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Protein production

Protein production is the biotechnological process of generating a specific protein. It is typically achieved by the manipulation of gene expression in an organism such that it expresses large amounts of a recombinant gene. This includes the transcription of the recombinant DNA to messenger RNA (mRNA), the translation of mRNA into polypeptide chains, which are ultimately folded into functional proteins and may be targeted to specific subcellular or extracellular locations.

Protein production systems (also known as expression systems) are used in the life sciences, biotechnology, and medicine. Molecular biology research uses numerous proteins and enzymes, many of which are from expression systems; particularly DNA polymerase for PCR, reverse transcriptase for RNA analysis, restriction endonucleases for cloning, and to make proteins that are screened in drug discovery as biological targets or as potential drugs themselves. There are also significant applications for expression systems in industrial fermentation, notably the production of biopharmaceuticals such as human insulin to treat diabetes, and to manufacture enzymes.

Protein production systems
Commonly used protein production systems include those derived from bacteria, yeast,  baculovirus/insect, mammalian cells,  and more recently filamentous fungi such as Myceliophthora thermophila. When biopharmaceuticals are produced with one of these systems, process-related impurities termed host cell proteins also arrive in the final product in trace amounts.

Cell-based systems
The oldest and most widely used expression systems are cell-based and may be defined as the "combination of an expression vector, its cloned DNA, and the host for the vector that provide a context to allow foreign gene function in a host cell, that is, produce proteins at a high level". Overexpression is an abnormally and excessively high level of gene expression which produces a pronounced gene-related phenotype.

There are many ways to introduce foreign DNA to a cell for expression, and many different host cells may be used for expression — each expression system has distinct advantages and liabilities. Expression systems are normally referred to by the host and the DNA source or the delivery mechanism for the genetic material. For example, common hosts are bacteria (such as E. coli, B. subtilis), yeast (such as S. cerevisiae ) or eukaryotic cell lines. Common DNA sources and delivery mechanisms are viruses (such as baculovirus, retrovirus, adenovirus), plasmids, artificial chromosomes and bacteriophage (such as lambda). The best expression system depends on the gene involved, for example the Saccharomyces cerevisiae is often preferred for proteins that require significant posttranslational modification. Insect or mammal cell lines are used when human-like splicing of mRNA is required. Nonetheless, bacterial expression has the advantage of easily producing large amounts of protein, which is required for X-ray crystallography or nuclear magnetic resonance experiments for structure determination.

Because bacteria are prokaryotes, they are not equipped with the full enzymatic machinery to accomplish the required post-translational modifications or molecular folding. Hence, multi-domain eukaryotic proteins expressed in bacteria often are non-functional. Also, many proteins become insoluble as inclusion bodies that are difficult to recover without harsh denaturants and subsequent cumbersome protein-refolding.

To address these concerns, expressions systems using multiple eukaryotic cells were developed for applications requiring the proteins be conformed as in, or closer to eukaryotic organisms: cells of plants (i.e. tobacco), of insects or mammalians (i.e. bovines) are transfected with genes and cultured in suspension and even as tissues or whole organisms, to produce fully folded proteins. Mammalian in vivo expression systems have however low yield and other limitations (time-consuming, toxicity to host cells,..). To combine the high yield/productivity and scalable protein features of bacteria and yeast, and advanced epigenetic features of plants, insects and mammalians systems, other protein production systems are developed using unicellular eukaryotes (i.e. non-pathogenic ' Leishmania ' cells).

Escherichia coli
E. coli is one of the most widely used expression hosts, and DNA is normally introduced in a plasmid expression vector. The techniques for overexpression in E. coli are well developed and work by increasing the number of copies of the gene or increasing the binding strength of the promoter region so assisting transcription.

For example, a DNA sequence for a protein of interest could be cloned or subcloned into a high copy-number plasmid containing the lac (often LacUV5) promoter, which is then transformed into the bacterium E. coli. Addition of IPTG (a lactose analog) activates the lac promoter and causes the bacteria to express the protein of interest.

E. coli strain BL21 and BL21(DE3) are two strains commonly used for protein production. As members of the B lineage, they lack lon and OmpT proteases, protecting the produced proteins from degradation. The DE3 prophage found in BL21(DE3) provides T7 RNA polymerase (driven by the LacUV5 promoter), allowing for vectors with the T7 promoter to be used instead.

Corynebacterium
Non-pathogenic species of the gram-positive Corynebacterium are used for the commercial production of various amino acids. The C. glutamicum species is widely used for producing glutamate and lysine, components of human food, animal feed and pharmaceutical products.

Expression of functionally active human epidermal growth factor has been done in C. glutamicum, thus demonstrating a potential for industrial-scale production of human proteins. Expressed proteins can be targeted for secretion through either the general, secretory pathway (Sec) or the twin-arginine translocation pathway (Tat).

Unlike gram-negative bacteria, the gram-positive Corynebacterium lack lipopolysaccharides that function as antigenic endotoxins in humans.

Pseudomonas fluorescens
The non-pathogenic and gram-negative bacteria, Pseudomonas fluorescens, is used for high level production of recombinant proteins; commonly for the development bio-therapeutics and vaccines. P. fluorescens is a metabolically versatile organism, allowing for high throughput screening and rapid development of complex proteins. P. fluorescens is most well known for its ability to rapid and successfully produce high titers of active, soluble protein.

Yeasts
Expression systems using either S. cerevisiae or Pichia pastoris allow stable and lasting production of proteins that are processed similarly to mammalian cells, at high yield, in chemically defined media of proteins.

Filamentous fungi
Filamentous fungi, especially Aspergillus and Trichoderma, have long been used to produce diverse industrial enzymes from their own genomes ("native", "homologous") and from recombinant DNA ("heterologous").

More recently, Myceliophthora thermophila C1 has been developed into an expression platform for screening and production of native and heterologous proteins.The expression system C1 shows a low viscosity morphology in submerged culture, enabling the use of complex growth and production media. C1 also does not "hyperglycosylate" heterologous proteins, as Aspergillus and Trichoderma tend to do.

Baculovirus-infected cells
Baculovirus-infected insect cells (Sf9, Sf21, High Five strains) or mammalian cells (HeLa, HEK 293) allow production of glycosylated or membrane proteins that cannot be produced using fungal or bacterial systems. It is useful for production of proteins in high quantity. Genes are not expressed continuously because infected host cells eventually lyse and die during each infection cycle.

Non-lytic insect cell expression
Non-lytic insect cell expression is an alternative to the lytic baculovirus expression system. In non-lytic expression, vectors are transiently or stably transfected into the chromosomal DNA of insect cells for subsequent gene expression. This is followed by selection and screening of recombinant clones. The non-lytic system has been used to give higher protein yield and quicker expression of recombinant genes compared to baculovirus-infected cell expression. Cell lines used for this system include: Sf9, Sf21 from Spodoptera frugiperda cells, Hi-5 from Trichoplusia ni cells, and Schneider 2 cells and Schneider 3 cells from Drosophila melanogaster cells. With this system, cells do not lyse and several cultivation modes can be used. Additionally, protein production runs are reproducible. This system gives a homogeneous product. A drawback of this system is the requirement of an additional screening step for selecting viable clones.

Excavata
Leishmania tarentolae (cannot infect mammals) expression systems allow stable and lasting production of proteins at high yield, in chemically defined media. Produced proteins exhibit fully eukaryotic post-translational modifications, including glycosylation and disulfide bond formation.

Mammalian systems
The most common mammalian expression systems are Chinese Hamster ovary (CHO) and Human embryonic kidney (HEK) cells.
 * Chinese hamster ovary cell
 * Mouse myeloma lymphoblstoid (e.g. NS0 cell)
 * Fully Human
 * Human embryonic kidney cells (HEK-293)
 * Human embryonic retinal cells (Crucell's Per.C6)
 * Human amniocyte cells (Glycotope and CEVEC)

Cell-free systems
Cell-free production of proteins is performed in vitro using purified RNA polymerase, ribosomes, tRNA and ribonucleotides. These reagents may be produced by extraction from cells or from a cell-based expression system. Due to the low expression levels and high cost of cell-free systems, cell-based systems are more widely used.

title: Protein production


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Hermeto Pascoal

Hermeto Pascoal (born June 22, 1936) is a Brazilian composer and multi-instrumentalist. He was born in Lagoa da Canoa, Alagoas, Brazil. Pascoal is best known in Brazilian music for his orchestration and improvisation, as well as for being a record producer and contributor to many Brazilian and international albums.

Early life and career
Pascoal comes from Northeastern Brazil, in an area that lacked electricity at the time he was born. He learned the accordion from his father and practiced for hours indoors, as, being albino, he was incapable of working in the countryside with the rest of his family.

His career began in 1964 with appearances on several Brazilian recordings alongside relatively unknown groups. These albums and the musicians involved (Edu Lobo, Elis Regina, Cesar Camargo Mariano) established widely influential new directions in post-bossa nova Brazilian jazz.

In 1966, he played in the Sambrasa Trio, with Airto Moreira and Humberto Clayber; they released only one album, Em Som Maior. Then he joined Trio Novo (Airto Moreira, Heraldo do Monte, Theo de Barros) and in 1967 the group, renamed Quarteto Novo, released an album that launched the careers of Pascoal and Moreira. Pascoal would then go on to join the multi-faceted group Brazilian Octopus.

International fame
Pascoal initially caught the international public's attention with an appearance on Miles Davis's 1971 album Live-Evil, which featured him on three pieces, which he also composed. Davis allegedly called Pascoal "the most impressive musician in the world". Later collaborations involved fellow Brazilian musicians Airto Moreira and Flora Purim. From the late 1970s onward, he has mostly led his own groups, playing at many prestigious venues, such as the Montreux Jazz Festival in 1979. Other members of the group have included bassist Itibere Zwarg, pianist Jovino Santos-Neto and percussionists Nene, Pernambuco, and Zabelê.

Known as o Bruxo (the Sorcerer), Pascoal often makes music with unconventional objects such as teapots, children's toys, and animals, as well as keyboards, button accordions, melodica, saxophones, guitars, flutes, voices, various brass and folkloric instruments. He uses nature as a basis for his compositions, as in his Música da Lagoa, in which the musicians burble water and play flutes while immersed in a lagoon: a Brazilian television broadcast from 1999 showed him soloing at one point by singing into a cup with his mouth partially submerged in water. Folk music from rural Brazil is another important influence in his work.

Between 1996 and 1997, Pascoal worked on a book project called Calendário do Som, which contains a song for every day of the year, including February 29, so that everyone would have a song for their birthday.

