Eukaryote

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Eukaryota
Temporal range: Statherian–Present 1650–0 Ma
Rhodomonas salina CCMP 322.jpg
Ranunculus asiaticus4LEST.jpg
Viridiplantae (plants)
Trypanosoma sp. PHIL 613 lores.jpg
Chaos carolinensis Wilson 1900.jpg
Ammonia tepida.jpg
Isotricha intestinalis.jpg
Osmia rufa couple (aka).jpg
Boletus edulis (Tillegem).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
(Chatton, 1925) Whittaker & Margulis, 1978
Subgroups
Synonyms

The eukaryotes ( /juːˈkærioʊts,-əts/ yoo-KARR-ee-ohts, -⁠əts) [4] constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes: the Bacteria and the Archaea. Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass is much larger than that of prokaryotes.

Contents

The eukaryotes seemingly emerged within the Asgard archaea, and are closely related to the Heimdallarchaeia. [5] This implies that there are only two domains of life, Bacteria and Archaea, with eukaryotes incorporated among the Archaea. Eukaryotes first emerged during the Paleoproterozoic, likely as flagellated cells. The leading evolutionary theory is they were created by symbiogenesis between an anaerobic Asgard archaean and an aerobic proteobacterium, which formed the mitochondria. A second episode of symbiogenesis with a cyanobacterium created the plants, with chloroplasts.

Eukaryotic cells contain membrane-bound organelles such as the nucleus, the endoplasmic reticulum, and the Golgi apparatus. Eukaryotes may be either unicellular or multicellular. In comparison, prokaryotes are typically unicellular. Unicellular eukaryotes are sometimes called protists. Eukaryotes can reproduce both asexually through mitosis and sexually through meiosis and gamete fusion (fertilization).

Diversity

Eukaryotes are organisms that range from microscopic single cells, such as picozoans under 3 micrometres across, [6] to animals like the blue whale, weighing up to 190 tonnes and measuring up to 33.6 metres (110 ft) long, [7] or plants like the coast redwood, up to 120 metres (390 ft) tall. [8] Many eukaryotes are unicellular; the informal grouping called protists includes many of these, with some multicellular forms like the giant kelp up to 200 feet (61 m) long. [9] The multicellular eukaryotes include the animals, plants, and fungi, but again, these groups too contain many unicellular species. [10] Eukaryotic cells are typically much larger than those of prokaryotes—the bacteria and the archaea—having a volume of around 10,000 times greater. [11] [12] Eukaryotes represent a small minority of the number of organisms, but, as many of them are much larger, their collective global biomass (468 gigatons) is far larger than that of prokaryotes (77 gigatons), with plants alone accounting for over 81% of the total biomass of Earth. [13]

The eukaryotes are a diverse lineage, consisting mainly of microscopic organisms. [14] Multicellularity in some form has evolved independently at least 25 times within the eukaryotes. [15] [16] Complex multicellular organisms, not counting the aggregation of amoebae to form slime molds, have evolved within only six eukaryotic lineages: animals, symbiomycotan fungi, brown algae, red algae, green algae, and land plants. [17] Eukaryotes are grouped by genomic similarities, so that groups often lack visible shared characteristics. [14]

Distinguishing features

Nucleus

The defining feature of eukaryotes is that their cells have nuclei. This gives them their name, from the Greek εὖ (eu, "well" or "good") and κάρυον (karyon, "nut" or "kernel", here meaning "nucleus"). [18] Eukaryotic cells have a variety of internal membrane-bound structures, called organelles, and a cytoskeleton which defines the cell's organization and shape. The nucleus stores the cell's DNA, which is divided into linear bundles called chromosomes; [19] these are separated into two matching sets by a microtubular spindle during nuclear division, in the distinctively eukaryotic process of mitosis. [20]

Biochemistry

Eukaryotes differ from prokaryotes in multiple ways, with unique biochemical pathways such as sterane synthesis. [21] The eukaryotic signature proteins have no homology to proteins in other domains of life, but appear to be universal among eukaryotes. They include the proteins of the cytoskeleton, the complex transcription machinery, the membrane-sorting systems, the nuclear pore, and some enzymes in the biochemical pathways. [22]

Internal membranes

Prokaryote cell.svg
Prokaryote, to same scale
Endomembrane system diagram en (edit).svg
Eukaryotic cell with endomembrane system
Eukaryotic cells are some 10,000 times larger than prokaryotic cells by volume, and contain membrane-bound organelles.

