarded as asexual, are anciently sexual and that the majority of present-day asexual groups likely arose recently and independently.[39]
Classification[edit]
Further information: wikispecies:Eukaryota
Phylogenetic and symbiogenetic tree of living organisms, showing a view of the origins of eukaryotes & prokaryotes
One hypothesis of eukaryotic relationships. The Opisthokonta group includes both animals (Metazoa) and fungi. Plants (Plantae) are placed in Archaeplastida.
A pie chart of described eukaryote species (except for Excavata), together with a tree showing possible relationships between the groups
In antiquity, the two lineages of animals and plants were recognized. They were given the taxonomic rank of Kingdom by Linnaeus. Though he included the fungi with plants with some reservations, it was later realized that they are quite distinct and warrant a separate kingdom, the composition of which was not entirely clear until the 1980s.[40] The various single-cell eukaryotes were originally placed with plants or animals when they became known. In 1830, the German biologist Georg A. Goldfuss coined the word protozoa to refer to organisms such as ciliates, and this group was expanded until it encompassed all single-celled eukaryotes, and given their own kingdom, the Protista, by Ernst Haeckel in 1866.[41][42] The eukaryotes thus came to be composed of four kingdoms:
Kingdom Protista
Kingdom Plantae
Kingdom Fungi
Kingdom Animalia
The protists were understood to be "primitive forms", and thus an evolutionary grade, united by their primitive unicellular nature.[42] The disentanglement of the deep splits in the tree of life only really started with DNA sequencing, leading to a system of domains rather than kingdoms as top level rank being put forward by Carl Woese, uniting all the eukaryote kingdoms under the eukaryote domain.[43] At the same time, work on the protist tree intensified, and is still actively going on today. Several alternative classifications have been forwarded, though there is no consensus in the field.
A classification produced in 2005 for the International Society of Protistologists,[44] which reflected the consensus of the time, divided the eukaryotes into six supposedly monophyletic 'supergroups'. However, in the same year (2005), doubts were expressed as to whether some of these supergroups were monophyletic, particularly the Chromalveolata,[45] and a review in 2006 noted the lack of evidence for several of the supposed six supergroups.[46] A revised classification in 2012[1] recognizes five supergroups.
Archaeplastida (or Primoplantae) Land plants, green algae, red algae, and glaucophytes
SAR supergroup Stramenopiles (brown algae, diatoms, etc.), Alveolata, and Rhizaria (Foraminifera, Radiolaria, and various other amoeboid protozoa).
Excavata Various flagellate protozoa
Amoebozoa Most lobose amoeboids and slime molds
Opisthokonta Animals, fungi, choanoflagellates, etc.
There are also smaller groups of eukaryotes whose position is uncertain or seems to fall outside the major groups[47] — in particular, Haptophyta, Cryptophyta, Centrohelida, Telonemia, Picozoa,[48] Apusomonadida, Ancyromonadida, Breviatea, and the genus Collodictyon.[49] Overall, it seems that, although progress has been made, there are still very significant uncertainties in the evolutionary history and classification of eukaryotes. As Roger & Simpson said in 2009 "with the current pace of change in our understanding of the eukaryote tree of life, we should proceed with caution."[50]
In an article published in Nature Microbiology in April 2016 the authors, "reinforced once again that the life we see around us – plants, animals, humans and other so-called eukaryotes – represent a tiny percentage of the world's biodiversity."[51] They classified eukaryote "based on the inheritance of their information systems as opposed to lipid or other cellular structures." Jillian F. Banfield of the University of California, Berkeley and fellow scientists used a super computer to generate a diagram of a new tree of life based on DNA from 3000 species including 2,072 known species and 1,011 newly reported microbial organisms, whose DNA they had gathered from diverse environments.[6][52] As the capacity to sequence DNA became easier, Banfield and team were able to do metagenomic sequencing—"sequencing whole communities of organisms at once and picking out the individual groups based on their genes alone."[51]
Phylogeny[edit]
The rRNA trees constructed during the 1980s and 1990s left most eukaryotes in an unresolved "crown" group (not technically a true crown), which was usually divided by the form of the mitochondrial cristae; see crown eukaryotes. The few groups that lack mitochondria branched separately, and so the absence was believed to be primitive; but this is now considered an artifact of long-branch attraction, and they are known to have lost them secondarily.[53][54]
As of 2011[update], there is widespread agreement that the Rhizaria belong with the Stramenopiles and the Alveolata, in a clade dubbed the SAR supergroup, so that Rhizaria is not one of the main eukaryote groups; also that the Amoebozoa and Opisthokonta are each monophyletic and form a clade, often called the unikonts.[55][56][57][58][59] Beyond this, there does not appear to be a consensus.
It has been estimated that there may be 75 distinct lineages of eukaryotes.[60] Most of these lineages are protists.
The known eukaryote genome sizes vary from 8.2 megabases (Mb) in Babesia bovis to 112,000–220,050 Mb in the dinoflagellate Prorocentrum micans, showing that the genome of the ancestral eukaryote has undergone considerable variation during its evolution.[60] The last common ancestor of all eukaryotes is believed to have been a phagotrophic protist with a nucleus, at least one centriole and cilium, facultatively aerobic mitochondria, sex (meiosis and syngamy), a dormant cyst with a cell wall of chitin and/or cellulose and peroxisomes.[60] Later endosymbiosis led to the spread of plastids in some lineages.
Five supergroups[edit]
A global tree of eukaryotes from a consensus of phylogenetic evidence (in particular, phylogenomics), rare genomic signatures, and morphological characteristics is presented in Adl et al. 2012[1] and Burki 2014/2016 with the Cryptophyta and picozoa having emerged within the Archaeplastida.[47][61] A similar inclusion of Glaucophyta, Cryptista (and also Haptista), Telonemia is recovered in Karnkowska et al with Rhodophyta/Telonemia as basal Archaeplastida.[62]
Eukaryotes
Diaphoretickes
Archaeplastida

Red algae (Rhodophyta)

picozoa

Glaucophyta

Green plants (Viridiplantae)

Cryptista

Haptista

Telonemia

SAR
Halvaria

Stramenopiles

Alveolata

Rhizaria

Discoba

Amorphea

Amoebozoa

Apusomonadida

Opisthokonta

Holomycota (inc. fungi)

Holozoa (inc. animals)

Hacrobia
In some analyses, the Hacrobia group (Haptophyta + Cryptophyta) is placed next to Archaeplastida,[55] but in other ones it is nested inside the Archaeplastida.[63] However, several recent studies have concluded that Haptophyta and Cryptophyta do not form a monophyletic group.[64] The former could be a sister group to the SAR group, the latter cluster with the Archaeplastida (plants in the broad sense).[65]
The division of the eukaryotes into two primary clades, bikonts (Archaeplastida + SAR + Excavata) and unikonts (Amoebozoa + Opisthokonta), derived from an ancestral biflagellar organism and an ancestral uniflagellar organism, respectively, had been suggested earlier.[63][66][67] A 2012 study produced a somewhat similar division, although noting that the terms "unikonts" and "bikonts" were not used in the original sense.[48]
A highly converged and congruent set of trees appears in Derelle et al (2015), Ren et al (2016), Yang et al (2017) and Cavalier-Smith (2015) including the supplementary information, resulting in a more conservative and consolidated tree. It is combined with some results from Cavalier-Smith for the basal Opimoda.[68][69][70][71][72] The main remaining controversies are the root, and the exact positioning of the Rhodophyta and the bikonts Rhizaria, Haptista, Cryptista, Picozoa and Telonemia, many of which may be eukaryote-eukaryote hybrids.[73] Archeaplastida developed the Chloroplasts probably by endosymbiosis of an ancestor related to current a Glaucophyte, Gloeomargarita lithophora.[74][75][76]
Eukaryotes
Diphoda
Diaphoretickes

