Eukaryotes () are
organisms whose
cells
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
have a
nucleus
Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to:
* Atomic nucleus, the very dense central region of an atom
*Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA
Nucl ...
.
All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the
three domains of life.
Bacteria and
Archaea (both
prokaryotes) make up the other two domains.
The eukaryotes are usually now regarded as having emerged in the Archaea or as a sister of the
Asgard
In Nordic mythology, Asgard (Old Norse: ''Ásgarðr'' ; "enclosure of the Æsir") is a location associated with the gods. It appears in a multitude of Old Norse sagas and mythological texts. It is described as the fortified home of the Æsir ...
archaea.
This implies that there are only
two domains of life, Bacteria and Archaea, with eukaryotes incorporated among archaea.
Eukaryotes represent a small minority of the number of organisms,
but, due to their generally much larger size, their collective global
biomass is estimated to be about equal to that of prokaryotes.
Eukaryotes emerged approximately 2.3–1.8 billion years ago, during the
Proterozoic eon, likely as
flagellated
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have fro ...
phagotrophs.
Their name comes from the
Greek εὖ (''eu'', "well" or "good") and
κάρυον (''karyon'', "nut" or "kernel").
Eukaryotic cells typically contain other
membrane-bound organelles such as
mitochondria and
Golgi apparatus.
Chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s can be found in
plants and
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
. Prokaryotic cells may contain
primitive organelles. Eukaryotes may be either
unicellular or
multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni ...
, and include many
cell types forming different kinds of
tissue. In comparison, prokaryotes are typically unicellular.
Animal
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s,
plants, and
fungi are the most familiar eukaryotes. Other eukaryotes are sometimes called
protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exclu ...
s.
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
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there a ...
to produce four
haploid daughter cells that act as sex cells or 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.
Cell features
Eukaryotic cells are typically much larger than those of
prokaryotes, having a volume of around 10,000 times greater than the prokaryotic cell. They have a variety of internal membrane-bound structures, called
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s, and a
cytoskeleton composed of
microtubules,
microfilament
Microfilaments, also called actin filaments, are protein filaments in the cytoplasm of eukaryotic cells that form part of the cytoskeleton. They are primarily composed of polymers of actin, but are modified by and interact with numerous other p ...
s, and
intermediate filament
Intermediate filaments (IFs) are cytoskeletal structural components found in the cells of vertebrates, and many invertebrates. Homologues of the IF protein have been noted in an invertebrate, the cephalochordate ''Branchiostoma''.
Intermedia ...
s, 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 membranes
Eukaryote cells include a variety of membrane-bound structures, collectively referred to as the
endomembrane system. Simple compartments, called
vesicles
Vesicle may refer to:
; In cellular biology or chemistry
* Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane
* Synaptic vesicle
; In human embryology
* Vesicle (embryology), bulge-like features o ...
and
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s, can form by budding off other membranes. Many cells ingest food and other materials through a process of
endocytosis
Endocytosis is a cellular process in which substances are brought into the cell. The material to be internalized is surrounded by an area of cell membrane, which then buds off inside the cell to form a vesicle containing the ingested material. E ...
, where the outer membrane
invaginates and then pinches off to form a vesicle. 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 known as the
nuclear envelope, with
nuclear pore
A nuclear pore is a part of a large complex of proteins, known as a nuclear pore complex that spans the nuclear envelope, which is the double membrane surrounding the eukaryotic cell nucleus. There are approximately 1,000 nuclear pore complexes ...
s that allow material to move in and out.
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. In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles (
cisterna
A cisterna (plural cisternae) is a flattened membrane vesicle found in the endoplasmic reticulum and Golgi apparatus. Cisternae are an integral part of the packaging and modification processes of proteins occurring in the Golgi.
Function
Protei ...
e), the
Golgi apparatus.
Vesicles may be specialized for various purposes. For instance,
lysosomes contain digestive
enzymes that break down most
biomolecule
A biomolecule or biological molecule is a loosely used term for molecules present in organisms that are essential to one or more typically biological processes, such as cell division, morphogenesis, or development. Biomolecules include large ...
s in the cytoplasm.
Peroxisome
A peroxisome () is a membrane-bound organelle, a type of microbody, found in the cytoplasm of virtually all eukaryotic cells. Peroxisomes are oxidative organelles. Frequently, molecular oxygen serves as a co-substrate, from which hydrogen pero ...
s are used to break down
peroxide
In chemistry, peroxides are a group of compounds with the structure , where R = any element. The group in a peroxide is called the peroxide group or peroxo group. The nomenclature is somewhat variable.
The most common peroxide is hydrogen p ...
, which is otherwise toxic. Many
protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
ns have contractile vacuoles, which collect and expel excess water, and
extrusome Extrusomes are membrane-bound structures in some eukaryotes which, under certain conditions, discharge their contents outside the cell. There are a variety of different types, probably not homologous, and serving various functions.
Notable extru ...
s, 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
Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane.
It is also defined as the measure of the tendency of a solution to take in a pure ...
.
Mitochondria
Mitochondria are organelles found in all but one eukaryote, and are commonly referred to as "the powerhouse of the cell". Mitochondria provide energy to the eukaryote cell by oxidising sugars or fats and releasing energy as
ATP. They have two surrounding
membranes
A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. Bi ...
, each a
phospholipid bi-layer; the
inner of which is folded into invaginations called
cristae
A crista (; plural cristae) is a fold in the inner membrane of a mitochondrion. The name is from the Latin for ''crest'' or ''plume'', and it gives the inner membrane its characteristic wrinkled shape, providing a large amount of surface area fo ...
where
aerobic respiration
Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
takes place.
The outer mitochondrial membrane is freely permeable and allows almost anything to enter into the
intermembrane space while the inner mitochondrial membrane is semi permeable so allows only some required things into the mitochondrial matrix.
