A eukaryote (/jukri.

ot/ or /jukrit/) is any organism whose cells have a nucleus and

other organelles enclosed within membranes. Eukaryotes belong to
the taxon Eucarya or Eukaryota. 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 and is enclosed by the nuclear
envelope.[2][3][4] The presence of a nucleus gives eukaryotes their name, which comes from
the Greek (eu, "well" or "true") and (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. Eukaryotic organisms may
be unicellular or multicellular. Only eukaryotes form multicellular organisms consisting of many kinds
of tissue made up of different 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 daughter cells (haploid cells) each with half the number of
chromosomes as the original parent cell. These act as sex cells (gametes each gamete has just
one complement 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 so makes up one of the three domains of
life. 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 1.62.1 billion years ago (during
the Proterozoic eon).

Contents
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1History of the eukaryote concept

2Cell features
o 2.1Internal membrane
o 2.2Mitochondria and plastids
o 2.3Cytoskeletal structures
o 2.4Cell wall
3Differences among eukaryotic cells
o 3.1Animal cell
o 3.2Plant cell
o 3.3Fungal cell
o 3.4Other eukaryotic cells
4Reproduction
5Classification
o 5.1Phylogeny
5.1.1Five supergroups
5.1.2Cavalier-Smith's tree
6Origin of eukaryotes
o 6.1Fossils
o 6.2Relationship to Archaea
o 6.3Endomembrane system and mitochondria
o 6.4Hypotheses for the origin of eukaryotes
6.4.1Autogenous models
 6.4.2Chimeric models
7See also
8References
9External links

History of the eukaryote concept[edit]

Konstantin Mereschkowskiproposed a symbiotic origin for cells with nuclei.

In 1905 and 1910, the Russian biologist Konstantin Mereschkowsky (18551921) 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]
Life timeline
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Orange labels: known ice ages.
Also see: Human timeline and Nature timeline

Eukaryotic cells are typically much larger than those of prokaryotes. 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]

Detail of the endomembrane system and its components

Eukaryote cells include a variety of membrane-bound structures, collectively referred to as
the endomembrane system.[11] Simple compartments, called vesicles or 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. 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.

A 3D rendering of an animal cell cut in half.

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. Various tube- and sheet-like
extensions of the nuclear membrane form what is called the endoplasmic reticulum or ER, which is
involved in protein transport and maturation. It includes the rough ER 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 ER. In most eukaryotes, these protein-carrying vesicles are
released and further modified in stacks of flattened vesicles, called Golgi bodies or dictyosomes.
Vesicles may be specialized for various purposes. For instance, lysosomes contain digestive
enzymes that break down the contents of food vacuoles, and peroxisomes are used to break
down peroxide, which is toxic otherwise. Many protozoa 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 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.[12] They are surrounded by two 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.
In 2016, Monocercomonoides, a metamonad flagellate which resides in the intestines of
the chinchilla, has been found to lack mitochondria entirely. Monocercomonoides obtains its energy
by enzymatic action on nutrients absorbed from the environment. It 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 is considered
to have been lost secondarily.[13][14]
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.
Endosymbiotic origins have also been proposed for the nucleus, for which see below, and for
eukaryotic flagella.[15]

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,[16] 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 basal body, also called a kinetosome or 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., -
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 of one or two,
called kinetids, 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 may also be associated
in the formation of a 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 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]