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Freedom of the seas
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This article is about the legal concept. For other uses, see Freedom of the Seas (disambiguation).
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Freedom of the seas (Latin: mare liberum, lit. "free sea") is a principle in the international law and sea. It stresses freedom to navigate the oceans. It also disapproves of war fought in water. The freedom is to be breached only in a necessary international agreement.
Freedom of the Seas, World War II US poster
This principle was one of U.S. President Woodrow Wilson's Fourteen Points proposed during the First World War. In his speech to the Congress, the president said:
Absolute freedom of navigation upon the seas, outside territorial waters, alike in peace and in war, except as the seas may be closed in whole or in part by international action for the enforcement of international covenants.
The United States' allies Britain and France were opposed to this point, as France was also a considerable naval power at the time. As with Wilson's other points, freedom of the seas was rejected by the German government.
Today, the concept of "freedom of the seas" can be found in the United Nations Convention on the Law of the Sea under Article 87(1) which states: "the high seas are open to all states, whether coastal or land-locked". Article 87(1) (a) to (f) gives a non-exhaustive list of freedoms including navigation, overflight, the laying of submarine cables, building artificial islands, fishing and scientific research.
Contents
[hide]
1 Historical background
2 High seas and registration
3 Trade opportunities
4 See also
5 References
6 Sources
Historical background[edit]
Between the end of the 15th century up until the 17th century various powers claimed sovereignty over parts of the sea. In 1609, Dutch jurist and philosopher Hugo Grotius wrote what is considered the foundation of international legal doctrine regarding the seas and oceans – Mare Liberum, a Latin title that translates to "freedom of the seas". The Grotian concept of "freedom of the sea" was only accepted after a century long debate between Grotius' ideas and John Selden's.[1]
While it is generally assumed that Grotius first propounded the principle of freedom of the seas, countries in the Indian Ocean and other Asian seas accepted the right of unobstructed navigation long before Grotius wrote his De Jure Praedae (On the Law of Spoils) in the year of 1604. Previously, in the 16th century, Spanish theologian Francisco de Vitoria postulated the idea of freedom of the seas in a more rudimentary fashion under the principles of jus gentium.[2]
During World War II, nations started to expand and claim many resources and water territories all over their surrounding coasts. There were four international treaties meticulously drafted in the late 1950s and onto the 1970s, but the issues were not resolved between nations until 1982 when the United Nations Convention on the Law of the Sea was introduced.
UNCLOS is a Law of the Sea treaty: an agreement of rights and responsibilities of nations and their use of the world’s ocean with guidelines of trade, environment, and the management of marine and open seas resources. UNCLOS replaced the four international treaties drafted in the late 1950s through 1970s. As of 2013, 165 countries and the European Union have joined the Convention.
High seas and registration[edit]
According to international law, Article 92 of the convention which describes ships shall sail under the flag of one state only and, save in exceptional cases expressly provided for in international treaties or in this Convention, shall be subject to its exclusive jurisdiction on the high seas;[3] however, when a ship is involved in certain criminal acts, such as piracy,[4] any nation can exercise jurisdiction under the doctrine of universal jurisdiction. High seas were defined as any part of the sea that was not either territorial sea or internal waters, territorial waters and exclusive economic zones. Article 88 of the 1982 Convention states that the high seas shall be reserved for peaceful purposes. Many countries engage in military maneuvers and the testing of conventional weapons and nuclear weapons on the high seas.[5][6] In order to deliver the right punishment to the right person or state, ships must be registered to a country to show proof of ownership.[7] The owner of the vessel sometimes prefers to pay the lower registration fees by picking countries such as Panama, Bermuda, Italy, Malta and the Netherlands. According to Cruise Lines International Association, 90% of commercial vessels calling on U.S. ports fly foreign flags.[8] To avoid the high cost with more rules and regulations, ships and tankers sometime prefer lower cost registration with a lower standard of inspection and regulation by picking a country that exercises less control over their registered ships,[9] though many ships are owned by individuals or companies in another country (most commonly Japan and Greece) under a system called "flag of convenience". Registering a ship in Panama means that the ship is governed by the maritime rules of Panama rather than the ship owner's country. Ship owners do this because Panama has low taxes and fewer labor and safety regulations than most other countries. Ship owners can make their staff work longer hours in less safe environments, and therefore maximize their profits. Other countries, including Liberia, Cyprus and the Bahamas also offer flags of convenience, but Panama has the most ships registered under the scheme. Ships registered with the US will cost more, and the employee wages will be even higher.[8] Freedom of the seas allows a ship to move freely on the ocean as long as it follows international law.