He was married to Ilza da Silva, to whom he dedicated many compositions, from 1954 until her death in 2000. They had six children, Jorge, Fábio, Flávia, Fátima, Fabiula, and Flávio, and many grandchildren. Hermeto was later married to Aline Morena from 2003 until 2016, while living in Curitiba, Paraná, Brazil.

He has since returned to the Jabour neighborhood in Bangu, Rio de Janeiro, where he spends much of his time composing, rehearsing and hosting musicians from all over the world.

In 2019, his album Hermeto Pascoal e Sua Visão Original do Forró won the Latin Grammy Award for Best Portuguese Language Roots Album.

As leader or member

 * 1964: Conjunto Som 4 (with Conjunto Som 4)
 * 1966: Em Som Maior (with Sambrasa Trio)
 * 1967: Quarteto Novo (with Quarteto Novo)
 * 1969: Brazilian Octopus (with Brazilian Octopus)
 * 1970: Hermeto Pascoal (solo debut, reissued on CD as Brazilian Adventure) (featuring Googie Coppola)
 * 1973: A música livre de Hermeto Pascoal
 * 1977: Slaves Mass
 * 1977: Trindade
 * 1979: Zabumbê-bum-á
 * 1979: Ao vivo Montreux Jazz Festival
 * 1979: Nova história da Música Popular Brasileira (compilation)
 * 1980: Cérebro magnético
 * 1981: Planetário da Gávea
 * 1982: Hermeto Pascoal & Grupo (reissued by Westwind Germany on CD as The Legendary Improviser. The reissue appears to be a copy from vinyl.)
 * 1984: Lagoa da Canoa, Município de Arapiraca
 * 1986: Brasil Universo
 * 1987: Só não toca quem não quer
 * 1988: Hermeto solo: por diferentes caminhos
 * 1992: Festa dos deuses
 * 1993: Instrumental no CCBB (with Renato Borghetti)
 * 1998: Música!: o melhor da música de Hermeto Pascoal (compilation)
 * 1999: Eu e eles
 * 2002: Mundo verde esperança
 * 2006: Chimarrão com rapadura (with Aline Morena)
 * 2010: Bodas de Latão (with Aline Morena)
 * 2013: Hermeto Pascoal: The Monash Sessions
 * 2017: No Mundo dos Sons
 * 2017: Viajando com o som (recorded in 1976)
 * 2017: Natureza Universal
 * 2018: Made of Music
 * 2018: E sua visão original do forró

As contributor

 * 1956: Ritmos Alucinantes, by Clovis Pereira
 * 1959: Batucando no Morro, by Pernambuco do Pandeiro e seu regional
 * 1970: Natural Feelings, by Airto Moreira
 * 1970: Electric Byrd, by Donald Byrd
 * 1970: Sergio Mendes Presents Lobo, by Edu Lobo
 * 1970: It Could Only Happen with You, by Duke Pearson
 * 1970: Live-Evil, by Miles Davis
 * 1971: Cantiga de Longe, by Edu Lobo
 * 1971: Seeds on the Ground, by Airto Moreira
 * 1975: Di Melo, by Di Melo
 * 1976: Imyra, Tayra, Ipy, by Taiguara
 * 1976: Open Your Eyes You Can Fly, by Flora Purim
 * 1976 Goldenwings, by Opa
 * 1977: Orós, by Raimundo Fagner
 * 1978: Robertinho no passo, by Robertinho de Recife
 * 1979: Sivuca, by Sivuca
 * 1979: Live in Montreux, by Elis Regina
 * 1980: Stone Alliance, by Márcio Montarroyos
 * 1983: Cordas vivas, by Heraldo do Monte
 * 1984: Ponto do músicos, by Nenê
 * 1986: Balãozinho, by Eduardo Gudin
 * 1986: Cordas mágicas, by Heraldo do Monte
 * 1986: Pindorama, by Pau Brasil
 * 1987: Flávio Pantoja, by Flavio Pantoja
 * 1987: Dharana, by Dharana
 * 1996: Brasil Musical - Série Música Viva - Pau Brasil E Hermeto Pascoal|Brasil Musical - Série Música Viva (with Pau Brasil)
 * 1998: Maritmo, by Adriana Calcanhotto (on track "Canção por Acaso")
 * 2000: Oferenda, by Aleuda
 * 2000: Nação Nordestina, by Zé Ramalho (on track "Violando com Hermeto")
 * 2003: Serenata: The Music of Hermeto Pascoal, by Mike Marshall and Jovino Santos Neto
 * 2006: Roda Carioca, by Jovino Santos Neto
 * 2023: Beams, by Dan Costa (composer)

license: Creative Commons - Attribution Share-Alike - https://creativecommons.org/licenses/by-sa/4.0/

title: Hermeto Pascoal


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Talk:Squonk

Comment
Why is Jorge Luis Borges' Book of Imaginary Beings not mentioned in Jorge Luis Borges ? — Preceding unsigned comment added by MartinHarper (talk • contribs) 09:51, 10 January 2003 (UTC)

 * Because no one has mentioned it. But since this is a wiki, you can mention it yourself.  You certainly know more about than I do; I'd never heard of it before.  :-)  KQ —Preceding undated comment added 10:02, 10 January 2003 (UTC)
 * It's there now. -GTBacchus —Preceding undated comment added 23:15, 11 January 2003 (UTC)
 * I know I've heard of this thing before, but I definitely haven't ever read that book. I'm sure this is mentioned in a book I have read. It's slowly driving me insane though. So that's a plus. —Preceding unsigned comment added by 203.73.254.205 (talk) 14:53, 30 April 2008 (UTC)
 * I know I've heard of this thing before, but I definitely haven't ever read that book. I'm sure this is mentioned in a book I have read. It's slowly driving me insane though. So that's a plus. —Preceding unsigned comment added by 203.73.254.205 (talk) 14:53, 30 April 2008 (UTC)

Squank
Another cultural reference it well could be "Squank", 3rd trak from the ZZ Top's first album in 1971, about a monster living in the swamps. — Preceding unsigned comment added by 86.9.158.101 (talk) 14:19, 18 March 2006 (UTC)

Steely Dan song
I was surprised not to see the reference to "Have you ever seen a Squonk's Tears well look at mine. A line from a Steely Dan Song - Any Major Dude (will tell you) on their Pretzel Logic Album of 1974. Anothersquonk (talk) 16:10, 17 March 2016 (UTC)

Black walnuts
Does anyone know if there is truth to the rumor that they can create canopy gaps by attempting suicide by eating the bark of black walnuts? Harry Meanwell (talk) 14:35, 13 September 2016 (UTC)

 * that could be because black walnuts have a chemical that harms plants. But of course the squank is 110% real!Realfakebezalbob (talk) 00:30, 13 October 2022 (UTC)

Samuel Johnson
See author Samuel Johnson’s famous 18th-century dictionary, where squonk is a very tearful animal who stands in a puddle of its own tears. — Preceding unsigned comment added by Wicketdg1 (talk • contribs) 21:06, 8 April 2020 (UTC)

Random Older Mention
I found a mention of a squonk from a 1878 settler's diary in Nebraska but they don't go into detail so I assume it's unrelated? Bit weird though as I'm not sure what else they could be referring to!

Ref is - "The Settler's Calendar, Prairie Schooner, Vol. 41, No. 2, NEBRASKA CENTENNIAL ISSUE (SUMMER 1967), pp. 133-136" PanydThe muffin is not subtle 11:43, 11 August 2020 (UTC)

Genesis: A Trick of the Tail
Squonk was also a song on the early Genesis album A Trick of the Tail. 2601:248:400:C780:94C5:6AEE:EBAE:F13 (talk) 20:56, 27 November 2022 (UTC)

title: Talk:Squonk


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Santalaceae

The Santalaceae, sandalwoods, are a widely distributed family of flowering plants (including small trees, shrubs, perennial herbs, and epiphytic climbers ) which, like other members of Santalales, are partially parasitic on other plants. Its flowers are bisexual or, by abortion ("flower drop"), unisexual. Modern treatments of the Santalaceae include the family Viscaceae (mistletoes), previously considered distinct.

The APG II system of 2003 recognises the family and assigns it to the order Santalales in the clade core eudicots. However, the circumscription by APG is much wider than accepted by previous classifications, including the plants earlier treated in families Eremolepidaceae and Viscaceae. It includes about 1,000 species in 43 genera. Many have reported traditional and cultural uses, including as medicine.

Genera
43 genera are accepted.

 * Acanthosyris (Eichler) Griseb.
 * Amphorogyne Stauffer & Hürl.
 * Antidaphne Poepp. & Endl. (previously Eremolepidaceae)
 * Arceuthobium M.Bieb. (previously Viscaceae)
 * Buckleya Torr.
 * Cervantesia Ruiz & Pav.
 * Choretrum R.Br.
 * Comandra Nutt.
 * Daenikera Hürl. & Stauffer
 * Dendromyza Danser
 * Dendrophthora Eichler (previously Viscaceae)
 * Dendrotrophe Miq.
 * Elaphanthera N.Hallé
 * Eubrachion Hook.f. (previously Eremolepidaceae)
 * Exocarpos Labill.
 * Geocaulon Fernald
 * Ginalloa Korth. (previously Viscaceae)
 * Henslowia Blume
 * Hylomyza Danser
 * Jodina Hook. & Arn. ex Meisn.
 * Korthalsella Tiegh.
 * Lacomucinaea Nickrent & M.A.García
 * Lepidoceras Hook.f. (previously Eremolepidaceae)
 * Leptomeria R.Br.
 * Mida R.Cunn. ex A.Cunn.
 * Myoschilos Ruiz & Pav.
 * Nanodea Banks ex C.F.Gaertn.
 * Nestronia Raf.
 * Notothixos Oliv. (previously Viscaceae)
 * Okoubaka Pellegr. & Normand
 * Omphacomeria A.DC.
 * Osyridicarpos A.DC.
 * Osyris L.
 * Phacellaria Benth.
 * Phoradendron Nutt. (previously Viscaceae)
 * Pilgerina Z.S.Rogers, Nickrent & Malécot
 * Pyrularia Michx.
 * Rhoiacarpos A.DC.
 * Santalum L.
 * Scleropyrum Arn.
 * Staufferia Z.S.Rogers, Nickrent & Malécot
 * Thesium L.
 * Viscum L. (previously Viscaceae)

Formerly placed here

 * Anthobolus R.Br. – to Opiliaceae
 * Arjona - to Schoepfiaceae
 * Quinchamalium - to Schoepfiaceae

title: Santalaceae


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Transcription (biology)

Transcription is the process of copying a segment of DNA into RNA. The segments of DNA transcribed into RNA molecules that can encode proteins produce messenger RNA (mRNA). Other segments of DNA are transcribed into RNA molecules called non-coding RNAs (ncRNAs).