Mitochondria contain
their own DNA, which has close structural similarities to bacterial DNA, and which encodes
rRNA
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
and
tRNA
Transfer RNA (abbreviated tRNA and formerly referred to as sRNA, for soluble RNA) is an adaptor molecule composed of RNA, typically 76 to 90 nucleotides in length (in eukaryotes), that serves as the physical link between the mRNA and the amino a ...
genes that produce RNA which is closer in structure to bacterial RNA than to eukaryote RNA.
They are now generally held to have developed from
endosymbiotic
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
prokaryotes, probably
Alphaproteobacteria
Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota (formerly Proteobacteria). The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse an ...
.
Some eukaryotes, such as the
metamonad
The metamonads are microscopic eukaryotic organisms, a large group of flagellate amitochondriate Loukozoa. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as ...
s such as ''
Giardia'' and ''
Trichomonas'', and the amoebozoan ''
Pelomyxa
''Pelomyxa'' is a genus of giant flagellar amoebae, usually 500-800 μm but occasionally up to 5 mm in length, found in anaerobic or microaerobic bottom sediments of stagnant freshwater ponds or slow-moving streams.Chistyakova, L. V., and A ...
'', appear to lack mitochondria, but all have been found to contain mitochondrion-derived organelles, such as
hydrogenosome
A hydrogenosome is a membrane-enclosed organelle found in some anaerobic ciliates, flagellates, and fungi. Hydrogenosomes are highly variable organelles that have presumably evolved from protomitochondria to produce molecular hydrogen and ATP i ...
s and
mitosome
A mitosome is an organelle found in some unicellular eukaryotic organisms, like in members of the supergroup Excavata. The mitosome was found and named in 1999, and its function has not yet been well characterized. It was termed a ''crypton'' by o ...
s, and thus have lost their 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
sulfur mobilisation system which provides the
clusters of iron and sulfur required for protein synthesis. The normal mitochondrial iron-sulfur cluster pathway has been lost secondarily.
Plastids
Plants and various groups of
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
also have
plastids. Plastids also have
their own DNA and are developed from
endosymbionts, in this case
cyanobacteria. They usually take the form of
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s which, like cyanobacteria, contain
chlorophyll
Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
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
Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory,) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibly ...
or ingestion. The capture and sequestering of photosynthetic cells and chloroplasts occurs in many types of modern eukaryotic organisms and is known as
kleptoplasty
Kleptoplasty or kleptoplastidy is a symbiotic phenomenon whereby plastids, notably chloroplasts from algae, are sequestered by host organisms. The word is derived from ''Kleptes'' (κλέπτης) which is Greek for thief. The alga is eaten normal ...
.
Endosymbiotic origins have also been proposed for the nucleus, and for eukaryotic
flagella.
Cytoskeletal structures
Many eukaryotes have long slender motile cytoplasmic projections, called
flagella, or similar structures called
cilia. Flagella and cilia are sometimes referred to as
undulipodia
An undulipodium or undulopodium (a Greek word meaning "swinging foot"), or a 9+2 organelle is a motile filamentous extracellular projection of eukaryotic cells. It is basically synonymous to flagella and cilia which are differing terms for simila ...
, and are variously involved in movement, feeding, and sensation. They are composed mainly of
tubulin
Tubulin in molecular biology can refer either to the tubulin protein superfamily of globular proteins, or one of the member proteins of that superfamily. α- and β-tubulins polymerize into microtubules, a major component of the eukaryotic cytoske ...
. These are entirely distinct from prokaryotic flagellae. They are supported by a bundle of microtubules arising from a
centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and are ...
, characteristically arranged as nine doublets surrounding two singlets. Flagella also may have hairs, or
mastigonemes
Mastigonemes are lateral "hairs" that attach to protistan flagella. Flimsy hairs attach to the flagella of euglenid flagellates, while stiff hairs occur in stramenopile and cryptophyte protists.Hoek, C. van den, Mann, D. G. and Jahns, H. M. (1 ...
, and scales connecting membranes and internal rods. Their interior is continuous with the cell's
cytoplasm
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. Th ...
.
Microfilamental structures composed of
actin
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...
and actin binding proteins, e.g., α-
actinin
Actinin is a microfilament protein. Alpha-actinin-1 is necessary for the attachment of actin myofilaments to the Z-lines in skeletal muscle cells, and to the dense bodies in smooth muscle cells. The functional protein is an anti-parallel dimer ...
,
fimbrin
Fimbrin also known as is plastin 1 is a protein that in humans is encoded by the PLS1 gene. Fimbrin is an actin cross-linking protein important in the formation of filopodia.
Structure
Fimbrin belongs to the calponin homology (CH) domain supe ...
,
filamin Filamins are a class of proteins that hold two actin filaments at large angles. Filamin protein in mammals is made up of an actin-binding domain at its N-terminus that is followed by 24 immunoglobulin-like repeat modules of roughly 95 amino acids. T ...
are present in submembranous cortical layers and bundles, as well.
Motor proteins of microtubules, e.g.,
dynein
Dyneins are a family of cytoskeletal motor proteins that move along microtubules in cells. They convert the chemical energy stored in ATP to mechanical work. Dynein transports various cellular cargos, provides forces and displacements important ...
or
kinesin and actin, e.g.,
myosins
Myosins () are a superfamily of motor proteins best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are ATP-dependent and responsible for actin-based motility.
The first myosin (M ...
provide dynamic character of the network.
Centrioles
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and a ...
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.
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
Chemotaxis (from '' chemo-'' + ''taxis'') is the movement of an organism or entity in response to a chemical stimulus. Somatic cells, bacteria, and other single-cell or multicellular organisms direct their movements according to certain chemi ...
and
chemokinesis Chemokinesis is chemically prompted kinesis, a motile response of unicellular prokaryotic or eukaryotic organisms to chemicals that cause the cell to make some kind of change in their migratory/swimming behaviour. Changes involve an increase or de ...
. Some
protists have various other microtubule-supported organelles. These include the
radiolaria and
heliozoa
Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes ( protists) with stiff arms (axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections fr ...
, which produce
axopodia
A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filament ...
used in flotation or to capture prey, and the
haptophyte
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
s, which have a peculiar flagellum-like organelle called the
haptonema
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
.
