In biological taxonomy , a domain ( / d ə ˈ m eɪ n / or / d oʊ ˈ m eɪ n / ) ( Latin : regio ), also dominion , superkingdom , realm , or empire , is the highest taxonomic rank of all organisms taken together. It was introduced in the three-domain system of taxonomy devised by Carl Woese , Otto Kandler and Mark Wheelis in 1990.
42-407: A bikont ("two flagella") is any of the eukaryotic organisms classified in the group Bikonta . Many single-celled and multi-celled organisms are members of the group, and these, as well as the presumed ancestor, have two flagella . Another shared trait of bikonts is the fusion of two genes into a single unit: the genes for thymidylate synthase (TS) and dihydrofolate reductase (DHFR) encode
84-405: A microtubular spindle during nuclear division, in the distinctively eukaryotic process of mitosis . Eukaryotes differ from prokaryotes in multiple ways, with unique biochemical pathways such as sterane synthesis. 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,
126-518: A "symbiosis-based phylogeny", giving the description "Eukarya (symbiosis-derived nucleated organisms)". By 2014, a rough consensus started to emerge from the phylogenomic studies of the previous two decades. 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,
168-430: 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 . There is considerable variation in this pattern. Plants have both haploid and diploid multicellular phases . Eukaryotes have lower metabolic rates and longer generation times than prokaryotes, because they are larger and therefore have
210-479: 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 . 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
252-406: 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. 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
294-527: A single protein with two functions. The genes are separately translated in unikonts . Some research suggests that a unikont (a eukaryotic cell with a single flagellum) was the ancestor of opisthokonts (Animals, Fungi, and related forms) and Amoebozoa , and a bikont was the ancestor of Archaeplastida (Plants and relatives), Excavata , Rhizaria , and Chromalveolata . Cavalier-Smith has suggested that Apusozoa , which are typically considered incertae sedis , are in fact bikonts. Relationships within
336-510: A smaller surface area to volume ratio. 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. 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. Since these two species are descendants of lineages that diverged early from
378-407: A synonym for the category of dominion (Lat. dominium ), introduced by Moore in 1974. Carl Linnaeus made the classification "domain" popular in the famous taxonomy system he created in the middle of the eighteenth century. This system was further improved by the studies of Charles Darwin later on but could not classify bacteria easily, as they have very few observable features to compare to
420-417: 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". In 1996, the evolutionary biologist Lynn Margulis proposed to replace kingdoms and domains with "inclusive" names to create
462-610: Is closer in structure to bacterial RNA than to eukaryote RNA. 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. They obtain energy by enzymatic action in the cytoplasm. 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
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#1732772186023504-536: 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"). 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 ; these are separated into two matching sets by
546-1186: The Excavata , has been abandoned as a formal group as it is paraphyletic . The proposed phylogeny below includes only one group of excavates ( Discoba ), and incorporates the 2021 proposal that picozoans are close relatives of rhodophytes. The Provora are a group of microbial predators discovered in 2022. Ancyromonadida [REDACTED] Malawimonada [REDACTED] CRuMs [REDACTED] Amoebozoa [REDACTED] Breviatea [REDACTED] Apusomonadida [REDACTED] Holomycota (inc. fungi) [REDACTED] Holozoa (inc. animals) [REDACTED] ? Metamonada [REDACTED] Discoba [REDACTED] Cryptista [REDACTED] Rhodophyta (red algae) [REDACTED] Picozoa [REDACTED] Glaucophyta [REDACTED] Viridiplantae (plants) [REDACTED] Hemimastigophora [REDACTED] Provora [REDACTED] Haptista [REDACTED] Telonemia [REDACTED] Rhizaria [REDACTED] Alveolata [REDACTED] Stramenopiles [REDACTED] [REDACTED] Domain (biology) According to
588-588: The SAR supergroup and viridiplantae are usually considered monophyletic , Archaeplastida may be paraphyletic, and the mutual relationships between these phyla are still to be fully resolved. Recent reconstructions placed Archaeplastida and Hacrobia together in an "HA supergroup" or "AH supergroup", which was a sister clade to the SAR supergroup within the SAR/HA supergroup . However, this seems to have fallen out of favor as