Archaeplastida (+ Gloeomargarita lithophora)

Glaucophyta

Rhodophyta

Viridiplantae

Hacrobia

Haptista

Cryptista

SAR
Halvaria

Stramenopiles

Alveolata

Rhizaria

Discoba

Opimoda
Loukozoa
Metamonada, Malawimonas

Podiata
Varisulca
Diphyllatea, Rigifilida, Discocelida, Glissodiscea, Ancyromonas

Amorphea

Amoebozoa

Obazoa
Apusozoa

Apusomonadida

Mantamonadida

Breviata

Opisthokonta

Cavalier-Smith's tree[edit]
Thomas Cavalier-Smith 2010,[77] 2013,[78] 2014[79] and 2017[80] places the eukaryotic tree's root between Excavata (with ventral feeding groove supported by a microtubular root) and the grooveless Euglenozoa, and monophyletic Chromista, correlated to a single endosymbyotic event of capturing a red-algae.
Eukaryotes

Tsukubea

Discicristata

Euglenozoa

Percolozoa

Orthokaryota

Jakobea

Neokaryota
Corticata
Plantae

Glaucophytes

Rhodophytes

Viridiplantae

Chromista (+ Rhodophyte)

Hacrobia

SAR

Scotokaryota

Metamonada

Opimoda

Malawimonadea

Podiata

Planomonadida

Mantamonadia

Diphyllatea

Amorphea

Amoebozoa

Obazoa

Breviatea

Apusomonadida

Opisthokonta

Origin of eukaryotes[edit]
The three-domains tree and the Eocyte hypothesis[81]
Phylogenetic tree showing a possible relationship between the eukaryotes and other forms of life;[82] eukaryotes are colored red, archaea green and bacteria blue.
Fossils[edit]
The origin of the eukaryotic cell is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. The timing of this series of events is hard to determine; Knoll (2006) suggests they developed approximately 1.6–2.1 billion years ago. Some acritarchs are known from at least 1.65 billion years ago, and the possible alga Grypania has been found as far back as 2.1 billion years ago.[83] The Geosiphon-like fossil fungus Diskagma has been found in paleosols 2.2 billion years old.[84]
Organized living structures have been found in the black shales of the Palaeoproterozoic Francevillian B Formation in Gabon, dated at 2.1 billion years old. Eukaryotic life could have evolved at that time.[85] Fossils that are clearly related to modern groups start appearing an estimated 1.2 billion years ago, in the form of a red alga, though recent work suggests the existence of fossilized filamentous algae in the Vindhya basin dating back perhaps to 1.6 to 1.7 billion years ago.[86]
Biomarkers suggest that at least stem eukaryotes arose even earlier. The presence of steranes in Australian shales indicates that eukaryotes were present in these rocks dated at 2.7 billion years old.[87][88]
Relationship to Archaea[edit]
The nuclear DNA and genetic machinery of eukaryotes is more similar to Archaea than Bacteria, leading to a controversial suggestion that eukaryotes should be grouped with Archaea in the clade Neomura. In other respects, such as membrane composition, eukaryotes are similar to Bacteria. Three main explanations for this have been proposed:
Eukaryotes resulted from the complete fusion of two or more cells, wherein the cytoplasm formed from a eubacterium, and the nucleus from an archaeon,[89] from a virus,[90][91] or from a pre-cell.[92][93]
Eukaryotes developed from Archaea, and acquired their eubacterial characteristics through the endosymbiosis of a proto-mitochondrion of eubacterial origin.[94]
Eukaryotes and Archaea developed separately from a modified eubacterium.
Alternative proposals include:
The chronocyte hypothesis postulates that a primitive eukaryotic cell was formed by the endosymbiosis of both archaea and bacteria by a third type of cell, termed a chronocyte.[95]
The universal common ancestor (UCA) of the current tree of life was a complex organism that survived a mass extinction event rather than an early stage in the evolution of life. Eukaryotes and in particular akaryotes (Bacteria and Archaea) evolved through reductive loss,[96] so that similarities result from differential retention of original features.
Assuming no other group is involved, there are three possible phylogenies for the Bacteria, Archaea and Eukaryota in which each is monophyletic. These are labelled 1 to 3 in the table below. The eocyte hypothesis is a modification of hypothesis 2 in which the Archaea are paraphyletic. (The table and the names for the hypotheses are based on Harish and Kurland, 2017.[97])
Alternative hypotheses for the base of the tree of life
1 – Two empires
2 – Three domains
3 – Gupta
4 – Eocyte
UCA

Eukaryota

Archaea

Bacteria

UCA

Eukaryota

Archaea

Bacteria

UCA

Eukaryota

Bacteria

Archaea

UCA

Eukaryota

Archaea-Crenarchaeota

Archaea-Euryarchaeota

Bacteria

In recent years, most researchers have favoured either the three domains (3D) or the eocyte hypotheses. An rRNA analyses supports the eocyte scenario, apparently with the Eukaryote root in Excavata.[98][77][78][79][80] A cladogram supporting the eocyte hypothesis, positioning eukaryotes within Archaea, based on phylogenomic analyses of the Asgard archaea, is:[99][100]
Proteoarchaeota
TACK