Cell wall
The cells of plants and algae, 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.
The major
polysaccharides making up the primary cell wall of
land plants
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
are
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
,
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
Xyloglucan is a hemicellulose that occurs in the primary cell wall of all vascular plants; however, all enzymes responsible for xyloglucan metabolism are found in Charophyceae algae.LEV Del Bem and M Vincentz (2010) Evolution of xyloglucan-related ...
.
Differences among eukaryotic cells
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
All animals are eukaryotic. Animal cells are distinct from those of other eukaryotes, most notably
plants
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
, as they lack
cell wall
A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
s and
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s and have smaller
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s. Due to the lack of a
cell wall
A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
, animal cells can transform into a variety of shapes. A
phagocytic cell can even engulf other structures.
Plant cell
Plant cells have a number of features that distinguish them from the cells of the other eukaryotic organisms. These include:
* A large
central vacuole (enclosed by a membrane, the
tonoplast), which maintains the cell's
turgor and controls movement of
molecules between the
cytosol
The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells (intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondrio ...
and
sap
Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant.
Sap is distinct from latex, resin, or cell sap; it is a separ ...
* A primary
cell wall
A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
containing
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
,
hemicellulose and
pectin, deposited by the
protoplast
Protoplast (), is a biological term coined by Hanstein in 1880 to refer to the entire cell, excluding the cell wall. Protoplasts can be generated by stripping the cell wall from plant, bacterial, or fungal cells by mechanical, chemical or enzy ...
on the outside of the
cell membrane; this contrasts with the cell walls of
fungi, which contain
chitin, and the
cell envelope
The cell envelope comprises the inner cell membrane and the cell wall of a bacterium. In gram-negative bacteria an outer membrane is also included. This envelope is not present in the Mollicutes where the cell wall is absent.
Bacterial cell enve ...
s of prokaryotes, in which
peptidoglycans are the main structural molecules
* The
plasmodesmata, pores in the cell wall that link adjacent cells and allow plant cells to communicate with adjacent cells.
Animals have a different but functionally analogous system of
gap junctions between adjacent cells.
*
Plastids, especially
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s,
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s that contain
chlorophyll
Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
, the pigment that gives
plants their green color and allows them to perform
photosynthesis
*
Bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
s and
seedless vascular plants only have flagellae and centrioles in the sperm cells.
Sperm of
cycad
Cycads are seed plants that typically have a stout and woody ( ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or ...
s and ''
Ginkgo'' are large, complex cells that swim with hundreds to thousands of flagellae.
*
Conifers
Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All exta ...
(Pinophyta) and
flowering plants (Angiospermae) lack the
flagellae and
centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and are ...
s that are present in animal cells.
Fungal cell
The cells of
fungi are similar to animal cells, with the following exceptions:
* A cell wall that contains
chitin
* Less compartmentation between cells; the
hypha
A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium.
Structure
A hypha consists of one or ...
e of higher fungi have porous partitions called
septa, which allow the passage of cytoplasm, organelles, and, sometimes, nuclei; so each organism is essentially a giant
multinucleate supercell – these fungi are described as
coenocytic
A coenocyte () is a multinucleate cell which can result from multiple nuclear divisions without their accompanying cytokinesis, in contrast to a syncytium, which results from cellular aggregation followed by dissolution of the cell membranes insi ...
. Primitive fungi have few or no septa.
* Only the most primitive fungi,
chytrid
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytrid ...
s, have flagella.
Other eukaryotic cells
Some groups of eukaryotes have unique organelles, such as the cyanelles (unusual plastids) of the
glaucophytes,
the haptonema of the
haptophyte
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for ''Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at t ...
s, or the
ejectosome An ejectosome is a cellular organelle responsible for ejecting their contents from the cell. Two unrelated types of ejectosomes are described in the literature:
# Cryptomonads have two types of characteristic ejectosomes known as extrusomes.
# Int ...
s of the
cryptomonad
The cryptomonads (or cryptophytes) are a group of algae, most of which have plastids. They are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, with an anterio ...
s. Other structures, such as
pseudopodia, are found in various eukaryote groups in different forms, such as the lobose
amoebozoa
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported classi ...
ns or the reticulose
foraminiferan
Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
s. Structures known as
cortical alveoli The cortical alveoli (singular: cortical alveolum) are cellular organelles composed of vesicles located under the cytoplasmic membrane, to which they give support. They have been defined as membrane sacs that strengthen the cellular cortex through ...
are vesicles present under the cell membrane of many
protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exclu ...
s such as
dinoflagellates,
ciliate
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...
s and
apicomplexan
The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. The ...
parasites.
Reproduction
Cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there a ...
generally takes place
asexually by
mitosis, a process that allows each daughter nucleus to receive one copy of each
chromosome. Most eukaryotes also have a life cycle that involves
sexual reproduction,
alternating between a
haploid phase, where only one copy of each chromosome is present in each cell and a
diploid phase, wherein two copies of each chromosome are present in each cell. The diploid phase is formed by fusion of two haploid gametes to form a zygote, which may divide by mitosis or undergo chromosome reduction by
meiosis. There is considerable variation in this pattern. Animals have no multicellular haploid phase, but each plant generation can consist of haploid and diploid multicellular phases.
Eukaryotes have a smaller surface area to volume ratio than prokaryotes, and thus have lower metabolic rates and longer generation times.
The
evolution of sexual reproduction may be a primordial and fundamental characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and
Roger
Roger is a given name, usually masculine, and a surname. The given name is derived from the Old French personal names ' and '. These names are of Germanic origin, derived from the elements ', ''χrōþi'' ("fame", "renown", "honour") and ', ' ...
proposed that facultative sex was present in the common ancestor of all eukaryotes. A core set of genes that function in meiosis is present in both ''
Trichomonas vaginalis'' and ''
Giardia intestinalis
''Giardia duodenalis'', also known as ''Giardia intestinalis'' and ''Giardia lamblia'', is a flagellated parasitic microorganism of the genus ''Giardia'' that colonizes the small intestine, causing a diarrheal condition known as giardiasis. The ...
