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A prokaryote ( / p r oʊ ˈ k ær i oʊ t , - ə t / ; less commonly spelled procaryote ) 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 (formerly Eubacteria) and Archaea (formerly Archaebacteria). Organisms with nuclei are placed in a third domain: Eukaryota .

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61-734: In the classification of eukaryotes (living organisms with a cell nucleus), Cabozoa was a taxon proposed by Cavalier-Smith . It was a putative clade comprising the Rhizaria and Excavata . More recent research places the Rhizaria with the Alveolata and Stramenopiles instead of the Excavata, however, so "Cabozoa" is polyphyletic. This Bikont -related article is a stub . You can help Misplaced Pages by expanding it . Eukaryote The eukaryotes ( / j uː ˈ k ær i oʊ t s , - ə t s / yoo- KARR -ee-ohts, -⁠əts ) constitute

122-409: A haploid chromosomal composition that is partially replicated, a condition known as merodiploidy . Prokaryotes lack mitochondria and chloroplasts . Instead, processes such as oxidative phosphorylation and photosynthesis take place across the prokaryotic cell membrane . However, prokaryotes do possess some internal structures, such as prokaryotic cytoskeletons . It has been suggested that

183-405: 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,

244-688: A paraphyletic group, just like dinosaurs without birds. Unlike the above assumption of a fundamental split between prokaryotes and eukaryotes, the most important difference between biota may be the division between Bacteria and the rest (Archaea and Eukaryota). For instance, DNA replication differs fundamentally between the Bacteria and Archaea (including that in eukaryotic nuclei), and it may not be homologous between these two groups. Moreover, ATP synthase , though common (homologous) in all organisms, differs greatly between bacteria (including eukaryotic organelles such as mitochondria and chloroplasts ) and

305-1088: A prokaryotic cytoskeleton that is more primitive than that of the eukaryotes. Besides homologues of actin and tubulin ( MreB and FtsZ ), the helically arranged building-block of the flagellum , flagellin , is one of the most significant cytoskeletal proteins of bacteria, as it provides structural backgrounds of chemotaxis , the basic cell physiological response of bacteria. At least some prokaryotes also contain intracellular structures that can be seen as primitive organelles. Membranous organelles (or intracellular membranes) are known in some groups of prokaryotes, such as vacuoles or membrane systems devoted to special metabolic properties, such as photosynthesis or chemolithotrophy . In addition, some species also contain carbohydrate-enclosed microcompartments, which have distinct physiological roles (e.g. carboxysomes or gas vacuoles). Most prokaryotes are between 1 μm and 10 μm, but they can vary in size from 0.2 μm ( Mycoplasma genitalium ) to 750 μm ( Thiomargarita namibiensis ). Prokaryotic cells have various shapes;

366-401: A taxon to be found nearby the then-unknown Asgard group). For example, histones which usually package DNA in eukaryotic nuclei, have also been found in several archaean groups, giving evidence for homology . This idea might clarify the mysterious predecessor of eukaryotic cells ( eucytes ) which engulfed an alphaproteobacterium forming the first eucyte ( LECA , l ast e ukaryotic c ommon

427-513: 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,

488-1198: A bacterium to bind, take up and recombine donor DNA into its own chromosome, it must first enter a special physiological state called competence . About 40 genes are required in Bacillus subtilis for the development of competence. The length of DNA transferred during B. subtilis transformation can be as much as a third to the whole chromosome. Transformation is a common mode of DNA transfer, and 67 prokaryotic species are thus far known to be naturally competent for transformation. Among archaea, Halobacterium volcanii forms cytoplasmic bridges between cells that appear to be used for transfer of DNA from one cell to another. Another archaeon, Sulfolobus solfataricus , transfers DNA between cells by direct contact. Frols et al. (2008) found that exposure of S. solfataricus to DNA damaging agents induces cellular aggregation, and suggested that cellular aggregation may enhance DNA transfer among cells to provide increased repair of damaged DNA via homologous recombination. While prokaryotes are considered strictly unicellular, most can form stable aggregate communities. When such communities are encased in