Trade opportunities[edit]
From the Vikings to the European, Central Asia, Africa and North and South America, trade has served an important role in history, and has been a key factor of a growing economy. Trade transfers the ownership of goods from one person or entity to another by getting a product or service in exchange from the buyer. When a ship sets sail, there may be many ports waiting for it to bring goods from all over the world for trade and sale. Free trade opening up markets to foreign suppliers increases competition. Without free trade, domestic companies may have enjoyed monopolies or oligopolies that enabled them to keep prices well above marginal costs. Trade liberalization will undermine that market power.[10]
The World Trade Organization states that "The ocean is a big part of the free trade happening around the world. Every day we see silk imported from China, fruits from Mexico, spices from India, chocolate from Africa and many more goods in our home were imported from another country. The ocean made it possible for many countries to put their cargoes on to ships and transfer them across the ocean. Hundreds of nations joined the World Trade Organization. With goods to be traded from one nation to another through the transportation on various size vessels from coast to coast, trade promoted more economic value to goods where goods and services are produced to support the supply and demand of the participants in the organization."[11]
The Merchant Marine Act of 1920, also known as the Jones Act (46 USC § 883) is a maritime law that controls coastal trade within the United States and determines which ships may lawfully engage in that trade and the rules under which they must operate.[12] The act is in place to protect jobs for US citizens and its people working at U.S. ports, and on U.S. vessels. Many vessels around the globe sail under many different flags, and have different crews from different parts of the world where the pay rate is much lower than the U.S.[citation needed] The Jones Act protects the Americans' jobs and restricts coastwise transportation of passengers. 46 USC § 12108 additionally restricts the use of foreign vessels to commercially catch or transport fish in U.S. waters.
See also[edit]
Water portal
Dominium maris baltici
Mare liberum
Passenger Vessel Services Act of 1886, a similar law concerning passenger transportation between US ports.
Seaman status in United States admiralty law
Treaty of Alcáçovas
Treaty of Tordesillas
References[edit]
Jump up ^ Trullio Treves, High Seas, Max Planck Foundation for International Peace and the Rule of Law (2009).
Jump up ^ Arthur Nussbaum (1947). A concise history of the law of nations. Macmillan Co. p. 62.
Jump up ^ UNCLOS article 92(1)
Jump up ^ UNCLOS article 105
Jump up ^ UNCLOS article 88(1)
Jump up ^ See Trinity (nuclear test) and Ivy Mike.
Jump up ^ "A Guide to Ship Registration" (PDF). Maritime New Zealand. 30 September 2010. Retrieved 12 December 2012.
^ Jump up to: a b "Why are cruise ships registered in foreign countries?". Special for USA TODAY. 8 January 2013. Retrieved 12 December 2012.
Jump up ^ Epps, V. International Law, (2009) pg. 247
Jump up ^ "A short tour of economic theory". A survey of world trade. Retrieved 2014-04-22.
Jump up ^ "The WTO in Brief". WTO. Retrieved 2014-04-22.
Jump up ^ "THE JONES ACT". Mike Vaughn, Attorney at Law. Retrieved 22 April 2014.