Both DNA and RNA are nucleic acids, which use base pairs of nucleotides as a complementary language. During transcription, a DNA sequence is read by an RNA polymerase, which produces a complementary, antiparallel RNA strand called a primary transcript.

In virology, the term transcription may also be used when referring to mRNA synthesis from an RNA molecule (i.e., equivalent to RNA replication). For instance, the genome of a negative-sense single-stranded RNA (ssRNA -) virus may be a template for a positive-sense single-stranded RNA (ssRNA +). This is because the positive-sense strand contains the sequence information needed to translate the viral proteins needed for viral replication. This process is catalyzed by a viral RNA replicase.

Background
A DNA transcription unit encoding for a protein may contain both a coding sequence, which will be translated into the protein, and regulatory sequences, which direct and regulate the synthesis of that protein. The regulatory sequence before (upstream from) the coding sequence is called the five prime untranslated regions (5'UTR); the sequence after (downstream from) the coding sequence is called the three prime untranslated regions (3'UTR).

As opposed to DNA replication, transcription results in an RNA complement that includes the nucleotide uracil (U) in all instances where thymine (T) would have occurred in a DNA complement.

Only one of the two DNA strands serves as a template for transcription. The antisense strand of DNA is read by RNA polymerase from the 3' end to the 5' end during transcription (3' → 5'). The complementary RNA is created in the opposite direction, in the 5' → 3' direction, matching the sequence of the sense strand except switching uracil for thymine. This directionality is because RNA polymerase can only add nucleotides to the 3' end of the growing mRNA chain. This use of only the 3' → 5' DNA strand eliminates the need for the Okazaki fragments that are seen in DNA replication. This also removes the need for an RNA primer to initiate RNA synthesis, as is the case in DNA replication.

The non-template (sense) strand of DNA is called the coding strand, because its sequence is the same as the newly created RNA transcript (except for the substitution of uracil for thymine). This is the strand that is used by convention when presenting a DNA sequence.

Transcription has some proofreading mechanisms, but they are fewer and less effective than the controls for copying DNA. As a result, transcription has a lower copying fidelity than DNA replication.

Major steps
Transcription is divided into initiation, promoter escape, elongation, and termination.

Enhancers, transcription factors, Mediator complex, and DNA loops in mammalian transcription
Setting up for transcription in mammals is regulated by many cis-regulatory elements, including core promoter and promoter-proximal elements that are located near the transcription start sites of genes. Core promoters combined with general transcription factors are sufficient to direct transcription initiation, but generally have low basal activity. Other important cis-regulatory modules are localized in DNA regions that are distant from the transcription start sites. These include enhancers, silencers, insulators and tethering elements. Among this constellation of elements, enhancers and their associated transcription factors have a leading role in the initiation of gene transcription. An enhancer localized in a DNA region distant from the promoter of a gene can have a very large effect on gene transcription, with some genes undergoing up to 100-fold increased transcription due to an activated enhancer.

Enhancers are regions of the genome that are major gene-regulatory elements. Enhancers control cell-type-specific gene transcription programs, most often by looping through long distances to come in physical proximity with the promoters of their target genes. While there are hundreds of thousands of enhancer DNA regions, for a particular type of tissue only specific enhancers are brought into proximity with the promoters that they regulate. In a study of brain cortical neurons, 24,937 loops were found, bringing enhancers to their target promoters. Multiple enhancers, each often at tens or hundred of thousands of nucleotides distant from their target genes, loop to their target gene promoters and can coordinate with each other to control transcription of their common target gene.

The schematic illustration in this section shows an enhancer looping around to come into close physical proximity with the promoter of a target gene. The loop is stabilized by a dimer of a connector protein (e.g. dimer of CTCF or YY1), with one member of the dimer anchored to its binding motif on the enhancer and the other member anchored to its binding motif on the promoter (represented by the red zigzags in the illustration). Several cell function specific transcription factors (there are about 1,600 transcription factors in a human cell ) generally bind to specific motifs on an enhancer and a small combination of these enhancer-bound transcription factors, when brought close to a promoter by a DNA loop, govern level of transcription of the target gene. Mediator (a complex usually consisting of about 26 proteins in an interacting structure) communicates regulatory signals from enhancer DNA-bound transcription factors directly to the RNA polymerase II (pol II) enzyme bound to the promoter.

Enhancers, when active, are generally transcribed from both strands of DNA with RNA polymerases acting in two different directions, producing two enhancer RNAs (eRNAs) as illustrated in the Figure. An inactive enhancer may be bound by an inactive transcription factor. Phosphorylation of the transcription factor may activate it and that activated transcription factor may then activate the enhancer to which it is bound (see small red star representing phosphorylation of transcription factor bound to enhancer in the illustration). An activated enhancer begins transcription of its RNA before activating transcription of messenger RNA from its target gene.

CpG island methylation and demethylation

Transcription regulation at about 60% of promoters is also controlled by methylation of cytosines within CpG dinucleotides (where 5' cytosine is followed by 3' guanine or CpG sites). 5-methylcytosine (5-mC) is a methylated form of the DNA base cytosine (see Figure). 5-mC is an epigenetic marker found predominantly within CpG sites. About 28 million CpG dinucleotides occur in the human genome. In most tissues of mammals, on average, 70% to 80% of CpG cytosines are methylated (forming 5-methylCpG or 5-mCpG). However, unmethylated cytosines within 5'cytosine-guanine 3' sequences often occur in groups, called CpG islands, at active promoters. About 60% of promoter sequences have a CpG island while only about 6% of enhancer sequences have a CpG island. CpG islands constitute regulatory sequences, since if CpG islands are methylated in the promoter of a gene this can reduce or silence gene transcription.

DNA methylation regulates gene transcription through interaction with methyl binding domain (MBD) proteins, such as MeCP2, MBD1 and MBD2. These MBD proteins bind most strongly to highly methylated CpG islands. These MBD proteins have both a methyl-CpG-binding domain as well as a transcription repression domain. They bind to methylated DNA and guide or direct protein complexes with chromatin remodeling and/or histone modifying activity to methylated CpG islands. MBD proteins generally repress local chromatin such as by catalyzing the introduction of repressive histone marks, or creating an overall repressive chromatin environment through nucleosome remodeling and chromatin reorganization. As noted in the previous section, transcription factors are proteins that bind to specific DNA sequences in order to regulate the expression of a gene. The binding sequence for a transcription factor in DNA is usually about 10 or 11 nucleotides long. As summarized in 2009, Vaquerizas et al. indicated there are approximately 1,400 different transcription factors encoded in the human genome by genes that constitute about 6% of all human protein encoding genes. About 94% of transcription factor binding sites (TFBSs) that are associated with signal-responsive genes occur in enhancers while only about 6% of such TFBSs occur in promoters.

EGR1 protein is a particular transcription factor that is important for regulation of methylation of CpG islands. An EGR1 transcription factor binding site is frequently located in enhancer or promoter sequences. There are about 12,000 binding sites for EGR1 in the mammalian genome and about half of EGR1 binding sites are located in promoters and half in enhancers. The binding of EGR1 to its target DNA binding site is insensitive to cytosine methylation in the DNA.

While only small amounts of EGR1 transcription factor protein are detectable in cells that are un-stimulated, translation of the EGR1 gene into protein at one hour after stimulation is drastically elevated. Production of EGR1 transcription factor proteins, in various types of cells, can be stimulated by growth factors, neurotransmitters, hormones, stress and injury. In the brain, when neurons are activated, EGR1 proteins are up-regulated and they bind to (recruit) the pre-existing TET1 enzymes that are produced in high amounts in neurons. TET enzymes can catalyse demethylation of 5-methylcytosine. When EGR1 transcription factors bring TET1 enzymes to EGR1 binding sites in promoters, the TET enzymes can demethylate the methylated CpG islands at those promoters. Upon demethylation, these promoters can then initiate transcription of their target genes. Hundreds of genes in neurons are differentially expressed after neuron activation through EGR1 recruitment of TET1 to methylated regulatory sequences in their promoters.

The methylation of promoters is also altered in response to signals. The three mammalian DNA methyltransferasess (DNMT1, DNMT3A, and DNMT3B) catalyze the addition of methyl groups to cytosines in DNA. While DNMT1 is a maintenance methyltransferase, DNMT3A and DNMT3B can carry out new methylations. There are also two splice protein isoforms produced from the DNMT3A gene: DNA methyltransferase proteins DNMT3A1 and DNMT3A2.

The splice isoform DNMT3A2 behaves like the product of a classical immediate-early gene and, for instance, it is robustly and transiently produced after neuronal activation. Where the DNA methyltransferase isoform DNMT3A2 binds and adds methyl groups to cytosines appears to be determined by histone post translational modifications.

On the other hand, neural activation causes degradation of DNMT3A1 accompanied by reduced methylation of at least one evaluated targeted promoter.

Initiation
Transcription begins with the RNA polymerase and one or more general transcription factors binding to a DNA promoter sequence to form an RNA polymerase-promoter closed complex. In the closed complex, the promoter DNA is still fully double-stranded.

RNA polymerase, assisted by one or more general transcription factors, then unwinds approximately 14 base pairs of DNA to form an RNA polymerase-promoter open complex. In the open complex, the promoter DNA is partly unwound and single-stranded. The exposed, single-stranded DNA is referred to as the "transcription bubble".

RNA polymerase, assisted by one or more general transcription factors, then selects a transcription start site in the transcription bubble, binds to an initiating NTP and an extending NTP (or a short RNA primer and an extending NTP) complementary to the transcription start site sequence, and catalyzes bond formation to yield an initial RNA product.

In bacteria, RNA polymerase holoenzyme consists of five subunits: 2 α subunits, 1 β subunit, 1 β' subunit, and 1 ω subunit. In bacteria, there is one general RNA transcription factor known as a sigma factor. RNA polymerase core enzyme binds to the bacterial general transcription (sigma) factor to form RNA polymerase holoenzyme and then binds to a promoter. (RNA polymerase is called a holoenzyme when sigma subunit is attached to the core enzyme which is consist of 2 α subunits, 1 β subunit, 1 β' subunit only). Unlike eukaryotes, the initiating nucleotide of nascent bacterial mRNA is not capped with a modified guanine nucleotide. The initiating nucleotide of bacterial transcripts bears a 5′ triphosphate (5′-PPP), which can be used for genome-wide mapping of transcription initiation sites.