549-430: 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

610-479: 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

671-399: A higher metabolic rate , a higher growth rate, and as a consequence, a shorter generation time than eukaryotes. There is increasing evidence that the roots of the eukaryotes are to be found in (or at least next to) the archaean Asgard group, perhaps Heimdallarchaeota (an idea which is a modern version of the 1984 eocyte hypothesis , eocytes being an old synonym for Thermoproteota ,

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732-521: A ncestor) according to endosymbiotic theory . There might have been some additional support by viruses, called viral eukaryogenesis . The non-bacterial group comprising archaea and eukaryota was called Neomura by Thomas Cavalier-Smith in 2002. However, in a cladistic view, Eukaryota are Archaea in the same sense as birds are dinosaurs because they evolved from the Maniraptora dinosaur group. In contrast, archaea without eukaryota appear to be

793-659: A nucleus. Both eukaryotes and prokaryotes contain large RNA / protein structures called ribosomes , which produce protein , but the ribosomes of prokaryotes are smaller than those of eukaryotes. Mitochondria and chloroplasts , two organelles found in many eukaryotic cells, contain ribosomes similar in size and makeup to those found in prokaryotes. This is one of many pieces of evidence that mitochondria and chloroplasts are descended from free-living bacteria. The endosymbiotic theory holds that early eukaryotic cells took in primitive prokaryotic cells by phagocytosis and adapted themselves to incorporate their structures, leading to

854-406: 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

915-690: A similar group of selfish individuals (see inclusive fitness and Hamilton's rule ). Should these instances of prokaryotic sociality prove to be the rule rather than the exception, it would have serious implications for the way we view prokaryotes in general, and the way we deal with them in medicine. Bacterial biofilms may be 100 times more resistant to antibiotics than free-living unicells and may be nearly impossible to remove from surfaces once they have colonized them. Other aspects of bacterial cooperation—such as bacterial conjugation and quorum-sensing-mediated pathogenicity , present additional challenges to researchers and medical professionals seeking to treat

976-473: A single founder (in the way that animals and plants are founded by single cells), which presents a number of theoretical issues. Most explanations of co-operation and the evolution of multicellularity have focused on high relatedness between members of a group (or colony, or whole organism). If a copy of a gene is present in all members of a group, behaviors that promote cooperation between members may permit those members to have (on average) greater fitness than

1037-510: 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

1098-679: A stabilizing polymer matrix ("slime"), they may be called " biofilms ". Cells in biofilms often show distinct patterns of gene expression (phenotypic differentiation) in time and space. Also, as with multicellular eukaryotes, these changes in expression often appear to result from cell-to-cell signaling , a phenomenon known as quorum sensing . Biofilms may be highly heterogeneous and structurally complex and may attach to solid surfaces, or exist at liquid-air interfaces, or potentially even liquid-liquid interfaces. Bacterial biofilms are often made up of microcolonies (approximately dome-shaped masses of bacteria and matrix) separated by "voids" through which

1159-417: 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

1220-404: Is evidence on Mars of fossil or living prokaryotes. However, this possibility remains the subject of considerable debate and skepticism. The division between prokaryotes and eukaryotes is usually considered the most important distinction or difference among organisms. The distinction is that eukaryotic cells have a "true" nucleus containing their DNA , whereas prokaryotic cells do not have

1281-610: 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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1342-618: 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 the Archaea. Eukaryotes first emerged during the Paleoproterozoic , likely as flagellated cells. The leading evolutionary theory

1403-532: 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

1464-864: 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 ). Eukaryotes are organisms that range from microscopic single cells , such as picozoans under 3 micrometres across, to animals like

1525-1248: 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] Prokaryote Prokaryotes evolved before eukaryotes, and lack nuclei, mitochondria , and most of

1586-518: 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

1647-583: 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

1708-548: 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

1769-510: 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

1830-608: The archaea/eukaryote nucleus group. The last common antecessor of all life (called LUCA , l ast u niversal c ommon a ncestor) should have possessed an early version of this protein complex. As ATP synthase is obligate membrane bound, this supports the assumption that LUCA was a cellular organism. The RNA world hypothesis might clarify this scenario, as LUCA might have been a ribocyte (also called ribocell) lacking DNA, but with an RNA genome built by ribosomes as primordial self-replicating entities . A Peptide-RNA world (also called RNP world) hypothesis has been proposed based on