Korarchaeota

Crenarchaeota

Aigarchaeota

Geoarchaeota

Thaumarchaeota

Bathyarchaeota

Asgard

Lokiarchaeota

Odinarchaeota

Thorarchaeota

Heimdallarchaeota

(+α─Proteobacteria)
Eukaryota

In this scenario, the Asgard group is seen as a sister taxon of the TACK group, which comprises Crenarchaeota (formerly named eocytes), Thaumarchaeota, and others.
In 2017, there has been significant pushback against this scenario, arguing that the eukaryotes did not emerge within the Archaea. Cunha et al. produced analyses supporting the three domains (3D) or Woese hypothesis (2 in the table above) and rejecting the eocyte hypothesis (4 above).[101] Harish and Kurland found strong support for the earlier two empires (2D) or Mayr hypothesis (1 in the table above), based on analyses of the coding sequences of protein domains. They rejected the eocyte hypothesis as the least likely.[102][97] A possible interpretation of their analysis is that the universal common ancestor (UCA) of the current tree of life was a complex organism that survived an evolutionary bottleneck, rather than a simpler organism arising early in the history of life.[96]
Endomembrane system and mitochondria[edit]
The origins of the endomembrane system and mitochondria are also unclear.[103] The phagotrophic hypothesis proposes that eukaryotic-type membranes lacking a cell wall originated first, with the development of endocytosis, whereas mitochondria were acquired by ingestion as endosymbionts.[104] The syntrophic hypothesis proposes that the proto-eukaryote relied on the proto-mitochondrion for food, and so ultimately grew to surround it. Here the membranes originated after the engulfment of the mitochondrion, in part thanks to mitochondrial genes (the hydrogen hypothesis is one particular version).[105]
In a study using genomes to construct supertrees, Pisani et al. (2007) suggest that, along with evidence that there was never a mitochondrion-less eukaryote, eukaryotes evolved from a syntrophy between an archaea closely related to Thermoplasmatales and an α-proteobacterium, likely a symbiosis driven by sulfur or hydrogen. The mitochondrion and its genome is a remnant of the α-proteobacterial endosymbiont.[106]
Hypotheses[edit]
Different hypotheses have been proposed as to how eukaryotic cells came into existence. These hypotheses can be classified into two distinct classes – autogenous models and chimeric models.
Autogenous models[edit]
An autogenous model for the origin of eukaryotes.
Autogenous models propose that a proto-eukaryotic cell containing a nucleus existed first, and later acquired mitochondria.[107] According to this model, a large prokaryote developed invaginations in its plasma membrane in order to obtain enough surface area to service its cytoplasmic volume. As the invaginations differentiated in function, some became separate compartments—giving rise to the endomembrane system, including the endoplasmic reticulum, golgi apparatus, nuclear membrane, and single membrane structures such as lysosomes.[108] Mitochondria are proposed to come from the endosymbiosis of an aerobic proteobacterium, and it's assumed that all the eukaryotic lineages that did not acquire mitochondria became extinct.[109] Chloroplasts came about from another endosymbiotic event involving cyanobacteria. Since all eukaryotes have mitochondria, but not all have chloroplasts, the serial endosymbiosis theory proposes that mitochondria came first.
Chimeric models[edit]
Chimeric models claim that two prokaryotic cells existed initially – an archaeon and a bacterium. These cells underwent a merging process, either by a physical fusion or by endosymbiosis, thereby leading to the formation of a eukaryotic cell. Within these chimeric models, some studies further claim that mitochondria originated from a bacterial ancestor while others emphasize the role of endosymbiotic processes behind the origin of mitochondria.
Based on the process of mutualistic symbiosis, the hypotheses can be categorized as – the serial endosymbiotic theory (SET),[110][111][112] the hydrogen hypothesis (mostly a process of symbiosis where hydrogen transfer takes place among different species),[113] and the syntrophy hypothesis.[114][115]
According to serial endosymbiotic theory (championed by Lynn Margulis), a union between a motile anaerobic bacterium (like Spirochaeta) and a thermoacidophilic crenarchaeon (like Thermoplasma which is sulfidogenic in nature) gave rise to the present day eukaryotes. This union established a motile organism capable of living in the already existing acidic and sulfurous waters. Oxygen is known to cause toxicity to organisms that lack the required metabolic machinery. Thus, the archaeon provided the bacterium with a highly beneficial reduced environment (sulfur and sulfate were reduced to sulfide). In microaerophilic conditions, oxygen was reduced to water thereby creating a mutual benefit platform. The bacterium on the other hand, contributed the necessary fermentation products and electron acceptors along with its motility feature to the archaeon thereby gaining a swimming motility for the organism. From a consortium of bacterial and archaeal DNA originated the nuclear genome of eukaryotic cells. Spirochetes gave rise to the motile features of eukaryotic cells. Endosymbiotic unifications of the ancestors of alpha-proteobacteria and cyanobacteria, led to the origin of mitochondria and plastids respectively. For example, Thiodendron has been known to have originated via an ectosymbiotic process based on a similar syntrophy of sulfur existing between the two types of bacteria – Desulphobacter and Spirochaeta. However, such an association based on motile symbiosis have never been observed practically. Also there is no evidence of archaeans and spirochetes adapting to intense acid-based environments.[116]
In the hydrogen hypothesis, the symbiotic linkage of an anaerobic and autotrophic methanogenic archaeon (host) with an alpha-proteobacterium (the symbiont) gave rise to the eukaryotes. The host utilized hydrogen (H2) and carbon dioxide (CO2) to produce methane while the symbiont, capable of aerobic respiration, expelled H2 and CO2 as byproducts of anaerobic fermentation process. The host's methanogenic environment worked as a sink for H2, which resulted in heightened bacterial fermentation. Endosymbiotic gene transfer (EGT) acted as a catalyst for the host to acquire the symbionts' carbohydrate metabolism and turn heterotrophic in nature. Subsequently, the host's methane forming capability was lost. Thus, the origins of the heterotrophic organelle (symbiont) are identical to the origins of the eukaryotic lineage. In this hypothesis, the presence of H2 represents the selective force that forged eukaryotes out of prokaryotes.[citation needed]