In archaea and eukaryotes, RNA polymerase contains subunits homologous to each of the five RNA polymerase subunits in bacteria and also contains additional subunits. In archaea and eukaryotes, the functions of the bacterial general transcription factor sigma are performed by multiple general transcription factors that work together. In archaea, there are three general transcription factors: TBP, TFB, and TFE. In eukaryotes, in RNA polymerase II-dependent transcription, there are six general transcription factors: TFIIA, TFIIB (an ortholog of archaeal TFB), TFIID (a multisubunit factor in which the key subunit, TBP, is an ortholog of archaeal TBP), TFIIE (an ortholog of archaeal TFE), TFIIF, and TFIIH. The TFIID is the first component to bind to DNA due to binding of TBP, while TFIIH is the last component to be recruited. In archaea and eukaryotes, the RNA polymerase-promoter closed complex is usually referred to as the "preinitiation complex".

Transcription initiation is regulated by additional proteins, known as activators and repressors, and, in some cases, associated coactivators or corepressors, which modulate formation and function of the transcription initiation complex.

Promoter escape
After the first bond is synthesized, the RNA polymerase must escape the promoter. During this time there is a tendency to release the RNA transcript and produce truncated transcripts. This is called abortive initiation, and is common for both eukaryotes and prokaryotes. Abortive initiation continues to occur until an RNA product of a threshold length of approximately 10 nucleotides is synthesized, at which point promoter escape occurs and a transcription elongation complex is formed.

Mechanistically, promoter escape occurs through DNA scrunching, providing the energy needed to break interactions between RNA polymerase holoenzyme and the promoter.

In bacteria, it was historically thought that the sigma factor is definitely released after promoter clearance occurs. This theory had been known as the obligate release model. However, later data showed that upon and following promoter clearance, the sigma factor is released according to a stochastic model known as the stochastic release model.

In eukaryotes, at an RNA polymerase II-dependent promoter, upon promoter clearance, TFIIH phosphorylates serine 5 on the carboxy terminal domain of RNA polymerase II, leading to the recruitment of capping enzyme (CE). The exact mechanism of how CE induces promoter clearance in eukaryotes is not yet known.

Elongation

One strand of the DNA, the template strand (or noncoding strand), is used as a template for RNA synthesis. As transcription proceeds, RNA polymerase traverses the template strand and uses base pairing complementarity with the DNA template to create an RNA copy (which elongates during the traversal). Although RNA polymerase traverses the template strand from 3' → 5', the coding (non-template) strand and newly formed RNA can also be used as reference points, so transcription can be described as occurring 5' → 3'. This produces an RNA molecule from 5' → 3', an exact copy of the coding strand (except that thymines are replaced with uracils, and the nucleotides are composed of a ribose (5-carbon) sugar whereas DNA has deoxyribose (one fewer oxygen atom) in its sugar-phosphate backbone).

mRNA transcription can involve multiple RNA polymerases on a single DNA template and multiple rounds of transcription (amplification of particular mRNA), so many mRNA molecules can be rapidly produced from a single copy of a gene. The characteristic elongation rates in prokaryotes and eukaryotes are about 10–100 nts/sec. In eukaryotes, however, nucleosomes act as major barriers to transcribing polymerases during transcription elongation. In these organisms, the pausing induced by nucleosomes can be regulated by transcription elongation factors such as TFIIS.

Elongation also involves a proofreading mechanism that can replace incorrectly incorporated bases. In eukaryotes, this may correspond with short pauses during transcription that allow appropriate RNA editing factors to bind. These pauses may be intrinsic to the RNA polymerase or due to chromatin structure.

Double-strand breaks in actively transcribed regions of DNA are repaired by homologous recombination during the S and G2 phases of the cell cycle. Since transcription enhances the accessibility of DNA to exogenous chemicals and internal metabolites that can cause recombinogenic lesions, homologous recombination of a particular DNA sequence may be strongly stimulated by transcription.

Termination
Bacteria use two different strategies for transcription termination – Rho-independent termination and Rho-dependent termination. In Rho-independent transcription termination, RNA transcription stops when the newly synthesized RNA molecule forms a G-C-rich hairpin loop followed by a run of Us. When the hairpin forms, the mechanical stress breaks the weak rU-dA bonds, now filling the DNA–RNA hybrid. This pulls the poly-U transcript out of the active site of the RNA polymerase, terminating transcription. In Rho-dependent termination, Rho, a protein factor, destabilizes the interaction between the template and the mRNA, thus releasing the newly synthesized mRNA from the elongation complex.

Transcription termination in eukaryotes is less well understood than in bacteria, but involves cleavage of the new transcript followed by template-independent addition of adenines at its new 3' end, in a process called polyadenylation.

Beyond termination by a terminator sequences (which is a part of a gene), transcription may also need to be terminated when it encounters conditions such as DNA damage or an active replication fork. In bacteria, the Mfd ATPase can remove a RNA polymerase stalled at a lesion by prying open its clamp. It also recruits nucleotide excision repair machinery to repair the lesion. Mfd is proposed to also resolve conflicts between DNA replication and transcription. In eukayrotes, ATPase TTF2 helps to suppress the action of RNAP I and II during mitosis, preventing errors in chromosomal segregation. In archaea, the Eta ATPase is proposed to play a similar role.

Role of RNA polymerase in post-transcriptional changes in RNA
RNA polymerase plays a very crucial role in all steps including post-transcriptional changes in RNA. As shown in the image in the right it is evident that the CTD (C Terminal Domain) is a tail that changes its shape; this tail will be used as a carrier of splicing, capping and polyadenylation, as shown in the image on the left.

Inhibitors
Transcription inhibitors can be used as antibiotics against, for example, pathogenic bacteria (antibacterials) and fungi (antifungals). An example of such an antibacterial is rifampicin, which inhibits bacterial transcription of DNA into mRNA by inhibiting DNA-dependent RNA polymerase by binding its beta-subunit, while 8-hydroxyquinoline is an antifungal transcription inhibitor. The effects of histone methylation may also work to inhibit the action of transcription. Potent, bioactive natural products like triptolide that inhibit mammalian transcription via inhibition of the XPB subunit of the general transcription factor TFIIH has been recently reported as a glucose conjugate for targeting hypoxic cancer cells with increased glucose transporter production.

Endogenous inhibitors
In vertebrates, the majority of gene promoters contain a CpG island with numerous CpG sites. When many of a gene's promoter CpG sites are methylated the gene becomes inhibited (silenced). Colorectal cancers typically have 3 to 6 driver mutations and 33 to 66 hitchhiker or passenger mutations. However, transcriptional inhibition (silencing) may be of more importance than mutation in causing progression to cancer. For example, in colorectal cancers about 600 to 800 genes are transcriptionally inhibited by CpG island methylation (see regulation of transcription in cancer). Transcriptional repression in cancer can also occur by other epigenetic mechanisms, such as altered production of microRNAs. In breast cancer, transcriptional repression of BRCA1 may occur more frequently by over-produced microRNA-182 than by hypermethylation of the BRCA1 promoter (see Low expression of BRCA1 in breast and ovarian cancers).

Transcription factories
Active transcription units are clustered in the nucleus, in discrete sites called transcription factories or euchromatin. Such sites can be visualized by allowing engaged polymerases to extend their transcripts in tagged precursors (Br-UTP or Br-U) and immuno-labeling the tagged nascent RNA. Transcription factories can also be localized using fluorescence in situ hybridization or marked by antibodies directed against polymerases. There are ~10,000 factories in the nucleoplasm of a HeLa cell, among which are ~8,000 polymerase II factories and ~2,000 polymerase III factories. Each polymerase II factory contains ~8 polymerases. As most active transcription units are associated with only one polymerase, each factory usually contains ~8 different transcription units. These units might be associated through promoters and/or enhancers, with loops forming a "cloud" around the factor.

History
A molecule that allows the genetic material to be realized as a protein was first hypothesized by François Jacob and Jacques Monod. Severo Ochoa won a Nobel Prize in Physiology or Medicine in 1959 for developing a process for synthesizing RNA in vitro with polynucleotide phosphorylase, which was useful for cracking the genetic code. RNA synthesis by RNA polymerase was established in vitro by several laboratories by 1965; however, the RNA synthesized by these enzymes had properties that suggested the existence of an additional factor needed to terminate transcription correctly.

Roger D. Kornberg won the 2006 Nobel Prize in Chemistry "for his studies of the molecular basis of eukaryotic transcription".

Measuring and detecting

Transcription can be measured and detected in a variety of ways:
 * G-Less Cassette transcription assay: measures promoter strength
 * Run-off transcription assay: identifies transcription start sites (TSS)
 * Nuclear run-on assay: measures the relative abundance of newly formed transcripts
 * KAS-seq: measures single-stranded DNA generated by RNA polymerases; can work with 1,000 cells.
 * RNase protection assay and ChIP-Chip of RNAP: detect active transcription sites
 * RT-PCR: measures the absolute abundance of total or nuclear RNA levels, which may however differ from transcription rates
 * DNA microarrays: measures the relative abundance of the global total or nuclear RNA levels; however, these may differ from transcription rates
 * In situ hybridization: detects the presence of a transcript
 * MS2 tagging: by incorporating RNA stem loops, such as MS2, into a gene, these become incorporated into newly synthesized RNA. The stem loops can then be detected using a fusion of GFP and the MS2 coat protein, which has a high affinity, sequence-specific interaction with the MS2 stem loops. The recruitment of GFP to the site of transcription is visualized as a single fluorescent spot. This new approach has revealed that transcription occurs in discontinuous bursts, or pulses (see Transcriptional bursting). With the notable exception of in situ techniques, most other methods provide cell population averages, and are not capable of detecting this fundamental property of genes.
 * Northern blot: the traditional method, and until the advent of RNA-Seq, the most quantitative
 * RNA-Seq: applies next-generation sequencing techniques to sequence whole transcriptomes, which allows the measurement of relative abundance of RNA, as well as the detection of additional variations such as fusion genes, post-transcriptional edits and novel splice sites
 * Single cell RNA-Seq: amplifies and reads partial transcriptomes from isolated cells, allowing for detailed analyses of RNA in tissues, embryos, and cancers

Reverse transcription

Some viruses (such as HIV, the cause of AIDS), have the ability to transcribe RNA into DNA. HIV has an RNA genome that is reverse transcribed into DNA. The resulting DNA can be merged with the DNA genome of the host cell. The main enzyme responsible for synthesis of DNA from an RNA template is called reverse transcriptase.