'', two organisms previously thought to be asexual.
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.
Eukaryotic species once thought to be asexual, such as parasitic protozoa of the genus ''
Leishmania'', have been shown to have a sexual cycle. 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.
Classification
In
antiquity
Antiquity or Antiquities may refer to:
Historical objects or periods Artifacts
*Antiquities, objects or artifacts surviving from ancient cultures
Eras
Any period before the European Middle Ages (5th to 15th centuries) but still within the histo ...
, the two lineages of
animal
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s and
plants were recognized. They were given the
taxonomic rank of
Kingdom
Kingdom commonly refers to:
* A monarchy ruled by a king or queen
* Kingdom (biology), a category in biological taxonomy
Kingdom may also refer to:
Arts and media Television
* ''Kingdom'' (British TV series), a 2007 British television drama s ...
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.
The various single-cell eukaryotes were originally placed with plants or animals when they became known. In 1818, the German biologist
Georg A. Goldfuss coined the word ''
protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
'' to refer to organisms such as
ciliate
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a differen ...
s, and this group was expanded until it encompassed all single-celled eukaryotes, and given their own kingdom, the
Protista
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
, by
Ernst Haeckel in 1866.
The eukaryotes thus came to be composed of four kingdoms:
* Kingdom
Protista
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
* Kingdom
Plantae
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
* Kingdom
Fungi
* Kingdom
Animalia
The protists were understood to be "primitive forms", and thus an
evolutionary grade, united by their primitive unicellular nature.
[ 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
Carl Richard Woese (; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal RNA, ...
, uniting all the eukaryote kingdoms under the eukaryote domain. 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.
Eukaryotes are a clade usually assessed to be sister to Heimdallarchaeota
Asgard or Asgardarchaeota is a proposed superphylum consisting of a group of archaea that includes Lokiarchaeota, Thorarchaeota, Odinarchaeota, and Heimdallarchaeota. It appears the eukaryotes emerged within the Asgard, in a branch containing ...
in the Asgard
In Nordic mythology, Asgard (Old Norse: ''Ásgarðr'' ; "enclosure of the Æsir") is a location associated with the gods. It appears in a multitude of Old Norse sagas and mythological texts. It is described as the fortified home of the Æsir ...
grouping in the Archaea. In one proposed system, the basal groupings are the Opimoda, Diphoda, the Discoba, and the Loukozoa
Loukozoa (+ Ancyromonads) (From Greek ''loukos'': groove) is a proposed taxon used in some classifications of eukaryotes, consisting of the Metamonada and Malawimonadea. Ancyromonads are closely related to this group, as sister of the entire gr ...
. The Eukaryote root is usually assessed to be near or even in Discoba.
A classification produced in 2005 for the International Society of Protistologists, 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
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes.
It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in ...
, and a review in 2006 noted the lack of evidence for several of the supposed six supergroups. A revised classification in 2012 recognizes five supergroups.
There are also smaller groups of eukaryotes whose position is uncertain or seems to fall outside the major groups – in particular, Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
, Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
, Centrohelida
The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth.
Characteristics
Individuals are unicellular and sphe ...
, Telonemia, Picozoa,[ ]Apusomonadida
The Apusomonadida are a group of protozoan zooflagellates that glide on surfaces, and mostly consume prokaryotes. They are of particular evolutionary interest because they appear to be the sister group to the Opisthokonts, the clade that includ ...
, Ancyromonadida
Ancyromonadida or Planomonadida is a small group of biflagellated protists found in the soil and in aquatic habitats, where they feed on bacteria.Cavalier-Smith, T. (2013)Early evolution of eukaryote feeding modes, cell structural diversity, and ...
, Breviatea
''Breviata anathema'' is a single-celled flagellate amoeboid eukaryote, previously studied under the name '' Mastigamoeba invertens''. The cell lacks mitochondria but has remnant mitochondrial genes, and possesses an organelle believed to be a modi ...
, and the genus ''Collodictyon
''Collodictyon'' is a genus of single-celled, omnivorous eukaryotes belonging to the collodictyonids, also known as diphylleids. Due to their mix of cellular components, Collodictyonids do not belong to any well-known kingdom-level grouping of th ...
''. 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
Roger is a given name, usually masculine, and a surname. The given name is derived from the Old French personal names ' and '. These names are of Germanic origin, derived from the elements ', ''χrōþi'' ("fame", "renown", "honour") and ', ' ...
& Simpson said in 2009 "with the current pace of change in our understanding of the eukaryote tree of life, we should proceed with caution." Newly identified protists, purported to represent novel, deep-branching lineages, continue to be described well into the 21st century; recent examples including ''Rhodelphis
''Rhodelphis'' is a single-celled archaeplastid that lives in aquatic environments and is the sister group to red algae and possibly Picozoa. While red algae have no flagellated stages and are generally photoautotrophic, ''Rhodelphis'' is a flage ...
'', putative sister group to Rhodophyta, and ''Anaeramoeba
Anaeramoeba is a genus of anaerobic protists on uncertain phylogenetic position, first described in 2016.
Description
As the name implies, ''Anaeramoeba'' are anaerobic amoeboid organisms which form a fan-like shape similar to that of '' Fl ...
'', anaerobic amoebaflagellates of uncertain placement.
Phylogeny
The rRNA
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
trees constructed during the 1980s and 1990s left most eukaryotes in an unresolved "crown" group (not technically a true crown
A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
), 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
In phylogenetics, long branch attraction (LBA) is a form of systematic error whereby distantly related lineages are incorrectly inferred to be closely related. LBA arises when the amount of molecular or morphological change accumulated within a lin ...
, and they are known to have lost them secondarily.
It has been estimated that there may be 75 distinct lineages of eukaryotes. Most of these lineages are protists.
The known eukaryote genome sizes vary from 8.2 megabases (Mb) in ''Babesia bovis
''Babesia bovis'' is an Apicomplexan single-celled parasite of cattle which occasionally infects humans. The disease it and other members of the genus ''Babesia'' cause is a hemolytic anemia known as babesiosis and colloquially called Texas cattl ...