The syntrophy hypothesis was developed in contrast to the hydrogen hypothesis and proposes the existence of two symbiotic events. According to this theory, the origin of eukaryotic cells was based on metabolic symbiosis (syntrophy) between a methanogenic archaeon and a delta-proteobacterium. This syntrophic symbiosis was initially facilitated by H2 transfer between different species under anaerobic environments. In earlier stages, an alpha-proteobacterium became a member of this integration, and later developed into the mitochondrion. Gene transfer from a delta-proteobacterium to an archaeon led to the methanogenic archaeon developing into a nucleus. The archaeon constituted the genetic apparatus, while the delta-proteobacterium contributed towards the cytoplasmic features. This theory incorporates two selective forces at the time of nucleus evolution – (a) presence of metabolic partitioning to avoid the harmful effects of the co-existence of anabolic and catabolic cellular pathways, and (b) prevention of abnormal protein biosynthesis due to a vast spread of introns in the archaeal genes after acquiring the mitochondrion and losing methanogenesis.[citation needed]
See also[edit]
Biology portal
Book: Eukaryote
Evolution of sexual reproduction
List of sequenced eukaryotic genomes
Parikaryote
Prokaryote
Thaumarchaeota
Vault (organelle)
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Jump up ^ Cox, C. J.; Foster, P. G.; Hirt, R. P.; Harris, S. R.; Embley, T. M. (2008). "The archaebacterial origin of eukaryotes". Proc Natl Acad Sci USA. 105 (51): 20356–61. Bibcode:2008PNAS..10520356C. doi:10.1073/pnas.0810647105. PMC 2629343  . PMID 19073919.
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Jump up ^ Knoll, Andrew H.; Javaux, E.J; Hewitt, D.; Cohen, P. (29 June 2006). "Eukaryotic organisms in Proterozoic oceans". Philosophical Transactions of the Royal Society B. 361 (1470): 1023–38. doi:10.1098/rstb.2006.1843. PMC 1578724  . PMID 16754612.
Jump up ^ Retallack, G.J.; Krull, E.S.; Thackray, G.D. & Parkinson, D. H. (2013). "Problematic urn-shaped fossils from a Paleoproterozoic (2.2 Ga) paleosol in South Africa". Precambrian Research. 235: 71–87. doi:10.1016/j.precamres.2013.05.015.
Jump up ^ Albani, A. E.; Bengtson, S.; Canfield, D. E.; Bekker, A.; MacChiarelli, R.; Mazurier, A.; Hammarlund, E. U.; Boulvais, P.; Dupuy, J. J.; Fontaine, C.; Fürsich, F. T.; Gauthier-Lafaye, F. O.; Janvier, P.; Javaux, E.; Ossa, F. O.; Pierson-Wickmann, A. C.; Riboulleau, A.; Sardini, P.; Vachard, D.; Whitehouse, M.; Meunier, A. (2010). "Large colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr ago". Nature. 466 (7302): 100–104. Bibcode:2010Natur.466..100A. doi:10.1038/nature09166. PMID 20596019.
Jump up ^ Bengtson, S; Belivanova, V; Rasmussen, B; Whitehouse, M (2009). "The controversial "Cambrian" fossils of the Vindhyan are real but more than a billion years older". Proceedings of the National Academy of Sciences of the United States of America. 106 (19): 7729–34. Bibcode:2009PNAS..106.7729B. doi:10.1073/pnas.0812460106. PMC 2683128  . PMID 19416859.
Jump up ^ Brocks JJ, Logan GA, Buick R, Summons RE (August 1999). "Archean molecular fossils and the early rise of eukaryotes". Science. 285 (5430): 1033–6. doi:10.1126/science.285.5430.1033. PMID 10446042.
Jump up ^ Ward P (9 Feb 2008). "Mass extinctions: the microbes strike back". New Scientist: 40–3.
Jump up ^ Martin W (December 2005). "Archaebacteria (Archaea) and the origin of the eukaryotic nucleus". Curr. Opin. Microbiol. 8 (6): 630–7. doi:10.1016/j.mib.2005.10.004. PMID 16242992.
Jump up ^ Takemura M (May 2001). "Poxviruses and the origin of the eukaryotic nucleus". J. Mol. Evol. 52 (5): 419–25. doi:10.1007/s002390010171. PMID 11443345.
Jump up ^ Bell PJ (September 2001). "Viral eukaryogenesis: was the ancestor of the nucleus a complex DNA virus?". J. Mol. Evol. 53 (3): 251–6. doi:10.1007/s002390010215. PMID 11523012.
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^ Jump up to: a b Harish, Ajith; Tunlid, Anders & Kurland, Charles G. (2013), "Rooted phylogeny of the three superkingdoms", Biochimie, 95 (8): 1593–1604, doi:10.1016/j.biochi.2013.04.016
^ Jump up to: a b Harish, Ajith; Kurland, Charles G. "Akaryotes and Eukaryotes are independent descendants of a universal common ancestor". Biochimie. 138: 168–183. doi:10.1016/j.biochi.2017.04.013.
Jump up ^ Hug, Laura A.; Baker, Brett J.; Anantharaman, Karthik; Brown, Christopher T.; Probst, Alexander J.; Castelle, Cindy J.; Butterfield, Cristina N.; Hernsdorf, Alex W.; Amano, Yuki (2016-04-11). "A new view of the tree of life". Nature Microbiology. 1 (5). doi:10.1038/nmicrobiol.2016.48.
Jump up ^ 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. doi:10.1038/nature14447.
Jump up ^ Zaremba-Niedzwiedzka, Katarzyna; Caceres, Eva F.; Saw, Jimmy H.; Bäckström, Disa; Juzokaite, Lina; Vancaester, Emmelien; Seitz, Kiley W.; Anantharaman, Karthik; Starnawski, Piotr; Kjeldsen, Kasper U.; Stott, Matthew B.; Nunoura, Takuro; Banfield, Jillian F.; Schramm, Andreas; Baker, Brett J.; Spang, Anja; Ettema, Thijs J. G. (2017). "Asgard archaea illuminate the origin of eukaryotic cellular complexity". Nature. 541 (7637): 353–358. doi:10.1038/nature21031.
Jump up ^ Cunha, Violette Da; Gaia, Morgan; Gadelle, Daniele; Nasir, Arshan; Forterre, Patrick (12 June 2017). "Lokiarchaea are close relatives of Euryarchaeota, not bridging the gap between prokaryotes and eukaryotes". PLOS Genetics. 13 (6): e1006810. doi:10.1371/journal.pgen.1006810.
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This article incorporates public domain material from the NCBI document "Science Primer".[dead link]
External links[edit]
Wikispecies has information related to: Eukaryota
Eukaryotes (Tree of Life web site)
Eukaryote at the Encyclopedia of Life
[show]
v
t
e
Eukaryota
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Excavata
Metamonad
Malawimonadea