Eukaryote cells include a variety of membrane-bound structures, together forming the endomembrane system. [23] Simple compartments, called vesicles and vacuoles, can form by budding off other membranes. Many cells ingest food and other materials through a process of endocytosis, where the outer membrane invaginates and then pinches off to form a vesicle. [24] Some cell products can leave in a vesicle through exocytosis. [25]

The nucleus is surrounded by a double membrane known as the nuclear envelope, with nuclear pores that allow material to move in and out. [26] 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, covered in ribosomes which synthesize proteins; these enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth endoplasmic reticulum. [27] In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles (cisternae), the Golgi apparatus. [28]

Vesicles may be specialized; for instance, lysosomes contain digestive enzymes that break down biomolecules in the cytoplasm. [29]

Mitochondria

Mitochondria are essentially universal in the eukaryotes, and with their own DNA somewhat resemble prokaryotic cells. Mitochondrion structure.svg
Mitochondria are essentially universal in the eukaryotes, and with their own DNA somewhat resemble prokaryotic cells.

Mitochondria are organelles in eukaryotic cells. The mitochondrion is commonly called "the powerhouse of the cell", [30] for its function providing energy by oxidising sugars or fats to produce the energy-storing molecule ATP. [31] [32] Mitochondria have two surrounding membranes, each a phospholipid bilayer, the inner of which is folded into invaginations called cristae where aerobic respiration takes place. [33]

Mitochondria contain their own DNA, which has close structural similarities to bacterial DNA, from which it originated, and which encodes rRNA and tRNA genes that produce RNA which is closer in structure to bacterial RNA than to eukaryote RNA. [34]

Some eukaryotes, such as the metamonads Giardia and Trichomonas , and the amoebozoan Pelomyxa , appear to lack mitochondria, but all contain mitochondrion-derived organelles, like hydrogenosomes or mitosomes, having lost their mitochondria secondarily. [35] They obtain energy by enzymatic action in the cytoplasm. [36] [35]

Plastids

The most common type of plastid is the chloroplast, which contains chlorophyll and produces organic compounds by photosynthesis. Chloroplast II.svg
The most common type of plastid is the chloroplast, which contains chlorophyll and produces organic compounds by photosynthesis.

Plants and various groups of algae have plastids as well as mitochondria. Plastids, like mitochondria, 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. [37] The capture and sequestering of photosynthetic cells and chloroplasts, kleptoplasty, occurs in many types of modern eukaryotic organisms. [38] [39]

Cytoskeletal structures

The cytoskeleton. Actin filaments are shown in red, microtubules in green. (The nucleus is in blue.) FluorescentCells.jpg
The cytoskeleton. Actin filaments are shown in red, microtubules in green. (The nucleus is in blue.)

The cytoskeleton provides stiffening structure and points of attachment for motor structures that enable the cell to move, change shape, or transport materials. The motor structures are microfilaments of actin and actin-binding proteins, including α-actinin, fimbrin, and filamin are present in submembranous cortical layers and bundles. Motor proteins of microtubules, dynein and kinesin, and myosin of actin filaments, provide dynamic character of the network. [40] [41]

Many eukaryotes have long slender motile cytoplasmic projections, called flagella, or multiple shorter structures called cilia. These organelles are variously involved in movement, feeding, and sensation. They are composed mainly of tubulin, and are entirely distinct from prokaryotic flagella. They are supported by a bundle of microtubules arising from a centriole, characteristically arranged as nine doublets surrounding two singlets. Flagella may have hairs (mastigonemes), as in many Stramenopiles. Their interior is continuous with the cell's cytoplasm. [42] [43]

Centrioles are often present, even in cells and groups that do not have flagella, but conifers and flowering plants have neither. They generally occur in groups that give rise to various microtubular roots. These form a primary component of the cytoskeleton, 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. [44]

Cell wall

The cells of plants, algae, fungi and most chromalveolates, but not animals, are surrounded by a cell wall. This is 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. [45]

The major polysaccharides making up the primary cell wall of land plants are cellulose, hemicellulose, and pectin. The cellulose microfibrils are linked together with hemicellulose, embedded in a pectin matrix. The most common hemicellulose in the primary cell wall is xyloglucan. [46]

Sexual reproduction

Sexual reproduction requires a life cycle that alternates between a haploid phase, with one copy of each chromosome in the cell, and a diploid phase, with two copies. In eukaryotes, haploid gametes are produced by meiosis; two gametes fuse to form a diploid zygote. Sexual cycle N-2N.svg
Sexual reproduction requires a life cycle that alternates between a haploid phase, with one copy of each chromosome in the cell, and a diploid phase, with two copies. In eukaryotes, haploid gametes are produced by meiosis; two gametes fuse to form a diploid zygote.