630-518: The archaea —having a volume of around 10,000 times greater. 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 . The eukaryotes are a diverse lineage, consisting mainly of microscopic organisms . Multicellularity in some form has evolved independently at least 25 times within
672-583: The blue whale , weighing up to 190 tonnes and measuring up to 33.6 metres (110 ft) long, or plants like the coast redwood , up to 120 metres (390 ft) tall. 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. The multicellular eukaryotes include the animals, plants, and fungi , but again, these groups too contain many unicellular species . Eukaryotic cells are typically much larger than those of prokaryotes —the bacteria and
714-562: 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 ). Eukaryotes are organisms that range from microscopic single cells , such as picozoans under 3 micrometres across, to animals like
756-514: 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. 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 , and this group
798-430: The viruses , is not included in this system. Alternatives to the three-domain system include the earlier two-empire system (with the empires Prokaryota and Eukaryota), and the eocyte hypothesis (with two domains of Bacteria and Archaea, with Eukarya included as a branch of Archaea). The term domain was proposed by Carl Woese , Otto Kandler , and Mark Wheelis (1990) in a three-domain system . This term represents
840-560: 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
882-627: The bacterial species on the Earth, or organize them into a tree-like structure (unless the structure includes cross-connections between branches, making it a "network" instead of a "tree"). Members of the domain Eukarya – called eukaryotes – have membrane-bound organelles (including a nucleus containing genetic material) and are represented by five kingdoms : Plantae , Protozoa , Animalia , Chromista , and Fungi . The three-domain system includes no form of non-cellular life . Stefan Luketa proposed
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#1732772186023924-1106: The basis of the three-domain system. While the presence of a nuclear membrane differentiates the Eukarya from the Archaea and Bacteria , both of which lack a nuclear envelope , the Archaea and Bacteria are distinct from each other due to differences in the biochemistry of their cell membranes and RNA markers. Archaea are prokaryotic cells, typically characterized by membrane lipids that are branched hydrocarbon chains attached to glycerol by ether linkages. The presence of these ether linkages in Archaea adds to their ability to withstand extreme temperatures and highly acidic conditions, but many archaea live in mild environments. Halophiles , organisms that thrive in highly salty environments, and hyperthermophiles , organisms that thrive in extremely hot environments, are examples of Archaea. Archaea evolved many cell sizes, but all are relatively small. Their size ranges from 0.1 μm to 15 μm diameter and up to 200 μm long. They are about
966-790: The bikonts are not yet clear. Cavalier-Smith has grouped the Excavata and Rhizaria into the Cabozoa and the Archaeplastida and Chromalveolata into the Corticata , but at least one other study has suggested that the Rhizaria and Chromalveolata form a clade . An alternative to the Unikont–Bikont division was suggested by Derelle et al. in 2015, where they proposed the acronyms Opimoda – Diphoda respectively, as substitutes to
1008-709: 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. 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
1050-404: 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 are cellulose , hemicellulose , and pectin . The cellulose microfibrils are linked together with hemicellulose, embedded in a pectin matrix. The most common hemicellulose in
1092-427: The complex transcription machinery, the membrane-sorting systems, the nuclear pore , and some enzymes in the biochemical pathways. Eukaryote cells include a variety of membrane-bound structures, together forming the endomembrane system. 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
1134-622: The cytoplasm. Mitochondria are organelles in eukaryotic cells. The mitochondrion is commonly called "the powerhouse of the cell", for its function providing energy by oxidising sugars or fats to produce the energy-storing molecule ATP . Mitochondria have two surrounding membranes , each a phospholipid bilayer , the inner of which is folded into invaginations called cristae where aerobic respiration takes place. 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
1176-489: The domain system, the tree of life consists of either three domains, Archaea , Bacteria , and Eukarya , or two domains , Archaea and Bacteria , with Eukarya included in Archaea. In the three-domain model , the first two are prokaryotes , single-celled microorganisms without a membrane-bound nucleus . All organisms that have a cell nucleus and other membrane-bound organelles are included in Eukarya and called eukaryotes . Non-cellular life , most notably