'' to 112,000–220,050 Mb in the dinoflagellate ''Prorocentrum micans
The Prorocentrales are a small order of dinoflagellates. They are distinguished by having their two flagella inserted apically, rather than ventrally as in other groups. One flagellum extends forward and the other circles its base, and there are ...
'', showing that the genome of the ancestral eukaryote has undergone considerable variation during its evolution. The last common ancestor of all eukaryotes is believed to have been a phagotrophic protist with a nucleus, at least one centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and are ...
and cilium
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projec ...
, facultatively aerobic mitochondria, sex ( meiosis and syngamy
Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proc ...
), a dormant cyst
A cyst is a closed sac, having a distinct envelope and division compared with the nearby tissue. Hence, it is a cluster of cells that have grouped together to form a sac (like the manner in which water molecules group together to form a bubble) ...
with a cell wall of chitin and/or cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
and peroxisome
A peroxisome () is a membrane-bound organelle, a type of microbody, found in the cytoplasm of virtually all eukaryotic cells. Peroxisomes are oxidative organelles. Frequently, molecular oxygen serves as a co-substrate, from which hydrogen pero ...
s. Later endosymbiosis led to the spread of plastids in some lineages.
Although there is still considerable uncertainty in global eukaryote phylogeny, particularly regarding the position of the root, a rough consensus has started to emerge from the phylogenomic studies of the past two decades. The majority of eukaryotes can be placed in one of two large clades dubbed Amorphea
Amorphea are members of a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the mem ...
(similar in composition to the unikont
Amorphea are members of a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the mem ...
hypothesis) and the Diaphoretickes, which includes plants and most algal lineages. A third major grouping, the Excavata
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...
, has been abandoned as a formal group in the most recent classification of the International Society of Protistologists due to growing uncertainty as to whether its constituent groups belong together. The proposed phylogeny below includes only one group of excavates ( Discoba), and incorporates the recent proposal that picozoans are close relatives of rhodophytes.
In some analyses, the Hacrobia group (Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
+ Cryptophyta
The cryptophyceae are a class of algae, most of which have plastids. About 220 species are known, and they are common in freshwater, and also occur in marine and brackish habitats. Each cell is around 10–50 μm in size and flattened in shape, ...
) is placed next to Archaeplastida
The Archaeplastida (or kingdom Plantae ''sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group ...
, but in others it is nested inside the Archaeplastida. However, several recent studies have concluded that Haptophyta and Cryptophyta do not form a monophyletic group. The former could be a sister group to the SAR group, the latter cluster with the Archaeplastida
The Archaeplastida (or kingdom Plantae ''sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group ...
(plants in the broad sense).
The division of the eukaryotes into two primary clades, bikonts
A bikont ("two flagella") is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled members of the group, and the presumed ancestor, have two flagella.
Enzymes
Another shared trait of bikonts is the fusion of two ge ...
(Archaeplastida
The Archaeplastida (or kingdom Plantae ''sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group ...
+ SAR + Excavata
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...
) and unikonts (Amoebozoa
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported classi ...
+ Opisthokonta), derived from an ancestral biflagellar organism and an ancestral uniflagellar organism, respectively, had been suggested earlier. A 2012 study produced a somewhat similar division, although noting that the terms "unikonts" and "bikonts" were not used in the original sense.
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. The main remaining controversies are the root, and the exact positioning of the Rhodophyta and the bikont
A bikont ("two flagella") is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled members of the group, and the presumed ancestor, have two flagella.
Enzymes
Another shared trait of bikonts is the fusion of two ge ...
s Rhizaria, Haptista, Cryptista, Picozoa and Telonemia, many of which may be endosymbiotic eukaryote-eukaryote hybrids. Archaeplastida acquired chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s probably by endosymbiosis of a prokaryotic ancestor related to a currently extant cyanobacterium
Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blue ...
, ''Gloeomargarita lithophora
''Gloeomargarita lithophora'' is a cyanobacterium, and is the proposed sister of the endosymbiotic plastids in the eukaryote group Archaeplastida (glaucophytes, plants, green and red algae). ''Gloeomargaritas relative would have ended up in an an ...
''.
Cavalier-Smith's tree
Thomas Cavalier-Smith 2010, 2013, 2014, 2017 and 2018 places the eukaryotic tree's root between Excavata
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...
(with ventral feeding groove supported by a microtubular root) and the grooveless Euglenozoa
Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by three major clades, i.e., Kinetoplastea, D ...
, and monophyletic Chromista, correlated to a single endosymbiotic event of capturing a red-algae. He et al. specifically supports rooting the eukaryotic tree between a monophyletic Discoba (Discicristata
Discicristata is a proposed eukaryotic clade. It consists of Euglenozoa plus Percolozoa.
It was proposed that Discicristata plus Cercozoa yielded Cabozoa. Another proposal is to group Discicristata with Jakobida into Discoba superphylum.
See ...
+ Jakobida) and an Amorphea
Amorphea are members of a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the mem ...
- Diaphoretickes clade.
Evolutionary history
Origin of eukaryotes
The origin of the eukaryotic cell, also known as eukaryogenesis, is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. A number of approaches have been used to find the first eukaryote and their closest relatives. The last eukaryotic common ancestor (LECA) is the hypothetical last common ancestor
In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of all living eukaryotes, and was most likely a biological population.
Eukaryotes have a number of features that differentiate them from prokaryotes, including an endomembrane system, and unique biochemical pathways such as sterane synthesis.[ A set of proteins called eukaryotic signature proteins (ESPs) was proposed to identify eukaryotic relatives in 2002: They have no homology to proteins known in other domains of life by then, but they appear to be universal among eukaryotes. They include proteins that make up the cytoskeleton, the complex transcription machinery, membrane-sorting systems, the nuclear pore, as well as some enzymes in the biochemical pathways.]
Fossils
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 acritarch
Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity refle ...
s are known from at least 1.65 billion years ago, and the possible alga ''Grypania
''Grypania'' is an early, tube-shaped fossil from the Proterozoic eon. The organism, with a size over one centimeter and consistent form, could have been a giant bacterium, a bacterial colony, or a eukaryotic alga. The oldest probable ''Grypania' ...