In the case of HIV, reverse transcriptase is responsible for synthesizing a complementary DNA strand (cDNA) to the viral RNA genome. The enzyme ribonuclease H then digests the RNA strand, and reverse transcriptase synthesises a complementary strand of DNA to form a double helix DNA structure (cDNA). The cDNA is integrated into the host cell's genome by the enzyme integrase, which causes the host cell to generate viral proteins that reassemble into new viral particles. In HIV, subsequent to this, the host cell undergoes programmed cell death, or apoptosis, of T cells. However, in other retroviruses, the host cell remains intact as the virus buds out of the cell.

Some eukaryotic cells contain an enzyme with reverse transcription activity called telomerase. Telomerase carries an RNA template from which it synthesizes a telomere, a repeating sequence of DNA, to the end of linear chromosomes. It is important because every time a linear chromosome is duplicated, it is shortened. With the telomere at the ends of chromosomes, the shortening eliminates some of the non-essential, repeated sequence, rather than the protein-encoding DNA sequence farther away from the chromosome end.

Telomerase is often activated in cancer cells to enable cancer cells to duplicate their genomes indefinitely without losing important protein-coding DNA sequence. Activation of telomerase could be part of the process that allows cancer cells to become immortal. The immortalizing factor of cancer via telomere lengthening due to telomerase has been proven to occur in 90% of all carcinogenic tumors in vivo with the remaining 10% using an alternative telomere maintenance route called ALT or Alternative Lengthening of Telomeres.

title: Transcription (biology)


text:
Talk:Transcription (biology)

Wiki Education Foundation-supported course assignment
This article was the subject of a Wiki Education Foundation-supported course assignment, between 9 January 2020 and 24 April 2020. Further details are available on the course page. Student editor(s): Charlov21. Peer reviewers: Magdala5.

Above undated message substituted from Template:Dashboard.wikiedu.org assignment by PrimeBOT (talk) 11:37, 17 January 2022 (UTC)

Lead section too long
The lead seems too long, and contains quite a few inaccuracies. A good lead should be brief and refer to the major sections of the article. This one is long and often too detailed. Some of the inaccuracies:
 * ...which produces a complementary, antiparallel RNA strand: RNA is mostly single-stranded, so it would be better to say that the RNA sequence is reverse complementary to the template strand of the DNA
 * Both RNA and DNA are nucleic acids, which use base pairs of nucleotides ...: Again, RNA usually does not contain base pairs. In double-stranded DNA, the information is stored twice (redundantly in the two strands), in RNA it is present once in a single strand.
 * Transcription proceeds in 5 or 6 steps, each moving like a wave along the DNA: This is a list that used to make sense, referring to steps in transcription elongation. In the current form, point 6 is the odd one out, certainly not moving like a wave along the DNA.
 * The other DNA strand is called the coding (lagging) strand...: One strand is called the template strand, and the other the non-template strand (or coding strand because its sequence, apart from the U vs T issue, is identical to that of the RNA). Why would it be called the lagging strand, though? This makes sense for replication, but not for transcription.

My suggestion would be to remove most of the material from the lead, especially the list of "5 or 6" steps. If the lead had brief explanations of DNA, RNA, RNA polymerase, the biological role of RNA, the steps of transcription and the importance of transcription regulation for gene expression, it could be concise and less riddled with errors or misleading statements.Theislikerice (talk) 04:23, 17 March 2013 (UTC)

Central Dogma
The "see also" section has a misstatement of the central dogma (as stated by Crick). In papers in both 1958 and 1970 he emphasized that it was "once information has got into a protein it can't get back out." The idea that we have DNA->RNA->protein and never the other way around seems to be due to Watson's misunderstanding of Crick's central dogma. So either don't call it Crick's central dogma or change it to what Crick meant when he said "the central dogma." —Preceding unsigned comment added by 67.180.16.146 (talk) 00:06, 22 May 2011 (UTC)

2008 Major upgrade?
New Article Hi i´m Lukas Müller and i´m coming from Köln in Germany. i´m Studying biology at the Heidelberg-University, one of the best universities in germany and i want to ask, if we can make the site "Transcription" new so we can get a perfect article.
 * Please see the page section called, below, for discussion of major changes. --JWSchmidt (talk) 21:18, 3 January 2008 (UTC)

Harvard Papers?
"By the late 1960s several papers that came out of the Harvard University Biological Laboratories established the basic mechanics of gene expression in bacteria."

This information is not related to this article in context and will be deleted. If you're going to include this you might as well include all the articles on pub med because the "basic mechanics of gene expression in bacteria" is a communitative effort not just exclusivly by faculty in Harvard. Although I presume they had made their contributions as well. It's just not correct to include one place that contributed to gene expression, so I assume the person who included this is maybe from the school itself.

Reverse transcriptase
According to the current article, transcription refers solely to DNA>RNA transcription, and even goes to far as to link to Crick's central dogma (DNA is transcribed to RNA which is translated to polypeptides, never the other way around). Someone should make it clear from the start that transcription, while usually used to refer to DNA>RNA transcription, might also refer to RNA>DNA transcription. -- Ec5618 23:3'''9, 4 December 2005 (UTC)

I added a beginning of informations plus a little scheme I made earlier ... I think it should be in another page, but I don't know how to do ;-P. Hope someone will ! -- Totophe64

Eukaryotic Transcription
Edit 4/2/06 explanation Most eukaryotes have 4 RNA polymerases, plants have 5. Although RNAP is a commonly used abbreviation for bacterial and bacteriophage RNA polymerases, it is almost never used for eukaryotic RNA polymerases. I have substituted Pol I, II and III. (This will also need to be done in the subsections.) I have also included Pol A, B, and C not only in recognition of the work of Pierre Chambon and Andre Sentenac (vs. the nomenclature of William Rutter and Robert Roeder), but at some point the gene names (eg., RPA1, RPB1, RPC1) will come up as this section expands, and this edit will help clarify this nomenclature. -opus118

Organization and additional items needed "Perhaps one way of organizing this section is to group the different RNA polymerases: Archaea, bacteria, chloroplast, and eukaryotic nuclear RNA polymerases form one related group.  Bacteriophage T7-like and mitochondrial RNA polymerases form a second, DNA polymerase I-derived group (does reverse transcriptase fall into this category?), and then a diverse mixture of bacteriophage and viral RNA polymerases that do not fall within the above two categories.  There should be some emphasis on common mechanisms and related subunits.  For example: 1) the β'βα2ω subunits of bacteral RNA polymerase have sequence-related homologs in chloroplast RNA polymerase, archaeal RNA polymerase and Pol I, II and III. 2) The 12 (?) subunits of archaeal RNA polymerase have homologs in Pol I, II and III (with Pol III containing 5 additional unique subunits); 3) TBP is a transcription factor for archaeal, Pol I, Pol II, and Pol III transcription. 4) A TFIIB-related factor is common for archaeal, Pol II and pol III transcription. 5)TFIIH functions for Pol I and II transcription. 6) Promoter opening occurs with an upstream to downstream polarity with all polymerases. 7) Stem-loop structures at sites of termination are common (with the possible exception of Pol III). 8) The mechanism of catalysis for RNA chain elongation is identical (the active site region is highly conserved). 9) bacterial GreA/GreB, TFIIS and Rpc11 all stimulate a 3' to 5' ribonucleolytic activity at the RNA polymerase active site by inserting a finger-like structure in the nucleotide substrate feeding pore of RNA polymerase to place a pair of acidic residues near the catalytic site that chelates an additional Mg2+ ion for hydrolysis. It is the polymerase-unique properties that might be best dealt with in subsections. -opus18

Termination

 * Garbeled section needs Editing. Something missing or duplicated etc in the "sentence(s)":
 * in the DNA template. or where a GC-rich inverted
 * 4 A residues. the inverted repeat forms
 * dissociate from the DNA template. where the -35 region

*Rho-dependent termination uses a termination factor called ρ factor(rho factor) to stop RNA synthesis at specific sites. This protein binds and runs along the mRNA towards the RNAP. When ρ-factor reaches the RNAP, it causes RNAP to dissociate from the DNA, terminating transcription.
 * See the following surround section.

Other termination mechanisms include where RNAP comes across a region with repetitious thymidine residues in the DNA template. or where a GC-rich inverted repeat followed by 4 A residues. the inverted repeat forms a stable stem loop structure in the Rna, which causes the RNA to dissociate from the DNA template.

where the -35 region and the -10 ("Pribnow box") region comprise the basic prokaryotic promoter, and |T| stands for the terminator. The DNA on the template strand between the +1 site and the terminator is transcribed into RNA, which is then translated into protein. DLH 17:37, 10 September 2006 (UTC)

category
Surely this article should be under Molecular Biology? answer: WHY? Where else should it be?

order?
Is there any reason why it shows "see also" and "external links" then more of the article, then another "external links" section? If I dont' get any replies within a week or so, I'll go ahead and move the rest of the article up, and combine the two "external links" sections. jf 22:20, 23 May 2006 (UTC)

Other
"For instance, in eukaryotes the genetic material (DNA), and therefore transcription, is primarily localized to the nucleus, where it is separated from the cytoplasm (where translation occurs) by the nuclear membrane." This sentence has a very unclear structure*.

I miss the pictures!!!
YES! I do!!!!!!! Somebody, add some, please, it would be VERY IMPORTANT!!! Myrmeleon formicarius 12:42, 12 March 2007 (UTC)

Contradictions in Direction
In transcription, DNA is read 3'-->5' and RNA is produced 5'-->3'.

So, in an image of DNA being read by RNA polymerase, short strands would be expected at the 3' end, and long strands at the 5' end. The first diagram has this backwards.

I can't see what I'm missing, and since this is marked as a high priority for improvement, I thought this could use clarification, but since I suspect some actual biologists edit this page, I won't do it myself.

(Didn't log on the first time I posted this.)

Triacylglyceride 17:46, 13 May 2007 (UTC)triacylglyceride

 * I believe the image is marked incorrectly. Where transcription started the transcripts will be longest; the picture indicates the opposite. BTW your username is kinda funny, since it's usually either triacylglycerol or triglyceride :P -Sakkura 19:27, 9 September 2007 (UTC)

 * I think Triacylglyceride is right. Short RNA strands will be found near the start of transcription (3' end on the template strand) and the longest RNA strands near the end.  Forluvoft 00:02, 10 September 2007 (UTC)

 * Aye, that was a brain fart by me. It certainly looks correct now, although maybe you shouldn't take my word for it :P Sakkura 00:54, 14 September 2007 (UTC)

Scientists Discover the Dynamics of Transcription
'''One surprising finding was how inefficient the transcription process really is, particularly during its first two stages,� says Dr. Singer. �It turns out that only one percent of polymerases that bind to the gene actually remain on to help in synthesizing an RNA molecule. Transcription is probably inefficient for a reason.''' Brian Pearson 02:10, 7 August 2007 (UTC)

5', 3', direction of transcription, etc....
We're making this much more complicated than it has to be. We're too fixated on the directionality from the point of view of the template strand (because it's the actual physical template for RNA). But in reality, when molecular biologists talk about gene regulation, they talk about directionality from the point of view of the coding (non-template) strand. Maybe this is because the coding strand has the sequence of the resulting protein in the correct orientation. Thus, to molecular biologists, the 5' end of a gene is always near the promoter.