Discoba
Euglenozoa
Jakobea
Percolozoa
Tsukubea
Diaphoretickes
Archaeplastida
Glaucophyta
Rhodophyta
Viridiplantae or plantae sensu stricto
Chlorophyta
Streptophyta
Cryptista
Cryptophyceae
Endohelea
Goniomonadea
Kathablepharidea
Palpitea
Picomonadea
Telonemea
Haptista
Centrohelea
Pavlovophyceae
Prymnesiophyceae
Rappemonada
SAR
Rhizaria
Cercozoa
Retaria
Acantharea
Foraminifera
Polycystinea
Sticholonchea
Halvaria
Stramenopiles
(heterokonts)
Bicosoecea
Developayellales
Hyphochytrea
Ochrophyta
Opalinata
Opalomonadea
Peronosporomycetes
Pirsoniales
Platysulca
Sagenista
Alveolata
Acavomonadia
Ciliates
Colponemidia
Myzozoa
Apicomonadea
Apicomplexa
Chromerida
Dinoflagellata
Perkinsozoa
Protoalveolata
Incertae sedis
Kamera lens
Podiata
Diphyllatea
Discocelida
Glissodiscea
Rigifilida
Ancyromonas
Amorphea
Amoebozoa
Conosa (Archamoebae, Semiconosia)
Lobosa (Cutosea, Discosea, Tubulinea)
Obazoa
Apusozoa
Breviatea
Apusomonadida
Mantamonadida
Opisthokonta
Holozoa
Teretosporea
Mesomycetozoea
Corallochytrea
Filasterea
Choanoflagellates
Metazoa or Animals
Holomycota
Cristidiscoidea
True fungi
Opisthosporidia
Aphelida
Cryptomycota
Microsporidia
Incertae sedis
Parakaryon myojinensis
†Acritarcha
†Charnia
†Gakarusia
†Galaxiopsis
†Grypania
†Leptoteichos
Major kingdoms are underlined. See also: protist. Sources and alternative views: Wikispecies.
[show]
v
t
e
Excavata
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Discoba
Discicristata
Euglenozoa
Postgaardea
Postgaardida
Postgaardidae
Diplonemea
Diplonemida
Diplonemidae
Kinetoplastea
Bordnamonadida
Hemistasiida
Trypanophidida
Prokinetoplastida
Neobodonida
Parabodonida
Bodonida
Trypanosomatida
Euglenida
Petalomonadida
Sphenomonadales
Ploeotiida
Heteronematales
Rhabdomonadales
Peranemida
Euglenomorphales
Eutreptiales
Euglenales
Symbiontida
Calkinsia
Percolozoa
Pharyngomonadidea
Pharyngomonadida
Pharyngomonadidae
Heterolobosea
Lyromonadida
Plaesiobystridae
Gruberellidae
Psalteriomonadidae
Acrasida
Acrasidae
Guttulinopsidae
Schizopyrenida
Naegleriidae
Vahlkampfiidae
Neovahlkampfiidae
Paravahlkampfiidae
Euhyperamoebidae
Percolatea
Percolomonas
Stephanopogon
Jakobea
Jakobida
Andaluciina
Andaluciidae
Stygiellidae
Histonina
Moramonadidae
Jakobidae
Histionidae
Tsukubea
Tsukubamonas globosa
Loukozoa
Metamonad
Trichozoa
Parabasalia
Chilomitea
Chilomitida
Chilomitidae
Trichonymphea
Lophomonadida
Lophomonadidae
Trichonymphida
Hoplonymphidae
Spirotrichosomidae
Staurojoeninidae
Teranymphidae
Trichonymphidae
Trichomonadea
Trichomonadida
Hypotrichomonadidae
Tricercomitidae
Hexamastigidae
Honigbergiellidae
Trichomonadidae
Trichocovinida
Trichocovinidae
Tritrichomonadida
Dientamoebidae
Monocercomonadidae
Simplicimonadidae
Tritrichomonadidae
Spirotrichonymphida
Spirotrichonymphidae
Cristamonadida
Calonymphidae
Devescovinidae
Fornicata
Eopharingea
Caviomonadidae
Diplomonadida
Giardiidae
Octomitidae
Spironucleidae
Hexamitidae
Retortamonadida
Retortamonadidae
Carpediemonadea
Carpediemonadida
Carpediemonadidae
Other
Kipferliidae
Dysnectida
Dysnectidae
Chilomastigida
Chilomastigidae
Anaeromonada
Anaeromonadea
Paratrimastigidae
Trimastigida
Trimastigidae
Oxymonadida
Polymastigidae
Saccinobaculidae
Pyrsonymphidae
Streblomastigidae
Oxymonadidae
Neolouka
Malawimonadea
Malawimonadida
Malawimonadidae
[show]
v
t
e
Classification of Archaeplastida / Plantae sensu lato
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Rhodophyta
(red algae)
Cyanidiophyceae
Porphyridiophyceae
Compsopogonophyceae
Stylonematophyceae
Rhodellophyceae
Bangiophyceae
Florideophyceae
Glaucocystophyta
(glaucophytes)
Glaucocystophyceae
Glaucocystis
Cyanophora
Gloeochaete
Viridiplantae
(green algae
& land plants)
Chlorophyta
Palmophyllales
Nephroselmidophyceae
Prasinophyceae
Pseudoscourfieldiales
Pyramimonadophyceae
Scourfieldiales
Pedinophyceae
Chlorodendrophyceae
UTC clade
Ulvophyceae
Trebouxiophyceae
Chlorophyceae
Streptophyta
(charophytes,
& land plants)
Mesostigmatophyceae
Chlorokybophyceae
Klebsormidiophyceae
Phragmo-
plastophyta
Charophyceae
Coleochaetophyceae
Zygnematophyceae
Embryophyta
(land plants)
Bryophytes
(non-vascular)
Marchantiophyta
Anthocerotophyta
Bryophyta "Moss"
†Horneophytopsida
Tracheophyta
(vascular)
Lycopodiophyta
(microphylls)
†Zosterophyllopsida
†Sawdoniales
Isoetopsida
Lycopodiopsida
Euphyllophyta
(megaphylls)
Moniliformopses
(ferns)
†Cladoxylopsida
†Stauropteridales
†Zygopteridales
Equisetopsida
Psilotopsida
Marattiopsida
Filicopsida
Spermatophyta
(seed plants)
†Seed ferns
Gymnosperms
Gnetopsida
Pinopsida
Cycadopsida
Ginkgoopsida
Angiosperms or flowering plants
Amborellales
Nymphaeales
Austrobaileyales
Magnoliids
Monocots
Eudicots
Other
†Trimerophytopsida
†Progymnosperm
Other
†Rhyniopsida
† = extinct. See also the list of plant orders.
[show]
v
t
e
Eukaryota: Hacrobia
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Cryptista
Corbihelia
Endohelea
Microhelida
Microhelidae
Heliomonadida
Heliomonadidae
Picomonadea
Picomonadida
Picomonas
Telonemea
Telonemida
Telonemidae
Cryptophyta
Palpitea
Palpitomonadida
Palpitomonas
Katablepharidea
Katablepharida
Katablepharidae
Goniomonadea
Goniomonadida
Goniomonas
Cryptophyceae
Tetragonidiales
Tetragonidiaceae
Cryptomonadales
Cryptomonadaceae
GPTgc (Plagioselmis)
other (Cryptomonas/Chilomonas/Goniomonas)
Hemiselmidaceae
CKHgc (Hemiselmis)
Pyrenomonadales
Chroomonadaceae
CKHgc (Chroomonas, Komma)
other (Falcomonas)
Geminigeraceae
GHgc (Guillardia, Hanusia)
GPTgc (Geminigera, Teleaulax)
other (Proteomonas)
Pyrenomonadaceae
RRSgc (Pyrenomonas, Rhinomonas, Rhodomonas, Storeatula)
Haptista
Centroheliozoa
Centrohelea
Pterocystida
Choanocystidae
Oxnerellidae
Heterophryidae
Pterocystidae
Acanthocystida
Marophryidae
Raphidiophryidae
Acanthocystidae
Haptophyta
Rappephyceae
Rappemonadales
Rappemonas
Pavlovophyceae
Pavlovales
Pavlovaceae
Prymnesiophyceae
Coccolithales (Pleurochrysis, Coccolithus)
Prymnesiales (Chrysochromulina, Prymnesium)
Isochrysidales (Chrysotila, Dicrateria, Emiliania, Gephyrocapsa, Isochrysis)
See also: coccolithophores
[show]
v
t
e
Eukaryota: SAR: Heterokonta or Stramenopiles
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Ochrophyta
(mostly
photosynthetic)
Phaeista
Hypogyristea
Pinguiophyceae
Pelagophyceae
Dictyochophyceae/
Actinochrysophyceae/
Axodines
Dictyochales/Silicoflagellates
Rhizochromulinales
Florenciellales
Actinodines
Pedinellales
Ciliophryida
Actinophryida
Chrysista