Sources[edit]
Freedom_of_the_seas by Hugo Grotius ((in English))
Retrieved from "https://en.wikipedia.org/w/index.php?title=Freedom_of_the_seas&oldid=838140535"
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This Is the Way It Goes and Goes and Goes
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This Is the Way It Goes & Goes & Goes
Studio album by Juno
Released
March 30, 1999
Recorded
November 1998
Genre
Indie rock
Length
53:50
Label
DeSoto
Producer
Kip Beelman
Juno chronology
This Is the Way It Goes & Goes & Goes
(1999) A Future Lived in Past Tense
(2001)A Future Lived in Past Tense2001
Professional ratings
Review scores
Source
Rating
Allmusic [1]
PandoMag [2]
Pitchfork Media (8.3/10) [3]
This Is the Way It Goes & Goes & Goes is the debut album by Seattle band Juno, released in 1999 on DeSoto Records.
Track listing[edit]
"The Great Salt Lake/Into the Lavender Crevices of Evening the Otters Have Been Pushed" - 5:31
"Rodeo Programmers" - 2:53
"The Young Influentials" - 4:00
"All Your Friends Are Comedians" - 3:27
"Leave a Clean Camp and a Dead Fire" - 9:51
"January Arms" - 8:33
"Venus on 9th Street" - 3:14
"A Listening Ear" - 7:19
"Are You Still There?" - 0:55
"The Sea Looked Like Lead" - 8:07
References[edit]
Jump up ^ Allmusic review
Jump up ^ PandoMag review
Jump up ^ Pitchfork Media review
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Valley of the Dolls
From Wikipedia, the free encyclopedia
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For other uses, see Valley of the Dolls (disambiguation).
Valley of the Dolls
First Hardcover Edition
Author
Jacqueline Susann
Country
United States
Language
English
Published
1966 (Bernard Geis Associates)
Media type
Print (Hardback & Paperback)
Pages
442 pp (hardcover edition)
ISBN
978-0802135193
Preceded by
Every Night, Josephine!
Followed by
The Love Machine
Valley of the Dolls is the first novel by American writer Jacqueline Susann. Published in 1966, the book was the biggest selling novel of its year.[1] To date, it has sold more than 31 million copies,[2] making it one of the best-selling works in publishing history.[3]
Contents
[hide]
1 Plot summary
2 Background
3 Reception
4 Adaptations
5 Notes
6 References
7 External links
Plot summary[edit]
The novel tells the story of three young women who become fast friends in the turbulent post-war worlds of Broadway and Hollywood: Anne Welles, a reserved New England beauty who sees New York as the romantic city of her dreams; Neely O'Hara, an ebullient vaudevillian with a talent she doesn't fully understand; and Jennifer North, a sweet-natured showgirl who wants only to be loved. As life becomes difficult, each woman grows increasingly dependent on "dolls," the amphetamines and barbiturates which, for a time, seem to help.
Crossing their paths are such people as Helen Lawson, a brilliant but ruthless Broadway legend; Lyon Burke, a magnetic but self-centered theatrical attorney; Tony Polar, a child-like but sexually aggressive pop singer; Kevin Gillmore, a powerful but lovesick cosmetics tycoon; and Ted Casablaca, a potent but opportunistic fashion designer.
Over the course of twenty eventful years (1945–1965), each of the women strives to ascend the Mount Everest of her dreams, only to find herself back in the valley of the dolls.