'' has been found as far back as 2.1 billion years ago. The '' Geosiphon''-like fossil fungus '' Diskagma'' has been found in paleosols 2.2 billion years old.
Organized living structures have been found in the black shale
Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...
s of the Palaeoproterozoic Francevillian B Formation in Gabon, dated at 2.1 billion years old. Eukaryotic life could have evolved at that time. Fossils that are clearly related to modern groups start appearing an estimated 1.2 billion years ago, in the form of a red algae, though recent work suggests the existence of fossilized filamentous algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micro ...
in the Vindhya basin dating back perhaps to 1.6 to 1.7 billion years ago.
The presence of eukaryotic-specific biomarkers
In biomedical contexts, a biomarker, or biological marker, is a measurable indicator of some biological state or condition. Biomarkers are often measured and evaluated using blood, urine, or soft tissues to examine normal biological processes, pa ...
( steranes) in Australian shales previously indicated that eukaryotes were present in these rocks dated at 2.7 billion years old, which was even 300 million years older than the first geological records of the appreciable amount of molecular oxygen during the Great Oxidation Event. However, these Archaean biomarkers were eventually rebutted as later contaminants. Currently, putatively the oldest biomarker records are only ~800 million years old. In contrast, a molecular clock analysis suggests the emergence of sterol biosynthesis as early as 2.3 billion years ago, and thus there is a huge gap between molecular data and geological data, which hinders a reasonable inference of the eukaryotic evolution through biomarker records before 800 million years ago. The nature of steranes as eukaryotic biomarkers is further complicated by the production of sterols by some bacteria.
Whenever their origins, eukaryotes may not have become ecologically dominant until much later; a massive uptick in the zinc composition of marine sediments has been attributed to the rise of substantial populations of eukaryotes, which preferentially consume and incorporate zinc relative to prokaryotes, approximately a billion years after their origin (at the latest).
In April 2019, biologists reported that the very large medusavirus
Medusavirus is a nucleocytoplasmic large DNA virus first isolated from a Japanese hot spring in 2019. It notably encodes all five types of histones — H1, H2A, H2B, H3, and H4 — which are involved in DNA packaging in eukaryotes, raising the p ...
, or a relative, may have been responsible, at least in part, for the evolutionary emergence of complex eukaryotic cells from simpler prokaryotic cells
A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
.[
* ]
Relationship to Archaea
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
Neomura is a possible clade composed of the two domains of life of Archaea and Eukaryota. The group was named by Thomas Cavalier-Smith in 2002. Its name means "new walls", reflecting his hypothesis that it evolved from Bacteria, and one of t ...
. 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 bacterium
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, and the nucleus from an archaeon, from a virus, or from a pre-cell A pre-cell (or protocell) is a hypothetical lipid-based structure that, under the RNA world hypothesis, could have confined RNA in ancient times. A pre-cell allowed the RNA to remain in close proximity with other RNA molecules, keeping them concentr ...
.
* Eukaryotes developed from Archaea, and acquired their bacterial characteristics through the endosymbiosis of a proto-mitochondrion
The proto-mitochondrion is the hypothetical ancestral bacterial endosymbiont from which all mitochondria in eukaryotes are thought to descend, after an episode of symbiogenesis which created the aerobic eukaryotes.
Phylogeny
The phylogenetic ana ...
of bacterial origin.
* Eukaryotes and Archaea developed separately from a modified bacterium.
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. This is mainly to account for the fact that eukaryotic signature proteins were not found anywhere else by 2002.
* 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, 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, with a modification of hypothesis 2 making the 4th column: The ''eocyte hypothesis'', in which the Archaea are paraphyletic. (The table and the names for the hypotheses are based on Harish & Kurland, 2017.][)
In recent years, most researchers have favoured either the three domains (3D) or the eocyte hypothesis. An ]rRNA
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
analysis supports the eocyte scenario, apparently with the Eukaryote root in Excavata
Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free- ...
.[ A cladogram supporting the eocyte hypothesis, positioning eukaryotes within Archaea, based on phylogenomic analyses of the ]Asgard
In Nordic mythology, Asgard (Old Norse: ''Ásgarðr'' ; "enclosure of the Æsir") is a location associated with the gods. It appears in a multitude of Old Norse sagas and mythological texts. It is described as the fortified home of the Æsir ...
archaea, is:
In this scenario, the Asgard group is seen as a sister taxon of the TACK
TACK is a group of archaea acronym for Thaumarchaeota (now Nitrososphaerota), Aigarchaeota, Crenarchaeota (now Thermoproteota), and Korarchaeota, the first groups discovered. They are found in different environments ranging from acidophilic ...
group, which comprises Thermoproteota (formerly named eocytes or Crenarchaeota), Nitrososphaerota
The Nitrososphaerota (syn. Thaumarchaeota) are a phylum of the Archaea proposed in 2008 after the genome of '' Cenarchaeum symbiosum'' was sequenced and found to differ significantly from other members of the hyperthermophilic phylum Thermoprot ...
(formerly Thaumarchaeota), and others. This group is reported contain many of the eukaryotic signature proteins and produce vesicles.
In 2017, there was 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). 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. 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
A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population due to environmental events such as famines, earthquakes, floods, fires, disease, and droughts; or human activities such as specicide, widespread violen ...
, rather than a simpler organism arising early in the history of life. On the other hand, the researchers who came up with Asgard re-affirmed their hypothesis with additional Asgard samples. Since then, the publication of additional Asgard archaeal genomes and the independent reconstruction of phylogenomic trees by multiple independent laboratories have provided additional support for an Asgard archaeal origin of eukaryotes.
Details of the relation of Asgard archaea members and eukaryotes are still under consideration, although, in January 2020, scientists reported that ''Candidatus Prometheoarchaeum syntrophicum
Lokiarchaeota is a proposed phylum of the Archaea. The phylum includes all members of the group previously named Deep Sea Archaeal Group (DSAG), also known as Marine Benthic Group B (MBG-B). Lokiarchaeota is part of the superphylum Asgard contai ...