Chrysophyceae,
golden algae
Chromulinales
Chrysosphaerales
Hibberdiales
Hydrurales
Phaeothamniales
Eustigmatophyceae
Eustigmataceae
Monodopsidaceae
Pseudocharaciopsidaceae
Phaeophyceae,
brown algae
Ascoseirales
Cutleriales
Desmarestiales
Dictyotales
Discosporangiales
Ectocarpales
Fucales
Ishigeales
Laminariales
Nemodermatales
Onslowiales
Ralfsiales
Scytosiphonales
Scytothamnales
Sphacelariales
Sporochnales
Syringodermatales
Tilopteridales
Phaeothamniophyceae
Phaeothamniales
Pleurochloridellales
Raphidophyceae
Chattonella, Fibrocapsa, Gonyostomum, Haramonas, Heterosigma, Vacuolaria
Synurophyceae
Heterogloeales
Ochromonadales
Rhizochloridales
Synurales
Xanthophyceae,
yellow-green algae
Botrydiales
Mischococcales
Tribonematales
Vaucheriales
Khakista
Bolidophyceae
Bolidomonas
Bacillariophyta,
diatoms
Coscinodiscophyceae
Biddulphiophycidae
Chaetocerotophycidae
Corethrophycidae
Coscinodiscophycidae
Cymatosirophycidae
Lithodesmiophycidae
Rhizosoleniophycidae
Thalassiosirophycidae
Bacillariophyceae
Bacillariales
Naviculales
Fragilariophyceae
Fragilariales
Heterotrophic
groups
Pseudofungi
Oomycetes
Hyphochytriomycetes
Bigyra
Bigyromonadea
Bicosoecea
Sagenista
Labyrinthulomycetes
Eogyrea
Opalinata/Slopalinida
Opalinidae
Proteromonadidae
Blastocystis
[show]
v
t
e
Eukaryota: SAR: Alveolata
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Acavomonidia
Acavomonadea
Acavomonadida (Acavomonas)
Myzozoa
Apicomplexa
Aconoidasida
Haemospororida
Garniidae (Garnia)
Haemoproteidae (Haemoproteus)
Leucocytozoidae (Leucocytozoon)
Plasmodiidae (Plasmodium)
Piroplasmida
Babesiidae (Babesia)
Theileriidae (Theileria)
Conoidasida
Coccidia
Agamococcidiorida
Gemmocystidae (Gemmocystis)
Rhytidocystidae (Rhytidocystis)
Eucoccidiorida
Adeleorina
Adeleidae
Dactylosomatidae (Babesiosoma, Dactylosoma)
Haemogregarinidae (Haemogregarina)
Hepatozoidae (Hepatozoon)
Karyolysidae (Karyolysus)
Klossiellidae (Klossiella)
Legerellidae (Legerella)
Eimeriorina
Aggregatidae (Aggregata, Grasseella, Merocystis, Ovivora, Pseudoklossia, Selysina)
Atoxoplasmatidae
Barrouxiidae
Calyptosporiidae
Caryotrophidae
Cryptosporidiidae (Cryptosporidium)
Eimeriidae (Cyclospora, Eimeria, Isospora)
Elleipsisomatidae
Lankesterellidae
Selenococcidiidae
Sarcocystidae
Sarcocystinae (Frenkelia, Sarcocystis)
Toxoplasmatinae (Besnoitia, Hammondia, Hyaloklossia, Nephroisospora, Neospora, Toxoplasma)
Ixorheorida
Ixorheidae (Ixorheis)
Protococcidiorida
Angeiocystidae (Angeiocystis)
Eleutheroschizonidae (Coelotropha, Defretinella, Eleutheroschizon)
Grelliidae (Coelotropha, Grellia)
Mackinnoniidae (Mackinnonia)
Myriosporidae (Myriosporides, Myriospora)
Gregarinia
Archigregarinorida
Exoschizonidae (Exoschizon)
Selenidioididae (Filipodium, Merogregarina, Meroselenidium, Platyproteum, Selenidioides, Veloxidium)
Eugregarinorida
Aseptatorina
Aikinetocystidae
Allantocystidae
Diplocystidae
Enterocystidae
Ganymedidae
Lecudinidae
Monocystidae (Monocystinae, Oligochaetocystinae, Rhynchocystinae, Stomatophorinae, Zygocystinae)
Schaudinnellidae
Selenidiidae
Thiriotiidae
Urosporidae
Blastogregarinorina
Siedleckiidae (Siedleckia)
Septatorina
Fusionicae (Fusionidae)
Gregarinicae (Cephaloidophoridae, Cephalolobidae, Didymophoridae, Gregarinidae, Hirmocystidae, Metameridae, Uradiophoridae)
Porosporicae (Porosporidae)
Stenophoricae (Acutidae, Amphiplatysporidae, Brustiophoridae, Cnemidosporidae, Dactylophoridae, Leidyanidae, Monoductidae, Monoicidae, Sphaerocystidae, Stenophoridae, Trichorhynchidae)
Stylocephaloidea (Actinocephalidae, Stylocephalidae)
Blabericolidae
Neogregarinorida
Schizogregarinina (Caulleryellidae, Ophryocystidae)
Gigaductidae (Gigaductus)
Lipotrophidae (Apicystis, Farinocystis, Lipotropha, Lipocystis, Mattesia, Menzbieria)
Schizocystidae (Lymphotropha, Machadoella, Schizocystis)
Syncystidae (Syncystis)
Apicomonadea
Chromerida
Chromeraceae (Chromera velia)
Vitrellaceae (Vitrella brassicaformis)
Colpodellida
Colpodellidae (Colpodella)
Voromonadida
Alphamonadidae (Alphamonas)
Voromonadidae (Voromonas)
Dinoflagellata
Dinokaryota
With a theca: Peridiniales (Pfiesteria, Peridinium)
Gonyaulacales (Ceratium, Gonyaulax)
Prorocentrales (Prorocentrum)
Dinophysiales (Dinophysis, Histioneis, Ornithocercus, Oxyphysis)
Without theca: Gymnodiniales (Gymnodinium, Karenia, Karlodinium, Amphidinium)
Suessiales (Polarella, Symbiodinium)
Noctilucea
Noctilucales (Noctiluca)
Syndinea
Syndiniales: Amoebophryaceae (Amoebophyra)
Duboscquellaceae (Duboscquella)
Syndiniaceae (Hematodinium, Syndinium)
Other
Acrocoelidae (Acrocoelus)
Ichthyodinium
Oxyrrhinaceae (Oxyrrhis)
Pronoctilucidae (Pronoctiluca)
Psammosidae (Psammosa)
Perkinsozoa
Perkinsea
Perkinsidae (Perkinsus)
Phagodinida (Phagodinium)
Rastromonadida (Parvilucifera, Rastrimonas)
Protoalveolata
Ellobiopsea
Ellobiopsidae (Elliobiocystis, Ellobiopsis, Parallobiopsis, Thalassomyces, Rhizellobiopsis)
Myzomonadea
Algovorida
Algovoridae (Algovora)
Chilovorida
Chilovoridae (Chilovora)
Squirmidea
Squirmidae (Platyproteum, Filipodium)
Ciliophora
Postciliodesmatophora
Heterotrichea (Stentor, Climacostomum, Blepharisma)
Karyorelictea (Loxodes, Tracheloraphis)
Intramacronucleata
Spirotrichea (Stylonychia, Euplotes)
Litostomatea (Balantidium, Dileptus)
Phyllopharyngea (Chilodonella, Tokophrya)
Armophorea (Metopus)
Nassophorea (Nassula)
Colpodea (Colpoda)
Oligohymenophorea (Tetrahymena, Ichthyophthirius, Vorticella, Paramecium)
Plagiopylea (Plagiopyla)
Prostomatea (Coleps, Holophrya)
Protocruziea (Protocruzia)
Cariacotrichea (Cariacothrix caudata)
Mesodiniea (Mesodinium, Myrionecta)
Colponemidia
Colponemadea
Colponemadida (Colponema)
[show]
v
t
e
Amoebozoa
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Lobosa
Cutosea
Squamocutida
Squamamoebidae
Sapocribridae
Glycopoda
Discosea
Hyalodiscida
Hyalodiscidae
Centramoebia
Centramoebida
Acanthamoebidae
Balamuthiidae
Cochliopodiida
Parvamoebina
Parvamoebidae
Tectiferina
Cochliopodiidae
Pellitina
Pellitidae
Goceviidae
Flabellinia
Thecamoebida
Stenamoebidae
Thecamoebidae
Mayorellidae
Mayorella
Dermamoebida
Dermamoebidae
Glycostylida
Vannellina
Vannellidae
Discamoebidae
Stygamoebina
Stygamoebidae
Dactylopodina
Paramoebidae
Vexilliferidae
Tubulinea
Echinamoebida
Echinamoebidae
Vermamoebidae
Leptomyxida
Flabellina
Flabellulidae
Leptomyxina
Gephyramoebidae
Leptomyxidae
Neolobosia
Trichosida
Trichosphaerium
Eulobosia
Euamoebida
Nolandellidae
Amoebidae
Hartmannellidae