1218-603: The ether linkages that Archaea have. Internally, bacteria have different RNA structures in their ribosomes , hence they are grouped into a different category. In the two- and three-domain systems, this puts them into a separate domain. There is a great deal of diversity in the domain Bacteria . That diversity is further confounded by the exchange of genes between different bacterial lineages. The occurrence of duplicate genes between otherwise distantly-related bacteria makes it nearly impossible to distinguish bacterial species, count
1260-406: The eukaryotes. 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 . Eukaryotes are grouped by genomic similarities, so that groups often lack visible shared characteristics. The defining feature of eukaryotes
1302-484: The eukaryotic evolutionary tree, core meiotic genes, and hence sex, were likely present in the common ancestor of eukaryotes. Species once thought to be asexual, such as Leishmania parasites, have a sexual cycle. Amoebae, previously regarded as asexual, may be anciently sexual; while present-day asexual groups could have arisen recently. In antiquity , the two lineages of animals and plants were recognized by Aristotle and Theophrastus . The lineages were given
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1344-646: 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. The capture and sequestering of photosynthetic cells and chloroplasts, kleptoplasty , occurs in many types of modern eukaryotic organisms. The cytoskeleton provides stiffening structure and points of attachment for motor structures that enable
1386-435: The monophyly of hacrobia has come under dispute. Eukaryote The eukaryotes ( / j uː ˈ k ær i oʊ t s , - ə t s / yoo- KARR -ee-ohts, -əts ) 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
1428-417: The older terms. The name Diphoda is formed from the letters of DIscoba and diaPHOretickes (shown in capitals). [suggested singular forms are Opneme-Dipheme respectively] A "classical" cladogram (data from 2012, 2015) is: Archaeplastida (inc. land plants) Cryptophyta Haptophyta Rhizaria Heterokont Alveolata Excavata Unikonta However, a cladogram (data from 2015, 2016) with
1470-573: The other domains. Carl Woese made a revolutionary breakthrough when, in 1977, he compared the nucleotide sequences of the 16s ribosomal RNA and discovered that the rank "domain" contained three branches, not two as scientists had previously thought. Initially, due to their physical similarities, Archaea and Bacteria were classified together and called "archaebacteria". However, scientists now know that these two domains are hardly similar and are internally distinctly different. Each of these three domains contains unique ribosomal RNA . This forms
1512-449: The outer membrane invaginates and then pinches off to form a vesicle. Some cell products can leave in a vesicle through exocytosis . The nucleus is surrounded by a double membrane known as the nuclear envelope , with nuclear pores 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
1554-417: The primary cell wall is xyloglucan . 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
1596-399: The root in Excavata is Archaeplastida (inc. land plants) Cryptista Haptista Rhizaria Heterokonta Alveolata Discoba Metamonada ? Metamonada ? Neolouka Collodictyon Varisulca Amoebozoa Apusomonadida Breviata Opisthokonta The corticates correspond roughly to the bikonts. While Haptophyta , Cryptophyta , Glaucophyta , Rhodophyta ,
1638-498: 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. In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles ( cisternae ), the Golgi apparatus . Vesicles may be specialized; for instance, lysosomes contain digestive enzymes that break down biomolecules in
1680-454: The size of bacteria, or similar in size to the mitochondria found in eukaryotic cells. Members of the genus Thermoplasma are the smallest of the Archaea. Cyanobacteria and mycoplasmas are two examples of bacteria. Even though bacteria are prokaryotic cells just like Archaea, their cell membranes are instead made of phospholipid bilayers . Bacteria cell membranes are distinct from Archean membranes: They characteristically have none of
1722-599: 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. The eukaryotes seemingly emerged within the Asgard archaea , and are closely related to the Heimdallarchaeia . This implies that there are only two domains of life , Bacteria and Archaea, with eukaryotes incorporated among
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1764-503: Was expanded until Ernst Haeckel made it a kingdom encompassing all single-celled eukaryotes, the Protista , in 1866. 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. Understanding of the oldest branchings in the tree of life only developed substantially with DNA sequencing , leading to
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