1891-478: The associated diseases. Prokaryotes have diversified greatly throughout their long existence. The metabolism of prokaryotes is far more varied than that of eukaryotes, leading to many highly distinct prokaryotic types. For example, in addition to using photosynthesis or organic compounds for energy, as eukaryotes do, prokaryotes may obtain energy from inorganic compounds such as hydrogen sulfide . This enables prokaryotes to thrive in harsh environments as cold as

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1952-450: The bacterial phylum Planctomycetota has a membrane around the nucleoid and contains other membrane-bound cellular structures. However, further investigation revealed that Planctomycetota cells are not compartmentalized or nucleated and, like other bacterial membrane systems, are interconnected. Prokaryotic cells are usually much smaller than eukaryotic cells. Therefore, prokaryotes have a larger surface-area-to-volume ratio , giving them

2013-421: The biofilm—has led some to speculate that this may constitute a circulatory system and many researchers have started calling prokaryotic communities multicellular (for example ). Differential cell expression, collective behavior, signaling, programmed cell death , and (in some cases) discrete biological dispersal events all seem to point in this direction. However, these colonies are seldom if ever founded by

2074-440: The bodies of other organisms, including humans. Prokaryotes have high populations in the soil - including the rhizosphere and rhizosheath . Soil prokaryotes are still heavily undercharacterized despite their easy proximity to humans and their tremendous economic importance to agriculture . In 1977, Carl Woese proposed dividing prokaryotes into the Bacteria and Archaea (originally Eubacteria and Archaebacteria) because of

2135-709: 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

2196-404: 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

2257-427: 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

2318-454: The current set of prokaryotic species may have evolved from more complex eukaryotic ancestors through a process of simplification. Others have argued that the three domains of life arose simultaneously, from a set of varied cells that formed a single gene pool. This controversy was summarized in 2005: There is no consensus among biologists concerning the position of the eukaryotes in the overall scheme of cell evolution. Current opinions on

2379-622: 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

2440-406: 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

2501-484: 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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2562-530: The evolution and interrelationships of the three domains of life. The division between prokaryotes and eukaryotes reflects the existence of two very different levels of cellular organization; only eukaryotic cells have an enveloped nucleus that contains its chromosomal DNA , and other characteristic membrane-bound organelles including mitochondria. Distinctive types of prokaryotes include extremophiles and methanogens ; these are common in some extreme environments. The distinction between prokaryotes and eukaryotes

2623-646: 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

2684-480: The formation of the Earth's crust. Eukaryotes only appear in the fossil record later, and may have formed from endosymbiosis of multiple prokaryote ancestors. The oldest known fossil eukaryotes are about 1.7 billion years old. However, some genetic evidence suggests eukaryotes appeared as early as 3 billion years ago. While Earth is the only place in the universe where life is known to exist, some have suggested that there

2745-909: The four basic shapes of bacteria are: The archaeon Haloquadratum has flat square-shaped cells. Bacteria and archaea reproduce through asexual reproduction, usually by binary fission . Genetic exchange and recombination still occur, but this is a form of horizontal gene transfer and is not a replicative process, simply involving the transference of DNA between two cells, as in bacterial conjugation . DNA transfer between prokaryotic cells occurs in bacteria and archaea, although it has been mainly studied in bacteria. In bacteria, gene transfer occurs by three processes. These are (1) bacterial virus ( bacteriophage )-mediated transduction , (2) plasmid -mediated conjugation , and (3) natural transformation . Transduction of bacterial genes by bacteriophage appears to reflect an occasional error during intracellular assembly of virus particles, rather than an adaptation of

2806-432: The host bacteria. The transfer of bacterial DNA is under the control of the bacteriophage's genes rather than bacterial genes. Conjugation in the well-studied E. coli system is controlled by plasmid genes, and is an adaptation for distributing copies of a plasmid from one bacterial host to another. Infrequently during this process, a plasmid may integrate into the host bacterial chromosome, and subsequently transfer part of

2867-537: The host bacterial DNA to another bacterium. Plasmid mediated transfer of host bacterial DNA (conjugation) also appears to be an accidental process rather than a bacterial adaptation. Natural bacterial transformation involves the transfer of DNA from one bacterium to another through the intervening medium. Unlike transduction and conjugation, transformation is clearly a bacterial adaptation for DNA transfer, because it depends on numerous bacterial gene products that specifically interact to perform this complex process. For