Arcellinida
Phryganellina
Eulobosina
Centropyxidae
Difflugidae
Conosa
Archamoebae
incertae sedis
Tricholimacidae
Endamoebidae
Entamoebida
Entamoebidae
Pelobiontida
Pelomyxina
Pelomyxidae
Mastigamoebina
Mastigamoebidae
Semiconosia
Variosea
(paraphyletic)
Holomastigida
Phalansteriida
Artodiscida
Varipodida
Mycetozoa
Myxogastria
Echinosteliopsida
Ceratiomyxida
Lucisporidia
Liceida
Trichiida
Echinostelida
Echinosteliidae
Clastodermidae
Fuscisporida
Lamprodermina
Stemonitina
Physarina
Others
Protosteliales
Protosporangiida
Dictyosteliida
Incertae sedis
Multicilia
[show]
v
t
e
Opisthokonta
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Holomycota
Cristidiscoidea
Nucleariida
Nucleariidae
Fonticulida
Fonticulaceae
O+F
Opisthosporidia
Aphelida
Aphelidea
Aphelidida
C+M
Cryptomycota
Rozellidea
Rozellida
Microsporidia
Metchnikovellea
Metchnikovellida
Microsporea
Chytridiopsida
Glugeida
Meiodihaplophasida
Dissociodihaplophasida
True Fungi
Neocallimastigomycota
Chytridiomycota
Blastocladiomycota
Olpidiomycota
Entomophthoromycota
Kickxellomycota
Mucoromycota
Glomeromycota
Entorrhizomycota
Ascomycota
Basidiomycota
Holozoa
Teretosporea
Corallochytrea
Corallochytriida
Corallochytriidae
Ichthyosporea
Dermocystida
Rhinosporidiaceae
Ichthyophonida
Sphaeroformina
Psorospermidae
Piridae
Creolimacidae
Trichomycina
Ichthyophonidae
Amoebidiidae
Palavasciaceae
Parataeniellaceae
Eccrinaceae
Filozoa
Filasterea
Ministeriida
Capsasporidae
Ministeriidae
Apoikozoa
Choanoflagellate
Acanthoecida
Stephanoecidae
Acanthoecidae
Craspedida
Codonosigidae
Salpingoecidae
Metazoa (Animalia)
Porifera
Ctenophora
Placozoa
Cnidaria
Xenacoelomorpha
Deuterostome
chordates
echinoderms
Protostome
Ecdysozoa inc. arthropods and nematodes
Spiralia inc. molluscs and annelids
Sources and alternative views: Wikispecies.
[show]
v
t
e
Extant Animal phyla
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
A
n
i
m
a
l
i
a
Ctenophora (comb jellies)
Parazoa
Porifera (sponges)
ParaHoxozoa
(Epitheliozoa)
Placozoa (Trichoplax)
Planulozoa
(Eumetazoa)
Cnidaria (jellyfish and relatives)
Bilateria
(Triploblasts)
(see below↓)
[hide]
Bilateria
Xenacoelomorpha
Xenoturbellida (Xenoturbella)
Acoelomorpha
acoels
nemertodermatids
N
e
p
h
r
o
z
o
a
Deuterostomia
Chordata
lancelets
tunicates
craniates / vertebrates
Ambulacraria
Echinodermata (starfish and relatives)
Hemichordata
acorn worms
pterobranchs
P
r
o
t
o
s
t
o
m
i
a
Ecdysozoa
Scalidophora
Kinorhyncha (mud dragons)
Priapulida (penis worms)
Loricifera
N+P
Nematoida
Nematoda (roundworms)
Nematomorpha (horsehair worms)
Panarthropoda
Arthropoda (arthropods)
Tardigrada (waterbears)
Onychophora (velvet worms)
S
p
i
r
a
l
i
a
Gnathifera¹
Chaetognatha (arrow worms)
Gnathostomulida (jaw worms)
Micrognathozoa (Limnognathia)
Syndermata
Rotifera
Acanthocephala
Platytrochozoa
R+M
Mesozoa
Orthonectida
Dicyemida or Rhombozoa
Rouphozoa¹
Platyhelminthes (flatworms)
Gastrotricha (hairybacks)
Lophotrochozoa
Cycliophora (Symbion)
Mollusca (molluscs)
A+N
Annelida (ringed worms)
Nemertea (ribbon worms)
Lophophorata
Bryozoa
Entoprocta or Kamptozoa
Ectoprocta (moss animals)
Brachiozoa
Brachiopoda (lamp shells)
Phoronida (horseshoe worms)
Major groups
within phyla
Sponges
Calcareous
Hexactinellid
Demosponge
Homoscleromorpha
Cnidarians
Anthozoa inc. corals
Medusozoa inc. jellyfish
Myxozoa
Vertebrates
Jawless fish
Cartilaginous fish
Bony fish
Amphibians
Reptiles/Birds
Mammals
Echinoderms
Sea lilies
Asterozoa inc. starfish
Echinozoa
Nematodes
Chromadorea
Enoplea
Secernentea
Arthropods
Chelicerates/Arachnids
Myriapods
Crustaceans
Hexapods/Insects
Platyhelminths
Turbellaria
Trematoda
Monogenea
Cestoda
Bryozoans
Phylactolaemata
Stenolaemata
Gymnolaemata
Annelids
Polychaetes
Clitellata
Echiura
Molluscs
Gastropods
Cephalopods
Bivalves
Chitons
Tusk shells
Phyla with ≥5000 extant species bolded
See also
Parazoa and Eumetazoa
Monoblastozoa (nomen dubium)
¹Platyzoa
[show]
v
t
e
Opisthokont: True fungi classification, fungal orders
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Dikarya
Ascomycota
(sac fungi)
Pezizomycotina
Leotiomyceta
Dothideomyceta
Coniocybomycetes
Lichinomycetes
Arthoniomycetes
Dothideomycetes
Eurotiomycetes
Lecanoromycetes
Sordariomyceta
Xylonomycetes
Geoglossomycetes
Leotiomycetes
Laboulbeniomycetes
Sordariomycetes
Other
Orbiliomycetes
Pezizomycetes
Saccharomycotina
Saccharomycetes
Taphrinomycotina
Archaeorhizomycetes
Neolectomycetes
Pneumocystidomycetes
Schizosaccharomycetes
Taphrinomycetes
Basidiomycota
(with basidia)
Pucciniomycotina
Tritirachiomycetes
Mixiomycetes
Agaricostilbomycetes
Cystobasidiomycetes
Microbotryomycetes
Classiculomycetes
Cryptomycocolacomycetes
Atractiellomycetes
Pucciniomycetes
Ustilaginomycotina
Monilielliomycetes
Malasseziomycetes
Ustilaginomycetes
Exobasidiomycetes
Agaricomycotina
Hymenomycete
Dacrymycetales
Agaricomycetes
Other
Wallemiomycetes
Bartheletiomycetes
Tremellomycetes
Entorrhizomycota
Entorrhizomycetes
Glomeromycota
Glomeromycetes
Zygomycota
(paraphyletic)
Mucoromycotina
Mortierellomycetes
Mucoromycetes
Kickxellomycotina
Zoopagomycetes
Kickxellomycetes
Entomophthoromycotina
Neozygitomycetes
Basidiobolomycetes
Entomophthoromycetes
Zoosporic fungi
(paraphyletic)
Olpidiomycota
Olpidiomycetes
Blastocladiomycota
Blastocladiomycetes
Chytridiomycota
Neocallimastigomycetes
Hyaloraphidiomycetes
Monoblepharidomycetes
Chytridiomycetes
Fungal phyla are underlined. See also: fungi imperfecti (polyphyletic group).
[show]
v
t
e
Organisms and comparable organic structures
Life
Bacteria
Archaea
Eukaryota (Animalia
Fungi
Plantae
Protista)
Non-cellular
agents
Virus
dsDNA virus
ssDNA virus
dsRNA virus
(+)ssRNA virus
(−)ssRNA virus
ssRNA-RT virus
dsDNA-RT virus
Viroid
Pospiviroidae
Avsunviroidae
Prion
Mammalian prion
Fungal prion
Virus
dependent
Satellite
ssRNA satellite virus
dsDNA satellite virus (Virophage)
ssDNA satellite
dsRNA satellite
ssRNA satellite (Virusoid)
Defective
interfering particle
Defective Interfering RNA
Defective interfering DNA
Taxon identifiers
Wd: Q19088
EoL: 2908256
NCBI: 2759
Authority control
GND: 4070991-7
Retrieved from "https://en.wikipedia.org/w/index.php?title=Eukaryote&oldid=810182707"
Categories:
Eukaryotes
Domains (biology)
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This page was last edited on 13 November 2017, at 20:13.
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Konstantin Mereschkowsky