Hopefully this will make things clearer (if I don't screw it up):

RNA is synthesized using the template strand of DNA. RNA is synthesized from the 5' end to the 3' end. On the DNA template strand, this is 3' to 5'.

On the coding strand, however, transcription proceeds from 5' to 3'. Although RNA might not even touch the coding strand, the RNA will be an exact replica of the coding strand. Since RNA translation into protein occurs from 5' to 3', the DNA coding strand also shows what the protein sequence will look like, in the correct orientation (no reverse complementation necessary). So scientists usually like to work with the coding strand. Forluvoft 01:31, 27 September 2007 (UTC)

Reverse transcription
I added reverse transcription to the "See also" list. Does it really need its own section in this article? Forluvoft 03:53, 27 September 2007 (UTC)

 * It's called reverse transcription for a reason, so yes, it should have its own section here since it is a form of transcription (albeit unusual) Sakkura 00:00, 7 October 2007 (UTC)

 * Ok. Forluvoft 00:48, 7 October 2007 (UTC)

Transcription cartoons
I made and added some transcription cartoons. Let me know if something needs to be tweaked. The cartoons are really simplistic and I would envision them instead on an "Introduction to Molecular Genetics" page instead of this one. If this page ever gets to FA status, it will need much more detailed and accurate diagrams. Forluvoft 18:03, 12 October 2007 (UTC)

 * Yeah, they are quite simplistic, but it's far better than nothing. Especially for laymen. The protein complex at the promoter does look more like a eukaryotic preinitiation complex than what typically goes on in prokaryotes, but at least enhancers and whatnot are excluded, so I think they can be used as a general illustration for both eukaryotes and prokaryotes. Sakkura 10:53, 17 October 2007 (UTC)

Major Edits - Procedure?
Hi, new here. I'm currently doing a great deal of study of the transcription / translation area as an undergraduate at Cambridge - not saying that my opinion is any more valid than the next undergraduate and it certainly isn't compared to a PHD student, but I noticed quite a few holes here. I wondered what the procedure would be if I wanted to perform some major edits, probably adding a great deal more about Eukaryotic transcription (basal transcription factors, pol I-III etc) and doing some major rearrangements. I don't want to step on anyone's toes, but I'd like to do something useful. Martyn Axon (talk) 16:27, 27 December 2007 (UTC)
 * It would be great if we could improve the Wikipedia transcription article to featured article quality. If you want to start from scratch, create a page called Martyn Axon/Transcription. Alternatively, we could just start making major changes to the current article. Examples of featured biology articles. --JWSchmidt (talk) 21:26, 3 January 2008 (UTC)

B-Class Quality?
The information provided here is good, it does provide a broad introduction to transcription, im not complaining there. However, reading through this article i find parts where i just skip over due to grammatical errors, or just stupidity (i am not trying to be rude here i just cannot think of another word to describe it at the moment). For instance, a line just before the contents reads "Enzyme is the start of transcription". What would that mean exactly? —Preceding unsigned comment added by 152.78.249.55 (talk) 15:58, 15 January 2009 (UTC)

 * This has been fixed. --Timemutt (talk) 17:44, 11 October 2009 (UTC)

Repetition
For example:

"A polymerase binds to the 3' end of a gene (promoter) on the DNA template strand and travels toward the 5' end."

&

"The DNA template strand is read 3' → 5' by RNA polymerase and the new RNA strand is synthesized in the 5'→ 3' direction."

as found in the much too long introduction.

 * I addressed the repetition and grammatical flaws of this article. The introduction could still be shortened by including the information elsewhere if someone is feeling ambitious. --Timemutt (talk) 17:44, 11 October 2009 (UTC)

Some thoughts from a layman
The whole 5->3, 3->5 thing caused me some trouble for a while. It wasn't until I realized that the two halves of a DNA strand are going in opposite directions that it made sense.

I also think that the article is fuzzy on what kinds of cells are being discussed at points. I had trouble during the pre-initiation section trying to parse out what would apply only to prokaryotes, what would apply to eurkaryotes, and what would apply to both. Also, information is scant on archea and the distinction between archea and prokaryotic bacteria is sometimes lost. As someone who is using wikipedia article on the topic as an introduction to the material, I'm hesitant to make significant edits. Mishlai (talk) 15:53, 24 June 2009 (UTC)

 * Ah, and to make it more confusing there's still disagreement over whether all bacteria are Prokaryotes, broken into Bacteria and Archea or whether there are just Prokaryotes and Archea. I prefer the latter, but I'm definitely not getting involved in that fight. Mishlai (talk) 15:57, 24 June 2009 (UTC)

 * Although not universally accepted, I believe the trend now is to separate archae from bacteria, with only bacteria considered prokaryotes now. I made these distinctions for the information already within the article but it would be best for someone to add information differentiating more between the 3 during each stage. --Timemutt (talk) 17:44, 11 October 2009 (UTC)

Grammatical Mistake in Introduction
Hello all,

I've noticed a small grammatical mistake in the introduction to this article and would like it fixed. However, since it is in the introduction, I can't fix it myself, so I would like someone with sufficient permissions to change it for me.

The fifth paragraph has within it a sentence reading "This means it's 5' end is created first in base pairing." This is an incorrect usage of the word "it's", which should be changed to "its" in this case.

My thanks to anyone who fixes this on my behalf. —Preceding unsigned comment added by Mnkyman (talk • contribs) 02:01, 11 October 2010 (UTC)

Promoter clearance
I note that the section on promoter clearance mentions that the sigma factor stays attached to the polymerase up to 80 bp after elongation starts. However I am not entirely certain this is an accurate assessment. According to Rodney Boyer's 'Concepts in Biochemistry' (third edition), chapter 11, page 339 it stays on for ten bp, and according to page 1004 in Lehninger's 'Principles of Biochemistry' 4th edition it can stay on for 8-9 bp.

The difference between 8-10 bp and 80 bp is hardly trivial. Is this a mistake in the article, or is the cited source credible? Elvegaro (talk) 14:32, 6 June 2011 (UTC)

Hello, This article claims that transcription is an energy-dependent process and so requires ATP. While transcription does require energy, my understanding is that this energy comes from hydrolysis of the triphosphate of the incorporated nucleotide, not specifically ATP. Perhaps this article should be modified. Molecular Biology of the Cell (4th edition) pg. 305 figure 6-8. — Preceding unsigned comment added by 169.230.76.4 (talk) 16:21, 13 July 2012 (UTC)

RNA Polymerase/Helicase Confusion
The page states, "1.Helicase unwinds/"unzips" the DNA by breaking the hydrogen bonds between complementary nucleotides". Yet on the Wikipedia page for RNA Polymerase, it states, "In contrast to DNA polymerase, RNAP includes helicase activity, therefore no separate enzyme is needed to unwind DNA."

Which is it--does helicase unwind the molecule or RNA polymerase? Or is there some confusion of the terms that would resolve this issue? — Preceding unsigned comment added by 74.51.196.146 (talk) 16:09, 28 November 2011 (UTC)

hello user above^ I am not sure how to reply to your comment, so am including my reply as an edit of your original comment. Your confusion about the perpetrator of this DNA helicase activity is understandable, and I think this article on DNA transcription is quite poorly written and should be entirely reformed with input from some academics or experts in the field of genetics. I am an undergraduate biology student and I immediately identified several erroneous claims and fuzzy descriptions of what I think are generally considered vitally important intracellular mechanisms involved in this process. The RNA polymerase RNA-P II- which is the one we're discussing if considering the transcription of mRNA- is a complex enzyme consisting of 10 subunits. The helicase activity is not catalyzed by the RNA-polymerase though, it's performed by a transcription factor (TF2H) that is bound to promotor region of the DNA, one of many co-factors that assembled on the promotor region, collectively forming the the activation complex (with the addition of the polymerase) before transcription can proceed. This transcription factor, TF2H (the 2 is normally designated with roman numerals) is the most complex of the several transcription factors that will bind to the polymerase during transcription, consisting of 9 subunits. One of these subunits is a DNA helicase, This subunit hydrolyzes ATP and 'unwinds' the helical structure of the DNA, exposing the template strand that will be read by the polymerase during the elongation phase of transcription. Hope this elucidates the topic somewhat for you. Someone should fix that damn article, it's an academic disgrace.

Renamed
This article was moved from 'Translation (genetics)' to 'Transcription (biology)' to match Translation (biology), based on discussion at WT:MCB. T.Shafee(Evo & Evo)talk 00:01, 30 November 2016 (UTC)

Should RNA replication be splitted as an article?
Nowadays RNA replication is a redirect page to that article. I think that transcription and RNA replication could and should be splitted into two differenciate articles. As a key concept, trascription is a main goal in all DNA based genotypes. But in fact, at least exist some "organisms" commonly called RNA viruses (i.e.) o molecules called ribozymes, that uses and replicate RNA without transcribing it from DNA and does not use it at all (or not in a closed relation in cellullar machinery). Therefore I think that that splitting should involve at least these redirection pages: RNA synthesis, RNA replication, Template strand, Rna elongation, and RNA synthesis inhibitor. There is any volunteer to do that. I'm not a good writer in english language. Thanks and debate if you do not see it clear. --Bestiasonica (talk) 09:51, 13 December 2016 (UTC)
 * RNA "replication", by viruses and some organelles, is the copying of RNA by RNA. Terms such as RNA "transcription" and, usually at least, RNA "synthesis" refer to the copying of DNA code as RNA and typically results in the production of more RNA. They are pretty much unrelated topics. I have changed RNA replication to redirect to RNA-dependent RNA polymerase, which is a little more informative than the ribozyme article mentioned. But I agree, RNA replication does deserve a standalone article to organize the various metabolic processes that exist - ribosomal, viral, transcriptive and for all I know some other arcane corners of life. — Cheers, Steelpillow (Talk) 14:40, 11 May 2019 (UTC)

 * [Update] The article on RNA-dependent RNA polymerase, to which RNA replicase now redirects, directly contrasts RNA "replication" with DNA "transcription". The majority of sources appear to treat "RNA transcription" as the process of copying DNA code as RNA, i.e. a synonym for "DNA transcription". On that basis the viral material in the present article should simply be deleted and at best replaced by a hatnote or similar. I have tagged the article in a couple of places and if nobody comes up with a sensible alternative, I will go ahead with that. — Cheers, Steelpillow (Talk) 15:39, 12 May 2019 (UTC)

external link
these two links show 404 redirects ( not found) or Dead link '''Interactive Java simulation of transcription initiation. Interactive Java simulation of transcription interference--a game of promoter dominance in bacterial virus'''. when I try to remove it someone undoes my editing takes a look audit. thank you. Amitsabhadiyaj (Talk)

Delete figure?
The figure at the top of the page is labeled "RNA transcription" but most of it shows RNA processing, not transcription. Furthermore, it is very eukaryotic-centric and therefore not appropriate for a general article on transcription. In addition, the figure only shows RNA polymerase II transcription producing mRNA. There are a lot of noncoding genes transcribed in eukaryotes by RNA polymerases I and III.