'', a type of cultured Asgard archaea, may be a possible link between simple prokaryotic and complex eukaryotic microorganisms about two billion years ago.
Endomembrane system and mitochondria
The origins of the endomembrane system and mitochondria are also unclear. 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. 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 The hydrogen hypothesis is a model proposed by William F. Martin and Miklós Müller in 1998 that describes a possible way in which the mitochondrion arose as an endosymbiont within a prokaryotic host in the archaea, giving rise to a symbiotic asso ...
is one particular version).
In a study using genomes to construct supertree A supertree is a single phylogenetic tree assembled from a combination of smaller phylogenetic trees, which may have been assembled using different datasets (e.g. morphological and molecular) or a different selection of taxa. Supertree algorithms c ...
s, Pisani ''et al.'' (2007) suggest that, along with evidence that there was never a mitochondrion-less eukaryote, eukaryotes evolved from a syntrophy In biology, syntrophy, synthrophy, or cross-feeding (from Greek ''syn'' meaning together, ''trophe'' meaning nourishment) is the phenomenon of one species feeding on the metabolic products of another species to cope up with the energy limitations by ...
between an archaea closely related to Thermoplasmatales
In taxonomy, the Thermoplasmatales are an order of the Thermoplasmata. All are acidophiles, growing optimally at pH below 2. '' Picrophilus'' is currently the most acidophilic of all known organisms, being capable of growing at a pH of -0.06. ...
and an alphaproteobacterium
Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota (formerly Proteobacteria). The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse an ...
, likely a symbiosis
Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
driven by sulfur or hydrogen. The mitochondrion and its genome is a remnant of the alphaproteobacterial endosymbiont. The majority of the genes from the symbiont have been transferred to the nucleus. They make up most of the metabolic and energy-related pathways of the eukaryotic cell, while the information system (DNA polymerase, transcription, translation) is retained from archaea.
Hypotheses
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
=
Autogenous models propose that a proto-eukaryotic cell containing a nucleus
Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to:
* Atomic nucleus, the very dense central region of an atom
*Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA
Nucl ...
existed first, and later acquired mitochondria. According to this model, a large prokaryote developed invaginations
Invagination is the process of a surface folding in on itself to form a cavity, pouch or tube. In developmental biology, invagination is a mechanism that takes place during gastrulation. This mechanism or cell movement happens mostly in the vegetal ...
in its plasma membrane in order to obtain enough surface area to service its cytoplasmic
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
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.
Mitochondria are proposed to come from the endosymbiosis of an aerobic proteobacterium
Pseudomonadota (synonym Proteobacteria) is a major phylum of Gram-negative bacteria. The renaming of phyla in 2021 remains controversial among microbiologists, many of whom continue to use the earlier names of long standing in the literature. The ...
after an eukaryote with a nucleus has evolved. This theory is less held onto because it requires extra assumptions to explain current conditions. For example, as every known eukaryote has a mitochondrion (or at least show signs of having an ancestor that had), one must assumed that all the eukaryotic lineages that did not acquire mitochondria became extinct. The theory also doesn’t explain why anaerobic variants of mitochondria have evolved.
=Chimeric models
=
Chimeric models claim that two prokaryotic cells existed initially – an archaeon
Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaeba ...
and a bacterium
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
. The closest living relatives of these appears to be Asgardarchaeota and (distantly related) the alphaproteobacteria
Alphaproteobacteria is a class of bacteria in the phylum Pseudomonadota (formerly Proteobacteria). The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse an ...
called the proto-mitochondrion
The proto-mitochondrion is the hypothetical ancestral bacterial endosymbiont from which all mitochondria in eukaryotes are thought to descend, after an episode of symbiogenesis which created the aerobic eukaryotes.
Phylogeny
The phylogenetic ana ...
.[
* ] 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.
=The inside-out hypothesis
=
The inside-out hypothesis suggests that the fusion between free-living mitochondria-like bacteria, and an archaeon into a eukaryotic cell happened gradually over a long period of time, instead of in a single phagocytotic event. In this scenario, an archaeon would trap aerobic bacteria with cell protrusions, and then keep them alive to draw energy from them instead of digesting them. During the early stages the bacteria would still be partly in direct contact with the environment, and the archaeon would not have to provide them with all the required nutrients. But eventually the archaeon would engulf the bacteria completely, creating the internal membrane structures and nucleus membrane in the process.[
* ]
It is assumed the archaean group called halophile
The halophiles, named after the Greek word for "salt-loving", are extremophiles that thrive in high salt concentrations. While most halophiles are classified into the domain Archaea, there are also bacterial halophiles and some eukaryotic species, ...
s went through a similar procedure, where they acquired as much as a thousand genes from a bacterium, way more than through the conventional horizontal gene transfer that often occurs in the microbial world, but that the two microbes separated again before they had fused into a single eukaryote-like cell.
An expanded version of the inside-out hypothesis proposes that the eukaryotic cell was created by physical interactions between two prokaryotic organisms and that the last common ancestor of eukaryotes got its genome from a whole population or community of microbes participating in cooperative relationships to thrive and survive in their environment. The genome from the various types of microbes would complement each other, and occasional horizontal gene transfer between them would be largely to their own benefit. This accumulation of beneficial genes gave rise to the genome of the eukaryotic cell, which contained all the genes required for independence.
=The serial endosymbiotic hypothesis
=
According to serial endosymbiotic theory (championed by Lynn Margulis
Lynn Margulis (born Lynn Petra Alexander; March 5, 1938 – November 22, 2011) was an American evolutionary biologist, and was the primary modern proponent for the significance of symbiosis in evolution. Historian Jan Sapp has said that "Lynn Ma ...
), a union between a motile anaerobic
Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to:
* Anaerobic adhesive, a bonding a ...
bacterium (like ''Spirochaeta'') and a thermoacidophilic crenarchaeon
The Thermoproteota (also known as crenarchaea) are archaea that have been classified as a phylum of the Archaea domain. Initially, the Thermoproteota were thought to be sulfur-dependent extremophiles but recent studies have identified characteri ...