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Edouard Chatton

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Nucleus (cell)

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Vesicle (biology)

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Aerobic respiration

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Lateral gene transfer

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interlanguage-link-target

values:
https://gv.wikipedia.org/wiki/Eukaryota
Eukaryota – Manx
gv
gv
interlanguage-link-target

values:
https://gl.wikipedia.org/wiki/Eukaryota
Eukaryota – Galician
gl
gl
interlanguage-link-target

values: Կորիզավորներ – Armenian

values: सुकेन्द्रिक – Hindi

values:
https://hr.wikipedia.org/wiki/Eukarioti
Eukarioti – Croatian
hr
hr
interlanguage-link-target

values:
https://io.wikipedia.org/wiki/Eukarioto
Eukarioto – Ido
io
io
interlanguage-link-target

values:
https://id.wikipedia.org/wiki/Eukariota
Eukariota – Indonesian
id
id
interlanguage-link-target

values:
https://ia.wikipedia.org/wiki/Eucaryote
Eucaryote – Interlingua
ia
ia
interlanguage-link-target

values:
https://it.wikipedia.org/wiki/Eukaryota
Eukaryota – Italian
it
it
interlanguage-link-target

values: איקריוטים – Hebrew

values:
https://jv.wikipedia.org/wiki/Eukariota
Eukariota – Javanese
jv
jv
interlanguage-link-target

values: ევკარიოტები – Georgian

values: Эукариоттар – Kazakh

values:
https://sw.wikipedia.org/wiki/Eukaryota
Eukaryota – Swahili
sw
sw
interlanguage-link-target

values:
https://ku.wikipedia.org/wiki/Eukariyot
Eukariyot – Kurdish
ku
ku
interlanguage-link-target

values: Эукариоттор – Kyrgyz

values:
https://la.wikipedia.org/wiki/Eukaryota
Eukaryota – Latin
la
la
interlanguage-link-target

values:
https://lv.wikipedia.org/wiki/Eikarioti
Eikarioti – Latvian
lv
lv
interlanguage-link-target

values:
https://lij.wikipedia.org/wiki/Eukarya
Eukarya – Ligurian
lij
lij
interlanguage-link-target

values:
https://li.wikipedia.org/wiki/Eukarya
Eukarya – Limburgish
li
li
interlanguage-link-target

values: Еукариоти – Macedonian

values: യൂക്കാരിയോട്ടുകൾ – Malayalam

values: सकेंद्रक सजीव – Marathi

values: ევკარიოტეფი – Mingrelian

values:
https://ms.wikipedia.org/wiki/Eukariot
Eukariot – Malay
ms
ms
interlanguage-link-target

values: Эукариот – Mongolian

values:
https://oc.wikipedia.org/wiki/Eukaryota
Eukaryota – Occitan
oc
oc
interlanguage-link-target

values: ਸੁਕੇਂਦਰੀ ਪ੍ਰਾਣੀ – Punjabi

values:
https://pt.wikipedia.org/wiki/Eukaryota
Eukaryota – Portuguese
pt
pt
interlanguage-link-target

values:
https://ro.wikipedia.org/wiki/Eucariote
Eucariote – Romanian
ro
ro
interlanguage-link-target

values: Эукариоты – Russian

values: Эукариот – Sakha

values:
https://sq.wikipedia.org/wiki/Eukarioti
Eukarioti – Albanian
sq
sq
interlanguage-link-target

values:
https://sk.wikipedia.org/wiki/Eukaryoty
Eukaryoty – Slovak
sk
sk
interlanguage-link-target

values: interlanguage-link interwiki-sk badge-Q17437796 badge-featuredarticle

values: Еукариоте – Serbian

values:
https://sh.wikipedia.org/wiki/Eukariote
Eukariote – Serbo-Croatian
sh
sh
interlanguage-link-target

values:
https://su.wikipedia.org/wiki/Eukariot
Eukariot – Sundanese
su
su
interlanguage-link-target

values:
https://tl.wikipedia.org/wiki/Eukaryote
Eukaryote – Tagalog
tl
tl
interlanguage-link-target

values: மெய்க்கருவுயிரி – Tamil

values: Эукариотлар – Tatar

values: ยูแคริโอต – Thai

values: Эукариот – Tajik

values: حقیقی المرکزیہ – Urdu

values:
https://wa.wikipedia.org/wiki/Eucariote
Eucariote – Walloon
wa
wa
interlanguage-link-target

values: אייקאריאט – Yiddish

values: Ẹ̀ùkáríọ́tì – Yoruba

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