2928-426: The idea that oligopeptides may have been built together with primordial nucleic acids at the same time, which also supports the concept of a ribocyte as LUCA. The feature of DNA as the material base of the genome might have then been adopted separately in bacteria and in archaea (and later eukaryote nuclei), presumably by help of some viruses (possibly retroviruses as they could reverse transcribe RNA to DNA). As

2989-424: The major differences in the structure and genetics between the two groups of organisms. Archaea were originally thought to be extremophiles, living only in inhospitable conditions such as extremes of temperature , pH , and radiation but have since been found in all types of habitats . The resulting arrangement of Eukaryota (also called "Eucarya"), Bacteria, and Archaea is called the three-domain system , replacing

3050-406: The medium (e.g., water) may flow easily. The microcolonies may join together above the substratum to form a continuous layer, closing the network of channels separating microcolonies. This structural complexity—combined with observations that oxygen limitation (a ubiquitous challenge for anything growing in size beyond the scale of diffusion) is at least partially eased by movement of medium throughout

3111-571: The mitochondria and chloroplasts. The genome in a prokaryote is held within a DNA/protein complex in the cytosol called the nucleoid , which lacks a nuclear envelope . The complex contains a single, cyclic, double-stranded molecule of stable chromosomal DNA, in contrast to the multiple linear, compact, highly organized chromosomes found in eukaryotic cells. In addition, many important genes of prokaryotes are stored in separate circular DNA structures called plasmids . Like eukaryotes, prokaryotes may partially duplicate genetic material, and can have

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3172-403: The nucleus, that eukaryotes arose without endosymbiosis, and that eukaryotes arose through a symbiotic event entailing a simultaneous endosymbiotic origin of the flagellum and the nucleus, in addition to many other models, which have been reviewed and summarized elsewhere. The oldest known fossilized prokaryotes were laid down approximately 3.5 billion years ago, only about 1 billion years after

3233-408: The origin and position of eukaryotes span a broad spectrum including the views that eukaryotes arose first in evolution and that prokaryotes descend from them, that eukaryotes arose contemporaneously with eubacteria and archaebacteria and hence represent a primary line of descent of equal age and rank as the prokaryotes, that eukaryotes arose through a symbiotic event entailing an endosymbiotic origin of

3294-873: The other distinct organelles that characterize the eukaryotic cell. It was once thought that prokaryotic cellular components were unenclosed within the cytoplasm except for an outer cell membrane , but bacterial microcompartments , which are thought to be quasi-organelles enclosed in protein shells (such as the encapsulin protein cages ), have been discovered, along with other prokaryotic organelles . While being unicellular, some prokaryotes, such as cyanobacteria , may form colonies held together by biofilms , and large colonies can create multilayered microbial mats . Others, such as myxobacteria , have multicellular stages in their life cycles . Prokaryotes are asexual , reproducing via binary fission without any fusion of gametes , although horizontal gene transfer may take place. Molecular studies have provided insight into

3355-449: 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

3416-417: 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

3477-498: 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

3538-427: The snow surface of Antarctica , studied in cryobiology , or as hot as undersea hydrothermal vents and land-based hot springs . Prokaryotes live in nearly all environments on Earth. Some archaea and bacteria are extremophiles , thriving in harsh conditions, such as high temperatures ( thermophiles ) or high salinity ( halophiles ). Many archaea grow as plankton in the oceans. Symbiotic prokaryotes live in or on

3599-426: The traditional two-empire system . According to the phylogenetic analysis of Hug (2016), the relationships could be the following: A widespread current model of the evolution of the first living organisms is that these were some form of prokaryotes, which may have evolved out of protocells , while the eukaryotes evolved later in the history of life. Some authors have questioned this conclusion, arguing that

3660-498: 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

3721-520: Was firmly established by the microbiologists Roger Stanier and C. B. van Niel in their 1962 paper The concept of a bacterium (though spelled procaryote and eucaryote there). That paper cites Édouard Chatton 's 1937 book Titres et Travaux Scientifiques for using those terms and recognizing the distinction. One reason for this classification was so that what was then often called blue-green algae (now called cyanobacteria ) would not be classified as plants but grouped with bacteria. Prokaryotes have

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