(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 alphaproteobacteria 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 In biology, syntrophy, synthrophy, or cross-feeding (from Greek ''syn'' meaning together, ''trophe'' meaning nourishment) is the phenomenon of one species feeding on the metabolic products of another species to cope up with the energy limitations by ...
of sulfur existing between the two types of bacteria – ''Desulfobacter
''Desulfobacter'' is a genus of bacteria from the family of Desulfobacteraceae. ''Desulfobacter'' has the ability to oxidize acetate
An acetate is a salt formed by the combination of acetic acid with a base (e.g. alkaline, earthy, metal ...
'' and ''Spirochaeta''.
However, such an association based on motile symbiosis has never been observed practically. Also there is no evidence of archaeans and spirochetes adapting to intense acid-based environments. In addition, the theory posits that mitochondrion-less eukaryotes have existed, tying back to the problem in the autogenous model.
=The hydrogen hypothesis
=
In the hydrogen hypothesis The hydrogen hypothesis is a model proposed by William F. Martin and Miklós Müller in 1998 that describes a possible way in which the mitochondrion arose as an endosymbiont within a prokaryotic host in the archaea, giving rise to a symbiotic asso ...
, the symbiotic linkage of an anaerobic and autotroph
An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Work ...
ic methanogenic archaeon (host) with an alphaproteobacterium (the symbiont) gave rise to the eukaryotes. The host used hydrogen (H2) and carbon dioxide () to produce methane while the symbiont, capable of aerobic respiration, expelled H2 and 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
A diatom ( Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising se ...
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.
=The syntrophy hypothesis
=
The syntrophy In biology, syntrophy, synthrophy, or cross-feeding (from Greek ''syn'' meaning together, ''trophe'' meaning nourishment) is the phenomenon of one species feeding on the metabolic products of another species to cope up with the energy limitations by ...
hypothesis was developed in contrast to the hydrogen hypothesis and proposes the existence of two symbiotic events. According to this model, the origin of eukaryotic cells was based on metabolic symbiosis (syntrophy) between a methanogenic archaeon and a deltaproteobacterium
The Myxococcota are a phylum of bacteria known as the fruiting gliding bacteria. All species of this group are Gram-negative. They are predominantly aerobic genera that release myxospores in unfavorable environments.
Phylogeny
The currently ac ...
. This syntrophic symbiosis was initially facilitated by H2 transfer between different species under anaerobic environments. In earlier stages, an alphaproteobacterium became a member of this integration, and later developed into the mitochondrion. Gene transfer from a deltaproteobacterium to an archaeon led to the methanogenic archaeon developing into a nucleus. The archaeon constituted the genetic apparatus, while the deltaproteobacterium contributed towards the cytoplasm
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. Th ...
ic features.
This theory incorporates two selective forces at the time of nucleus evolution
* presence of metabolic partitioning to avoid the harmful effects of the co-existence of anabolic
Anabolism () is the set of metabolic pathways that construct molecules from smaller units. These reactions require energy, known also as an endergonic process. Anabolism is the building-up aspect of metabolism, whereas catabolism is the breakin ...
and catabolic
Catabolism () is the set of metabolic pathways that breaks down molecules into smaller units that are either oxidized to release energy or used in other anabolic reactions. Catabolism breaks down large molecules (such as polysaccharides, lipids, ...
cellular pathways, and
* prevention of abnormal protein biosynthesis
Protein biosynthesis (or protein synthesis) is a core biological process, occurring inside cells, balancing the loss of cellular proteins (via degradation or export) through the production of new proteins. Proteins perform a number of critical ...
due to a vast spread of introns in the archaeal genes after acquiring the mitochondrion and losing methanogenesis
Methanogenesis or biomethanation is the formation of methane coupled to energy conservation by microbes known as methanogens. Organisms capable of producing methane for energy conservation have been identified only from the domain Archaea, a group ...
.
=6+ serial endosymbiosis scenario
=
A complex scenario of 6+ serial endosymbiotic events of archaea and bacteria has been proposed in which mitochondria and an asgard related archaeota were acquired at a late stage of eukaryogenesis, possibly in combination, as a secondary endosymbiont. The findings have been rebuked as an artifact.
See also
* Eukaryote hybrid genome
Eukaryote hybrid genomes result from interspecific hybridization, where closely related species mate and produce offspring with admixed genomes. The advent of large-scale genomic sequencing has shown that hybridization is common, and that it ma ...
* Evolution of sexual reproduction
* List of sequenced eukaryotic genomes
This list of "sequenced" eukaryotic genomes contains all the eukaryotes known to have publicly available complete nuclear and organelle genome sequences that have been sequenced, assembled, annotated and published; draft genomes are not included, ...
* ''Parakaryon myojinensis
''Parakaryon myojinensis'', also known as the Myojin parakaryote, is a highly unusual species of single-celled organism known only from a single specimen, described in 2012. It has features of both prokaryotes and eukaryotes but is apparently d ...
''
* Prokaryote
* Nitrososphaerota
The Nitrososphaerota (syn. Thaumarchaeota) are a phylum of the Archaea proposed in 2008 after the genome of '' Cenarchaeum symbiosum'' was sequenced and found to differ significantly from other members of the hyperthermophilic phylum Thermoprot ...
* Vault (organelle)
The vault or vault cytoplasmic ribonucleoprotein is a eukaryotic organelle whose function is not yet fully understood. Discovered and isolated by Nancy Kedersha and Leonard Rome in 1986, vaults are cytoplasmic organelles which, when negative- ...
Notes
References
External links
"Eukaryotes"
(Tree of Life Web Project
The Tree of Life Web Project is an Internet project providing information about the diversity and phylogeny of life on Earth.
This collaborative peer reviewed project began in 1995, and is written by biologists from around the world. The site h ...
)
*
Attraction and sex among our microbial Last Eukaryotic Common Ancestors
The Atlantic, November 11, 2020
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Eukaryote