Eukaryotes 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, with two copies of each chromosome in each cell. The diploid phase is formed by fusion of two haploid gametes, such as eggs and spermatozoa, to form a zygote; this may grow into a body, with its cells dividing by mitosis, and at some stage produce haploid gametes through meiosis, a division that reduces the number of chromosomes and creates genetic variability. [47] There is considerable variation in this pattern. Plants have both haploid and diploid multicellular phases. [48] Eukaryotes have lower metabolic rates and longer generation times than prokaryotes, because they are larger and therefore have a smaller surface area to volume ratio. [49]

The evolution of sexual reproduction may be a primordial characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and Roger have proposed that facultative sex was present in the group's common ancestor. [50] A core set of genes that function in meiosis is present in both Trichomonas vaginalis and Giardia intestinalis , two organisms previously thought to be asexual. [51] [52] Since these two species are descendants of lineages that diverged early from the eukaryotic evolutionary tree, core meiotic genes, and hence sex, were likely present in the common ancestor of eukaryotes. [51] [52] Species once thought to be asexual, such as Leishmania parasites, have a sexual cycle. [53] Amoebae, previously regarded as asexual, may be anciently sexual; while present-day asexual groups could have arisen recently. [54]

Evolution

Tree of eukaryotes showing major subgroups and thumbnail diagrams of representative members of each group. Updated synthesis based on recent (as of 2023) phylogenomic reconstructions. Openly available illustrations as tools to describe eukaryotic microbial diversity - Journal.pbio.3002395.g001.tif
Tree of eukaryotes showing major subgroups and thumbnail diagrams of representative members of each group. Updated synthesis based on recent (as of 2023) phylogenomic reconstructions.

History of classification

In antiquity, the two lineages of animals and plants were recognized by Aristotle and Theophrastus. The lineages were given the taxonomic rank of Kingdom by Linnaeus in the 18th century. Though he included the fungi with plants with some reservations, it was later realized that they are quite distinct and warrant a separate kingdom. [56] 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 to refer to organisms such as ciliates, [57] and this group was expanded until Ernst Haeckel made it a kingdom encompassing all single-celled eukaryotes, the Protista, in 1866. [58] [59] [60] The eukaryotes thus came to be seen as four kingdoms:

The protists were at that time thought to be "primitive forms", and thus an evolutionary grade, united by their primitive unicellular nature. [59] Understanding of the oldest branchings in the tree of life only developed substantially with DNA sequencing, leading to a system of domains rather than kingdoms as top level rank being put forward by Carl Woese, Otto Kandler, and Mark Wheelis in 1990, uniting all the eukaryote kingdoms in the domain "Eucarya", stating, however, that "'eukaryotes' will continue to be an acceptable common synonym". [2] [61] In 1996, the evolutionary biologist Lynn Margulis proposed to replace Kingdoms and Domains with "inclusive" names to create a "symbiosis-based phylogeny", giving the description "Eukarya (symbiosis-derived nucleated organisms)". [3]

Phylogeny

By 2014, a rough consensus started to emerge from the phylogenomic studies of the previous two decades. [10] [62] The majority of eukaryotes can be placed in one of two large clades dubbed Amorphea (similar in composition to the unikont hypothesis) and the Diphoda (formerly bikonts), which includes plants and most algal lineages. A third major grouping, the Excavata, has been abandoned as a formal group as it is paraphyletic. [63] The proposed phylogeny below includes only one group of excavates (Discoba), [64] and incorporates the 2021 proposal that picozoans are close relatives of rhodophytes. [65] The Provora are a group of microbial predators discovered in 2022. [1]

Eukaryotes
2200 mya

One view of the great kingdoms and their stem groups. [64] [66] [67] [14] The Metamonada are hard to place, being sister possibly to Discoba or to Malawimonada [14] or being a paraphyletic group external to all other eukaryotes. [68]

Origin of eukaryotes

In the theory of symbiogenesis, a merger of an archaean and an aerobic bacterium created the eukaryotes, with aerobic mitochondria; a second merger added chloroplasts, creating the green plants. Symbiogenesis 2 mergers.svg
In the theory of symbiogenesis, a merger of an archaean and an aerobic bacterium created the eukaryotes, with aerobic mitochondria; a second merger added chloroplasts, creating the green plants.