I think we should remove this figure and replace it with one that just covers the basics of transcription in bacteria and eukaryotes.

What do others think? Genome42 (talk) 21:13, 5 June 2023 (UTC)

title: Talk:Transcription (biology)


text:
Santander, Spain

Santander is the capital of the autonomous community of Cantabria, Spain. It has a population of 172,000 (2017). It is a port city located in the northern coast of the Iberian Peninsula, facing the Cantabrian Sea. It is believed to have been a port since ancient times, due to its favourable location, and is documented as far back as the 11th century. Much of the medieval city was lost in the Great Fire of 1941. Today, its remaining old town, beach and other attractions are popular with tourists and other visitors and its economy is mainly service based. The port is still very active and a regular ferry service operates to the United Kingdom. Fish and seafood dominate the local cuisine. Santander notably houses the headquarters of multinational bank Banco Santander, which was founded there. The city has a mild climate typical of the Spanish northern coastline with frequent rainfall and stable temperatures. Cold snaps and heat waves are very rare.

Origins, Roman period and Middle Ages
The origin of the earliest human settlements in the current Santander is not easy to establish because there is little written and little archaeological data. However, there would appear to be good practical reasons for ancient settlers to have chosen the north side of the bay, sheltered from it and safer from the storms of the Bay of Biscay, on the north side of the promontory of Somorrostro and along the ancient Becedo estuary. Moreover, the hillside provided good visibility for spotting potential attackers, making this the ideal place for the foundation of a stable settlement, which was to evolve throughout the Middle Ages.

Although it is mentioned for the first time in 1068, in a draft document made by King Sancho II, in the 9th century Alfonso II the Chaste founded the Abadía de los Cuerpos Santos ("Abbey of the Holy Bodies") in the existing chapel on the hill of Somorrostro, housing as holy relics the heads of Saint Emeterius and Saint Celedonius and the graves of other unknown martyrs, giving the abbey its name.

Alfonso VIII of Castile granted the city a fuero (charter) in 1187. During the 12th and 13th centuries the population was contained within the walls of two different pueblas. La Puebla, the older, on the hill overlooking the city facing the bay, included the old castle, the Abbey of the Holy Bodies and the cloister. It had three rows of houses, separated by Rua Carnicerias and Rua Mayor, where the homes of prominent people of the town were, as well as those of the Abbot's canons. Meanwhile, the Puebla Nueva contained the convent of Santa Clara and San Francisco, which gave its name to one of the main streets; other important streets were the Rua de la Sal, The cavalcade Palace, Ribera, Don Gutierre, Puerta de la Sierra, Gallows and the Arcillero Rua. The two pueblas were joined by a bridge over the river that divided Becedo and flowed down to the shipyards, which were ordered by the king to take timber from the Cantabrian forests for shipbuilding. The villa was required to give the monarchy a ship per year.

By the end of the 15th century Santander had a population of about 4,000 or 5,000 inhabitants.

The city owes its existence to the excellent harbour of the Bay of Santander. Santander was an important port for Castile in the later Middle Ages, and also for trade with the New World. It officially became a city in 1755.

Cabo Machichaco explosions

On 3 November 1893 a steamship, SS Cabo Machichaco, caught fire while she was being unloaded at a pier in the heart of the city. A crowd of 2,000 to 3,000 people watched as crew and firefighters fought the fire. About 40 tonnes of dynamite being carried in her forward holds exploded, killing about 590 people, injuring about 2,000, sinking the ship and destroying at least 65 buildings near the harbour.

On 21 March 1894 a salvage diver working to salvage cargo from Cabo Machichaco's wreck accidentally detonated about 11 tonnes of dynamite that were submerged in the after hold of the ship. 18 people were killed and 11 injured.

Great Fire of 1941
Santander fell victim to a great fire in 1941. Fanned by a strong south wind, the fire burned for two days. The fire started in Cádiz Street, next to the harbour, the Cathedral and the medieval quarter. The fire destroyed the Old Town Hall, Jesús de Monasterio and Vargas streets and Atarazanas square buildings. It led to a major change in the architecture of Santander, away from the older small stone and wood buildings with balconies to the enormous blocks of flats built during the reconstruction.

There was only one casualty of the fire, a firefighter from Madrid killed in the line of duty, but thousands of families were left homeless and the city was plunged into chaos. The fire destroyed the greater part of the medieval town centre and gutted the city's Romanesque cathedral.

Geography
The city is located on the northern side of the Bay of Santander.

Climate
The city of Santander has an oceanic climate (Köppen climate classification Cfb), the annual thermal oscillation of the average monthly temperatures reaching around 10 °C (50 °F).

The maximum temperature reached in Santander Airport was 37.8 °C on 27 June 2009, and the minimum temperature -5.4 °C on 21 January 1957. The highest maximum daytime average for a month was in August 2003, with 27.1 C. Warm months (mean above 22 C) are however rare. The highest temperature recorded in downtown is 40.2 C in August 1940.

Sunshine hours are very low by comparison with the rest of mainland and southern Spain. Even compared with other areas of northern Spain, such as Galicia, which have many more hours of sunshine in coastal cities such as Vigo or Pontevedra. With annual averages of approximately 1650 hours of sunshine, Santander's southern areas are about as sunny as London and Paris, and quite a bit less sunny than most of England's south coastal regions. The area closer to the coast has higher sunshine time but lower summer afternoon temperatures.

Tourism and sights
The bars and restaurants of the old town are popular with tourists, as well as the El Sardinero beach a couple of kilometres away.

The Cathedral of Santander: The lower temple, called "cripta del Cristo" was built around 1200 on other earlier Roman buildings. It is 31 m long and 18 m wide, organised into three naves. Its style is a transition from Romanesque to Gothic.

The Lighthouse of Cabo Mayor presides over the entrance to the Bay of Santander.

Parque de la Vaguada de las Llamas is one of the largest parks in northern Spain, covering 11 hectare of the city.

Santander is pilot for a smart city. It is embedded with 12,000 sensors.

Politics and government
The People's Party were the leading party in the municipal elections of 1999, 2003, 2007, 2011, 2015, and 2019.

Economy
As a service centre at the regional level, Santander contains important public institutions and private organisations with a large number of employees, including Marqués de Valdecilla University Hospital, the University of Cantabria and Grupo Santander. Activities related to culture, leisure and tourism are an important part of the city's economy, and the regional and municipal authorities look to augment the summer tourist trade with additional offerings, including conventions, conferences, cultural festivals and cruises. Banco Santander, Spain's largest bank and corporation, has had its legal headquarters located in the city since its foundation.

Transport
There are ferry services to and from Portsmouth and Plymouth in the United Kingdom and Cork in Ireland, all operated by Brittany Ferries. Santander railway station serves three million annual passengers.

The city is served by the Seve Ballesteros–Santander Airport (SDR), located 4 km south of the city centre.

Education

 * University of Cantabria is the largest university in Cantabria.
 * European University of the Atlantic is a private university founded in 2013.
 * Universidad Internacional Menéndez Pelayo (UIMP) specializes in teaching Spanish and culture to foreign students.
 * Central Library of Cantabria, founded in 1839

Culture
Santander has a great tradition and cultural activity, with events that play an important role in cultural and social life of the city. UIMP is a major international summer university and organizes large festivals of music and dance. The Festival Internacional de Santander (FIS), Festival Internacional de Música de Órgano (FiMÓC), Encuentro de Música y Academia and the Paloma O'Shea International Piano Competition are main cultural events.

Diet
Santander's cuisine is characteristic of Cantabria in that it is based mainly on seafood. Popular shellfish include almejas (clams) and navajas (razor clams); fish include seabream, red mullet, anchovies, seabass and sardines; and squid and cuttlefish are also commonly eaten.

Some typical dishes from the city of Santander are the fried calamari called rabas, double donuts, bean stew called cocido montañés, and seafood dishes ranging from seabass and sardine to products such as morguera.