Background[edit]
Susann had apparently been thinking about the novel for some time. Some years earlier, she had begun a show business novel, Underneath the Pancake, with her friend, actress Beatrice Cole (c. 1910–1999).[4] Later, she considered writing a novel about drug usage in show business to be called The Pink Dolls.[5]
Valley of the Dolls is widely considered to be a roman a clef, with its characters based on such famous figures as Judy Garland, Carole Landis, Dean Martin, and Ethel Merman.[6] In 1973, after publication of her third novel, Susann said, "They can keep calling it that [roman a clef]. It'll only make my books sell, I don't care."[7] Susann insisted that she began each book with a theme: "Then I start asking, what kind of a personality? And because I have a good ear, I unconsciously pick up certain people."[7]
Susann dedicated the book to her poodle, Josephine, and to her husband, Irving Mansfield.[8]
Reception[edit]
The book was published by Bernard Geis Associates on February 10, 1966, and "took off like a Cape Canaveral space shot." [9]
Although Publishers Weekly, in an advance review, called the novel "big, brilliant and sensational" (if "poorly written"),[10] the book received largely negative reviews. Gloria Steinem panned the book in The New York Herald Tribune [11] as did the reviewer in The New York Times.[12] Time magazine called it the "Dirty Book of the Month," and said, "it might more accurately be described as a highly effective sedative, a living doll."[13]
Despite the poor reviews, the book was a commercial juggernaut. On May 8, 1966, in its ninth week on the list, the book reached #1 on the New York Times Best Seller List, where it remained for 28 consecutive weeks.[14] With a total of 65 weeks on the list, the book became the best selling novel of 1966.[1] By the time of Susann's death in 1974, it had entered the Guinness Book of World Records as the best selling novel in publishing history, with more than 17 million copies sold.[15] By 2016, the book had sold more than 31 million copies.[2]
Adaptations[edit]
In 1967 the book was adapted into a film of the same name, directed by Mark Robson (Peyton Place), and starring Barbara Parkins (as Anne), Patty Duke (Neely), Paul Burke (Lyon), Sharon Tate (Jennifer), and Susan Hayward (Helen). The movie was written by Helen Deutsch (National Velvet) and Dorothy Kingsley (Seven Brides for Seven Brothers), and produced by Robson and David Weisbart. Reviews were scathing,[16] but the film was an enormous box-office hit, becoming the sixth most popular of the year with $44 million at the domestic box office.[17] Susann, who had a cameo as a news reporter, hated the film, reportedly telling director Robson that it was "a piece of shit."[18]
The novel was adapted for television in 1981 as Jacqueline Susann's Valley of the Dolls, a mini-series executive-produced by Susann's widower Irving Mansfield and directed by Walter Grauman. This version stars Catherine Hicks, Lisa Hartman, and Veronica Hamel. In 1994 a late-night, syndicated television soap opera, Valley of the Dolls, ran for one season and 65 episodes. The premise was a loose adaptation of the novel.[19] BBC Radio 4 broadcast a 15-episode dramatisation scripted by Yvonne Antrobus over three weeks in August and September 2005. It was part of the Woman's Hour programme's ongoing fifteen-minute daily drama slot,[20] and has been rebroadcast several times on BBC Radio 4 Extra in three 70-minute omnibus episodes.[21]
Notes[edit]
References[edit]
^ Jump up to: a b The 20th-Century American Bestsellers Database: 1960s. University of Virginia, via Publishers Weekly, 2016. Retrieved January 10, 2017.
^ Jump up to: a b Symonds, Alexandria. 'Valley of the Dolls,' by the numbers. T: The New York Times Style Magazine. February 9, 2016. Retrieved January 10, 2017.
Jump up ^ Best Seller. Encyclopedia Britannica. February 23, 2011. Retrieved January 10, 2017.
Jump up ^ Seaman, Barbara. Lovely Me: The Life of Jacqueline Susann. 2nd ed. (New York: Seven Stories Press, 1996), p. 197.
Jump up ^ Seaman. Lovely Me, p. 251-252.
Jump up ^ Collins, Amy Fine. Once Was Never Enough. Vanity Fair, January 2000. Retrieved January 5, 2017.
^ Jump up to: a b Kasindorf, Martin. Jackie Susann Picks Up the Marbles. The New York Times, August 12, 1973. Retrieved January 5, 2017.
Jump up ^ Susann, Jacqueline. Valley of the Dolls. (New York: Bernard Geis Associates, 1966).
Jump up ^ Daniels, Mary. Susann's Best Love Story a Private Affair. Chicago Tribune. August 15, 1976. Retrieved January 10, 2017.
Jump up ^ Mansfield, Irving and Jean Libman Block. Life with Jackie. (New York: Bantam Books, 1983), p. 132.
Jump up ^ Seaman, Lovely Me, p. 314.
Jump up ^ Fremont-Smith, Eliot. Thank You, Franz Kafka! The New York Times. February 4, 1966. Retrieved January 10, 2017.