The origin of the eukaryotic cell, or eukaryogenesis, is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. The last eukaryotic common ancestor (LECA) is the hypothetical origin of all living eukaryotes, [70] and was most likely a biological population, not a single individual. [71] The LECA is believed to have been a protist with a nucleus, at least one centriole and flagellum, facultatively aerobic mitochondria, sex (meiosis and syngamy), a dormant cyst with a cell wall of chitin or cellulose, and peroxisomes. [72] [73] [74]

An endosymbiotic union between a motile anaerobic archaean and an aerobic alphaproteobacterium gave rise to the LECA and all eukaryotes, with mitochondria. A second, much later endosymbiosis with a cyanobacterium gave rise to the ancestor of plants, with chloroplasts. [69]

The presence of eukaryotic biomarkers in archaea points towards an archaeal origin. The genomes of Asgard archaea have plenty of Eukaryotic signature protein genes, which play a crucial role in the development of the cytoskeleton and complex cellular structures characteristic of eukaryotes. In 2022, cryo-electron tomography demonstrated that Asgard archaea have a complex actin-based cytoskeleton, providing the first direct visual evidence of the archaeal ancestry of eukaryotes. [75]

Fossils

The timing of the origin of eukaryotes is hard to determine but the discovery of Qingshania magnificia, the earliest multicelluar eukaryote from North China which lived during 1.635 billion years ago, suggests that the crown group eukaryotes would have originated from the late Paleoproterozoic (Statherian); the earliest unequivocal unicellular eukaryotes which lived during approximately 1.65 billion years ago are also discovered from North China: Tappania plana, Shuiyousphaeridium macroreticulatum, Dictyosphaera macroreticulata, Germinosphaera alveolata, and Valeria lophostriata. [76]

Some acritarchs are known from at least 1.65 billion years ago, and a fossil, Grypania , which may be an alga, is as much as 2.1 billion years old. [77] [78] The "problematic" [79] fossil Diskagma has been found in paleosols 2.2 billion years old. [79]

Reconstruction of the problematic Diskagma buttonii, a terrestrial fossil less than 1mm high, from rocks around 2.2 billion years old Diskagma butonii.jpg
Reconstruction of the problematic Diskagma buttonii , a terrestrial fossil less than 1mm high, from rocks around 2.2 billion years old

Structures proposed to represent "large colonial organisms" have been found in the black shales of the Palaeoproterozoic such as the Francevillian B Formation, in Gabon, dubbed the "Francevillian biota" which is dated at 2.1 billion years old. [80] [81] However, the status of these structures as fossils is contested, with other authors suggesting that they might represent pseudofossils. [82] The oldest fossils than can unambiguously be assigned to eukaryotes are from the Ruyang Group of China, dating to approximately 1.8-1.6 billion years ago. [83] Fossils that are clearly related to modern groups start appearing an estimated 1.2 billion years ago, in the form of red algae, though recent work suggests the existence of fossilized filamentous algae in the Vindhya basin dating back perhaps to 1.6 to 1.7 billion years ago. [84]

The presence of steranes, eukaryotic-specific biomarkers, in Australian shales previously indicated that eukaryotes were present in these rocks dated at 2.7 billion years old, [21] [85] but these Archaean biomarkers have been rebutted as later contaminants. [86] The oldest valid biomarker records are only around 800 million years old. [87] In contrast, a molecular clock analysis suggests the emergence of sterol biosynthesis as early as 2.3 billion years ago. [88] The nature of steranes as eukaryotic biomarkers is further complicated by the production of sterols by some bacteria. [89] [90]

Whenever their origins, eukaryotes may not have become ecologically dominant until much later; a massive increase in the zinc composition of marine sediments 800  million years ago 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). [91]

See also

Related Research Articles

<span class="mw-page-title-main">Cell (biology)</span> Basic unit of many life forms

The cell is the basic structural and functional unit of all forms of life. Every cell consists of cytoplasm enclosed within a membrane; many cells contain organelles, each with a specific function. The term comes from the Latin word cellula meaning 'small room'. Most cells are only visible under a microscope. Cells emerged on Earth about 4 billion years ago. All cells are capable of replication, protein synthesis, and motility.