Historical figures

 * Francisco Marroquín (1499–1563), first bishop of Guatemala and provisional Governor of Guatemala
 * Toribio de Peñalva (c.1606-c.1685), Spanish military man, Procurator General of Buenos Aires during the Viceroyalty of Peru
 * Manuel Pardo (1774–?), a Spanish soldier, the Interim Governor of Spanish Texas in 1817
 * José de Madrazo y Agudo (1781–1859), a Spanish painter and engraver, an exponent of the Neoclassical

19th century

 * Jenaro Quesada, 1st Marquis of Miravalles (1818–1889) Grandee of Spain and Spanish soldier
 * José Antonio Mijares (1819–1847) a Mexican Army Lieutenant who led the Mexican resistance force against the American garrison of San José del Cabo in the Battle of San José del Cabo
 * Rafael Izquierdo y Gutiérrez (1820–1883) a Spanish Military Officer, politician, and statesman; Governor-General of the Philippines from 1871 to 1873
 * Marcelino Sanz de Sautuola (1831–1888) a Spanish jurist, amateur archaeologist, owned the land of the Cave of Altamira
 * Marcelino Menéndez y Pelayo (1856–1912) a Spanish scholar, historian and literary critic.
 * Francisco Iturrino (1864–1924) a Spanish Post-impressionist painter of Basque ancestry, sometimes called a Fauvist
 * Concha Espina (c.1877–1955) a Spanish writer nominated for a Nobel prize in literature 25 times in nine years
 * Marcial Solana González-Camino (1880–1958) a Spanish scholar, writer and politician; historian of philosophy
 * María Gutiérrez Blanchard (1881–1932) a Spanish painter, developed a unique style of Cubism
 * Ángel Herrera Oria (1886–1968) a Spanish journalist, Roman Catholic politician and later a cardinal
 * Gerardo Diego (1896–1987) a Spanish poet, a member of the Generation of '27

20th century

 * Emilio Botín (1903–1993), a Spanish banker, the chairman of Santander Group from 1950 to 1986
 * José Luis Zamanillo (1903–1980), a Traditionalist politician and leader of Carlist paramilitary Requeté structures
 * Francisco de Borbón y Borbón (1912–1995), a Spanish aristocrat, Lieutenant General of the cavalry in the Spanish army
 * Matilde Camus (1919–2012), a Spanish poet who also wrote non-fiction
 * Elena Quiroga (1921–1995), Spanish writer, explored the themes of childhood and adolescence
 * Daniel Gil (1930–2004), one of the leading Spanish graphic designers of the 20th century
 * Emilio Botín (1934–2014), a Spanish banker, executive chairman of Spain's Grupo Santander
 * Juan Carlos Calderon (1938–2012), a Spanish singer-songwriter and musician
 * Álvaro Pombo (born 1939), a Spanish poet, novelist and activist
 * Juan Navarro Baldeweg (born 1939), architect and professor at the Superior Technical School of Architecture of Madrid
 * Alfonso Vallejo (1943–2021), playwright, poet, painter and neurologist
 * Germán Gullón (born 1945), literary critic, writer and professor of Spanish literature at the University of Amsterdam
 * Domingo Sarrey (born 1948), a visual artist and video artist.
 * José Antonio Rodríguez Vega (1957–2002), nicknamed El Mataviejas (The Old Lady Killer), was a Spanish serial killer who raped and killed at least 16 elderly women in and around Santander between August 1987 and April 1988
 * Paco San José (born 1958), Cantabrian music composer
 * Álvaro Longoria (born 1968), a film director, executive producer and actor

Athletes

 * Francisco "Paco" Gento López (21 October 1933 - 18 January 2022), nicknamed Paco, was a Spanish footballer or Spanish soccer player, 437 appearances between Racing Santander and Real Madrid and scored a total of 129 goals in his senior career. He played for the National Spanish team in 43 aapearances. and scored 5 goals.
 * Marcos Alonso Imaz (1933–2012), nicknamed Marquitos, was a Spanish footballer, 272 pro appearances
 * José Pérez Francés (1936–2021), a Spanish former professional road racing cyclist
 * Francisco Javier Aguilar Garcia (1949–2020), a Spanish retired professional footballer 300 pro appearances
 * Seve Ballesteros (1957–2011), a Spanish professional golfer and World No. 1
 * Quique Setién (born 1958), a Spanish retired footballer, 518 pro appearances, former coach of FC Barcelona
 * Marcos Alonso Peña (1959–2023), a Spanish retired footballer, and a coach, 309 pro appearances
 * Iván Helguera (born 1975), a Spanish football player, 326 pro appearances
 * Pedro Munitis (born 1975), a Spanish football player, 475 pro appearances
 * Iván de la Peña (born 1976), a Spanish football player, 331 pro appearances
 * Mario Bermejo (born 1978), a Spanish retired professional footballer, 546 pro appearances
 * Ruth Beitia (born 1979), high jumper, gold medallist at the 2016 Olympic Games
 * Gonzalo Colsa (born 1979), a Spanish retired footballer 394 pro appearances
 * Mateo Joseph (born 2003), a Spanish footballer for Leeds United and the Spain national under-21 football team.

Sports
Racing de Santander is the main football team in the city, playing their home games at the Campos de Sport de El Sardinero.

Some elite teams of Santander:

title: Santander, Spain


text:
Talk:Power (social and political)

Merge
In the German page, the "bases of power" have already been integrated to the article on power.

 * I agree with the sugestion of merging the 2 articles about power. The ideas discussed are very similar.
 * Thanks

Pace?
what is meant by pace in "(pace advocates of empowerment)"? --Reagle 01:57, 28 Apr 2004 (UTC)

Quote
I can't see how the above quote belongs in this article. It seems to me they should create their own article if they want to advertise/promote their organization.

"the creation in 1990 of a permanent coalition of progressive organizations — the ... is a coalition of labor and community organizations with broad demographic representation of the state." Jim 16:01, 20 Jun 2005 (UTC)

Correct citation for Galbraith?
The article cites An Anatomy of Power by Galbraith JK; the only citation I can find is for The anatomy of power / John Kenneth Galbraith; Publisher info Boston : Houghton Mifflin, 1983. I wonder if the cite is wrong in the article, or if this is the one intended?

Steven Lukes
Steven Lukes found the assumption about power is “that A in some way affects B” to be too broad because everyone is affecting others in countless manners. In order to specify when a notion of power is significant in order to be useful for analysis the conception needs to be specified. A more specific conception of power that specifies when it is significant for analysis is when “A exercises power over B when A affects B in a manner contrary to B's interests”. Note that a distinction can be made between B's real interests and B's subjective interests. This applies to situations where B does not express or even is not conscious of real interests. Admitted, it is difficult to ascertain what the real interests of B are. Only B can identify real interests under conditions of relative autonomy and independent of A's power.

The first dimension of Lukes three-dimensional view of power focuses on observable behaviour where decisions are made over issues with an observable conflict of interest. This is akin to the definition Max Weber gave to power; “Power is the probability of individuals realizing their wills despite the resistance of others”. This notion of power can be analysed fairly easy, because it focuses on observable data. It merely requires the analyst to take note of every successful attempt of A to “get B to do something that B would otherwise not do”. However, this notion alone is too limited in various aspects. These shortcomings are addressed in the other two dimensions of Lukes view on power.

In the second dimension of Lukes view on power the behavioural aspect of the first dimension is critiqued. It asserts that B can be prevented by A from taking decisions on issues “over which there is an observable conflict of interests”. The action where A suppresses B to voice an opinion is termed nondecision. One way nondecisionmaking can be achieved is if A engages in creating or reinforcing structural issues, such as social and political values and institutional practices, limiting the political process to only those issues that are in A's interest. This is akin to Elmer Eric Schattschneider statement that “organization is the mobilization of bias”.

Nondecision-making can be achieved by controlling the political agenda, but it can also be achieved by securing the compliance of B through various means. These means are coercion, influence, authority, force, and manipulation. Coercion is understood as the threat of deprivation. Influence exists if A changes B's course of action without threat. Authority exists if B experiences A's command to be compatible with his own values, either because of legitimate and reasonable content or because it arrive through a legitimate and reasonable procedure. In the case of force A removes B's choice of compliance or noncompliance. Manipulation is an aspect of force, because compliance is gained absent B's recognition of the source or exact nature of A's demand.

In both the first and second dimension of Lukes view on power there is a necessity for observable conflict between interests. This becomes problematic if A successfully suppresses B's voice to be heard. However, Lukes ascertains that although the interests of B are excluded from public attention and controversy, they can still be observed outside the political system by an investigator.

The third dimension of Lukes view on power is addressing power where there appears to be an absence of conflict. This dimension has a more collectivist perspective on power, asserting that mobilization of bias is sustained by social and cultural behaviour of groups, and institutional practices. It recognizes that leaders are not only responding to preferences, but are also shaping them. Shaping preferences can be achieved by controlling information, and the processes of socialization. The absence of conflict is achieved, because A influences, shapes, or determines the perceptions, cognitions, and preferences of B. This can be done either through manipulation or legitimate authority. This dimension appears to be similar to the concept of hegemony developed by Antonio Gramsci, which is power obtained through the manufacture of consent. In the case of consent through manipulation there is still an absence of conflict, despite the decisions are going against B's real interests. This is because B's “perceptions, cognitions, and preferences are shaped in such a way that they accept their role in the existing order of things, either because they can see or imagine no alternative to it, or because they see it as natural and unchangeable, or because they value it as divinely ordained and beneficial”. It is also possible that B gives consent through authority, even is the decisions are going against B's real interest. This can be explained if B values collective interests, or the individual interests of A, as being of higher importance than his own interests.

The concept of inequality is understood as a difference of power between actors. This difference can be analysed by looking at differences between power resources that actors have available, can access, and are able to utilize. Power resources can exist in the form of capital, property rights, political influence, discursive resources, production and access to information, violence and force, authority, weapons of the weak (to be understood as every day resistance in the form of lifestyle choices. For example, consumption habits) and social category (gender, ethnicity, age, class, nationality). Inequality can arise among any or all of these power resources, directly affecting the power an actor has in relation to other actors. In turn the power an actor has can affect the power resources, creating a potential vicious or virtuous cycle. This suggests that increases or decreases in inequality do not always happen intentionally, but can be a result of such vicious or virtuous cycles.

title: Talk:Power (social and political)


text:
Quandong

Quandong, quandang or quondong is a common name for the species Santalum acuminatum (desert, sweet, Western quandong), especially its edible fruit, but may also refer to:

 * Aceratium concinnum (highroot quandong)
 * Peripentadenia mearsii (buff, grey quandong)
 * Elaeocarpus angustifolius (blue quandong)
 * Elaeocarpus arnhemicus (Arnhem Land, bony quandong)
 * Elaeocarpus bancroftii (Kuranda quandong)
 * Elaeocarpus coorangooloo (brown quandong)
 * Elaeocarpus eumundi (eumundi, smooth-leaved quandong)
 * Elaeocarpus ferruginiflorus (rusty leaf quandong)
 * Elaeocarpus foveolatus (white, Northern quandong)
 * Elaeocarpus grandis (brush, blue, white quandong; quandong)
 * Elaeocarpus holopetalus (mountain quandong)
 * Elaeocarpus johnsonii (Kuranda quandong)
 * Elaeocarpus kirtonii (brown-hearted quandong)
 * Elaeocarpus kirtonii (white quandong)
 * Elaeocarpus largiflorens (tropical quandong)
 * Elaeocarpus obovatus (hard quandong)
 * Elaeocarpus reticulatus (Ash quandong)
 * Elaeocarpus ruminatus (brown, grey quandong)
 * Elaeocarpus sericopetalus (Northern, hard, Northern hard quandong)
 * Elaeocarpus williamsianus (hairy quandong)
 * Santalum acuminatum (desert quandong)
 * Santalum lanceolatum (desert quandong)
 * Santalum murrayanum (bitter quandong)
 * Terminalia cunninghamii (pindan quandong)
 * The specific epithet of Amyema quandang, discovered growing on Santalum acuminatum

title: Quandong