Jump up ^ Books: Dirty Book of the Month. Time. April 22, 1966. Retrieved January 10, 2017.
Jump up ^ Adult New York Times Best Seller Lists for 1966. Hawes Publications. [n.d.] Retrieved January 10, 2017.
Jump up ^ Johnston, Laurie. Jacqueline Susann Dead at 53; Novelist Wrote 'Valley of Dolls'. The New York Times. September 23, 1974. Retrieved January 9, 2017.
Jump up ^ Seaman. Lovely Me, p. 349.
Jump up ^ Top Grossing Films of 1967. Listal. June 24, 2012. Retrieved January 13, 2017.
Jump up ^ Green, David B. This Day in Jewish History 1974: Jacqueline Susann, Who Knew What You Really Want to Read, Dies. Haaretz. September 21, 2016. Retrieved January 13, 2017.
Jump up ^ Gardner, Eriq (March 16, 2012). "Lawsuit Threatens Lee Daniels' 'Valley of the Dolls' TV Series". The Hollywood Reporter.
Jump up ^ Radio Listings
Jump up ^ BBC Radio omnibus edition catalogue entry
External links[edit]
Valley of the Dolls The official website, run by the estate of Jacqueline Susann.
[hide]
v
t
e
Jacqueline Susann
Books
Every Night, Josephine! (1963)
Valley of the Dolls (1966)
The Love Machine (1969)
Once Is Not Enough (1973)
Dolores (1976)
Yargo (1979)
Adaptations
Valley of the Dolls (1967)
The Love Machine (1971)
Once Is Not Enough (1975)
Valley of the Dolls (TV series, 1994)
Related articles
Isn't She Great
Jacqueline Susann's Open Door
Just This Once
Irving Mansfield
Retrieved from "https://en.wikipedia.org/w/index.php?title=Valley_of_the_Dolls&oldid=799961415"
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Widget (beer)
From Wikipedia, the free encyclopedia
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This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (September 2010) (Learn how and when to remove this template message)
Guinness floating widget
Guinness beer bottle widget
A widget is a device placed in a container of beer to manage the characteristics of the beer's head. The original widget was patented in Ireland by Guinness. The "floating widget" is found in cans of beer as a hollow plastic sphere, approximately 3 cm in diameter (similar in appearance to table tennis ball, but smaller) with at least one small hole and a seam. The "rocket widget" is found in bottles, 7 cm in length with the small hole at the bottom.[1]
Contents
[hide]
1 Background
2 Method
3 Development
4 Beer glass widget
5 References
6 Bibliography
Background[edit]
Draught Guinness, as it is known today, was first produced in 1964. With Guinness keen to produce draught beer packaged for consumers to drink at home, Bottled Draught Guinness was formulated in 1978 and launched into the Irish market in 1979. It was never actively marketed internationally as it required an "initiator" device, which looked rather like a syringe, to make it work.
Method[edit]
Some canned beers are pressurized by adding liquid nitrogen, which vaporises and expands in volume after the can is sealed, forcing gas and beer into the widget's hollow interior through a tiny hole—the less beer the better for subsequent head quality. In addition, some nitrogen dissolves in the beer which also contains dissolved carbon dioxide. Oxygen is generally excluded as its presence can cause flavour deterioration.
The presence of dissolved nitrogen allows smaller bubbles to be formed, which increases the creaminess of the head. This is because the smaller bubbles need a higher internal pressure to balance the greater surface tension, which is inversely proportional to the radius of the bubbles. Achieving this higher pressure would not be possible with just dissolved carbon dioxide, as the greater solubility of this gas compared to nitrogen would create an unacceptably large head.
When the can is opened, the pressure in the can quickly drops, causing the pressurised gas and beer inside the widget to jet out from the hole. This agitation on the surrounding beer causes a chain reaction of bubble formation throughout the beer. The result, when the can is then poured out, is a surging mixture in the glass of very small gas bubbles and liquid.