<span class="mw-page-title-main">Microorganism</span> Microscopic living organism

A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells.

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 suffix -elle being a diminutive. Organelles are either separately enclosed within their own lipid bilayers or are spatially distinct functional units without a surrounding lipid bilayer. Although most organelles are functional units within cells, some function units that extend outside of cells are often termed organelles, such as cilia, the flagellum and archaellum, and the trichocyst.

<span class="mw-page-title-main">Kingdom (biology)</span> Taxonomic rank

In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.

<span class="mw-page-title-main">Symbiogenesis</span> Evolutionary theory holding that eukaryotic organelles evolved through symbiosis with prokaryotes

Symbiogenesis 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 other organelles of eukaryotic cells are descended from formerly free-living prokaryotes taken one inside the other in endosymbiosis. Mitochondria appear to be phylogenetically related to Rickettsiales bacteria, while chloroplasts are thought to be related to cyanobacteria.

<span class="mw-page-title-main">Plastid</span> Plant cell organelles that perform photosynthesis and store starch

A plastid is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. Plastids are considered to be intracellular endosymbiotic cyanobacteria.

<span class="mw-page-title-main">Three-domain system</span> Hypothesis for classification of life

The three-domain system is a taxonomic classification system that groups all cellular life into three domains, namely Archaea, Bacteria and Eukarya, introduced by Carl Woese, Otto Kandler and Mark Wheelis in 1990. The key difference from earlier classifications such as the two-empire system and the five-kingdom classification is the splitting of Archaea from Bacteria as completely different organisms. It has been challenged by the two-domain system that divides organisms into Bacteria and Archaea only, as Eukaryotes are considered as a clade of Archaea.

<span class="mw-page-title-main">Chromista</span> Eukaryotic biological kingdom

Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.

<span class="mw-page-title-main">Unicellular organism</span> Organism that consists of only one cell

A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and eukaryotic organisms. Most prokaryotes are unicellular and are classified into bacteria and archaea. Many eukaryotes are multicellular, but some are unicellular such as protozoa, unicellular algae, and unicellular fungi. Unicellular organisms are thought to be the oldest form of life, with early protocells possibly emerging 3.5–4.1 billion years ago.

<span class="mw-page-title-main">Multicellular organism</span> Organism that consists of more than one cell

A multicellular organism is an organism that consists of more than one cell, unlike unicellular organisms. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and partially multicellular, like slime molds and social amoebae such as the genus Dictyostelium.

<span class="mw-page-title-main">Green algae</span> Paraphyletic group of eukaryotes

The green algae are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as a sister of the Zygnematophyceae. Since the realization that the Embryophytes emerged within the green algae, some authors are starting to include them. The completed clade that includes both green algae and embryophytes is monophyletic and is referred to as the clade Viridiplantae and as the kingdom Plantae. The green algae include unicellular and colonial flagellates, most with two flagella per cell, as well as various colonial, coccoid (spherical), and filamentous forms, and macroscopic, multicellular seaweeds. There are about 22,000 species of green algae, many of which live most of their lives as single cells, while other species form coenobia (colonies), long filaments, or highly differentiated macroscopic seaweeds.

<span class="mw-page-title-main">Viridiplantae</span> Clade of archaeplastids including green algae and the land plants

Viridiplantae constitute a clade of eukaryotic organisms that comprises approximately 450,000–500,000 species that play important roles in both terrestrial and aquatic ecosystems. They include the green algae, which are primarily aquatic, and the land plants (embryophytes), which emerged from within them. Green algae traditionally excludes the land plants, rendering them a paraphyletic group. However it is accurate to think of land plants as a kind of alga. Since the realization that the embryophytes emerged from within the green algae, some authors are starting to include them. They have cells with cellulose in their cell walls, and primary chloroplasts derived from endosymbiosis with cyanobacteria that contain chlorophylls a and b and lack phycobilins. Corroborating this, a basal phagotroph archaeplastida group has been found in the Rhodelphydia.

<span class="mw-page-title-main">Archaeplastida</span> Clade of eukaryotes containing land plants and some algae

The Archaeplastida are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans. The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by phagocytosis of a cyanobacterium. All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae. Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.

<span class="mw-page-title-main">Prokaryote</span> Unicellular organism lacking a membrane-bound nucleus

A prokaryote is a single-cell organism whose cell lacks a nucleus and other membrane-bound organelles. The word prokaryote comes from the Ancient Greek πρό (pró), meaning 'before', and κάρυον (káruon), meaning 'nut' or 'kernel'. In the two-empire system arising from the work of Édouard Chatton, prokaryotes were classified within the empire Prokaryota. However in the three-domain system, based upon molecular analysis, prokaryotes are divided into two domains: Bacteria and Archaea. Organisms with nuclei are placed in a third domain: Eukaryota.

<span class="mw-page-title-main">Lokiarchaeota</span> Phylum of archaea

Lokiarchaeota is a proposed phylum of the Archaea. The phylum includes all members of the group previously named Deep Sea Archaeal Group, also known as Marine Benthic Group B. Lokiarchaeota is part of the superphylum Asgard containing the phyla: Lokiarchaeota, Thorarchaeota, Odinarchaeota, Heimdallarchaeota, and Helarchaeota. A phylogenetic analysis disclosed a monophyletic grouping of the Lokiarchaeota with the eukaryotes. The analysis revealed several genes with cell membrane-related functions. The presence of such genes support the hypothesis of an archaeal host for the emergence of the eukaryotes; the eocyte-like scenarios.

<span class="mw-page-title-main">Ocelloid</span>

An ocelloid is a subcellular structure found in the family Warnowiaceae (warnowiids), which are members of a group of unicellular organisms known as dinoflagellates. The ocelloid is analogous in structure and function to the eyes of multicellular organisms, which focus, process and detect light. The ocelloid is much more complex than the eyespot, a light-sensitive structure also found in unicellular organisms, and is in fact one of the most complex known subcellular structures. It has been described as a striking example of convergent evolution.

<span class="mw-page-title-main">Eukaryogenesis</span> Process of forming the first eukaryotic cell

Eukaryogenesis, the process which created the eukaryotic cell and lineage, is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. The process is widely agreed to have involved symbiogenesis, in which an archeon and a bacterium came together to create the first eukaryotic common ancestor (FECA). This cell had a new level of complexity and capability, with a nucleus, at least one centriole and cilium, facultatively aerobic mitochondria, sex, a dormant cyst with a cell wall of chitin and/or cellulose and peroxisomes. It evolved into a population of single-celled organisms that included the last eukaryotic common ancestor (LECA), gaining capabilities along the way, though the sequence of the steps involved has been disputed, and may not have started with symbiogenesis. In turn, the LECA gave rise to the eukaryotes' crown group, containing the ancestors of animals, fungi, plants, and a diverse range of single-celled organisms.

A plastid is a membrane-bound organelle found in plants, algae and other eukaryotic organisms that contribute to the production of pigment molecules. Most plastids are photosynthetic, thus leading to color production and energy storage or production. There are many types of plastids in plants alone, but all plastids can be separated based on the number of times they have undergone endosymbiotic events. Currently there are three types of plastids; primary, secondary and tertiary. Endosymbiosis is reputed to have led to the evolution of eukaryotic organisms today, although the timeline is highly debated.

<span class="mw-page-title-main">Marine prokaryotes</span> Marine bacteria and marine archaea

Marine prokaryotes are marine bacteria and marine archaea. They are defined by their habitat as prokaryotes that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. All cellular life forms can be divided into prokaryotes and eukaryotes. Eukaryotes are organisms whose cells have a nucleus enclosed within membranes, whereas prokaryotes are the organisms that do not have a nucleus enclosed within a membrane. The three-domain system of classifying life adds another division: the prokaryotes are divided into two domains of life, the microscopic bacteria and the microscopic archaea, while everything else, the eukaryotes, become the third domain.

<span class="mw-page-title-main">Two-domain system</span> Biological classification system

The two-domain system is a biological classification by which all organisms in the tree of life are classified into two domains, Bacteria and Archaea. It emerged from development of knowledge of archaea diversity and challenges the widely accepted three-domain system that classifies life into Bacteria, Archaea, and Eukarya. It was preceded by the eocyte hypothesis of James A. Lake in the 1980s, which was largely superseded by the three-domain system, due to evidence at the time. Better understanding of archaea, especially of their roles in the origin of eukaryotes through symbiogenesis with bacteria, led to the revival of the eocyte hypothesis in the 2000s. The two-domain system became more widely accepted after the discovery of a large group (superphylum) of archaea called Asgard in 2017, which evidence suggests to be the evolutionary root of eukaryotes, thereby making eukaryotes members of the domain Archaea.

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