This is the case with certain types of draught beer such as draught stouts. In the case of these draught beers, which before dispensing also contain a mixture of dissolved nitrogen and carbon dioxide, the agitation is caused by forcing the beer under pressure through small holes in a restrictor in the tap. The surging mixture gradually settles to produce a very creamy head.
Development[edit]
Expired British Patent No 1266351, filed 27 January 1969
Diagram of Acorn Can
In 1969 two Guinness brewers at Guinness's St James's Gate brewery in Dublin, Tony Carey and Sammy Hildebrand, developed a system for producing draught type Guinness from cans or bottles through the discharge of gas from an internal compartment. It was patented in British Patent No 1266351, filed 27 January 1969, with a complete specification published 8 March 1972.
Development work on a can system under Project ACORN (Advanced Cans Of Rich Nectar) focused on an arrangement whereby a false lid underneath the main lid formed the gas chamber (see diagram below right).
Technical difficulties led to this approach being put on hold, and Guinness instead concentrated on bottles using external initiators. Subsequently, Guinness allowed this patent to lapse and it was not until Ernest Saunders centralised the company's research and development in 1984 that work restarted on this invention, under the direction of Alan Forage.
Development sequence for canned draught Guinness
Development sequence for bottled draught Guinness
The design of an internal compartment that could be readily inserted during the canning process was devised by Alan Forage and William Byrne, and work started on the widget during the period 1984–85. The plan was to introduce a plastic capsule into the can, pressurise it during the filling process and then allow it to release this pressure in a controlled manner when the can was opened. This would be sufficient to initiate the product and give it the characteristic creamy head. However, Tony Carey observed that this resulted in beer being forced into the widget during pressurisation, which reduced the quality of the head. He suggested overcoming this by rapidly inverting the can after the lid was seamed on. This extra innovation proved successful.
The first samples sent to Dublin were labelled "Project Dynamite", which caused some delay before customs and excise would release the samples.[citation needed] Because of this the name was changed to Oaktree in recognition of the earlier ACORN project. Another name that changed was "inserts"; the operators called them "widgets" almost immediately after they arrived on site, a name that has now stuck with the industry.[citation needed]
The development of ideas continued and more than one hundred alternatives were considered. The blow-moulded widget was to be pierced with a laser and a blower was then necessary to blow away the plume created by the laser burning through the polypropylene. This was abandoned and instead it was decided to gas-exchange air for nitrogen on the filler,[2] and produce the inserts with a hole in place using straightforward and cheaper injection-moulding techniques.
Commissioning began January 1988, with a national launch date of March 1989. This first-generation widget was a plastic disc held in place by friction in the bottom of the can. This method worked fine if the beer was served cold; when served warm the can would overflow when opened. The floating widget, which Guinness calls the "Smoothifier", was launched in 1997 and does not have this problem.
The diagrams on the left show the development sequences for canned and bottled draught Guinness from 1969 to 1988.
The idea for the widget soon became popular. John Smith's started to include widgets in their cans in 1994 and many beer brands in the UK now use widgets, often alongside regular carbonated products.
Although patented by Guinness, the widget was actually invented by John Lunn, MD of Mclennons of Birmingham, who went on to invent a second for Whitbread and Heineken, so that Whitbread could launch Draught Boddingtons in a can and Murphy's. Lunn then later invented a third widget, the floating one, with two one-way valves, which is the widget that all brewers use now.
Beer glass widget[edit]
Circular widget etched in the base of a standard pint glass
Comparison of bubbles formed in a glass containing a widget (left) and one with a smooth base (right).
The term widget glass can be used to refer to a laser-etched pattern at the bottom of a beer glass which aids the release of carbon dioxide bubbles.[3] The pattern of the etching can be anything from a simple circular or chequered design to a logo or text.
The widget in the base of a beer glass works by creating a nucleation point, allowing the CO2 to be released from the liquid which comes into contact with it, thus assisting in maintaining head on the beer. This has become increasingly popular with Fosters, Estrella and others using them in public houses in the UK.
References[edit]
Jump up ^ Daithí Ó hAnluain (17 December 2001). "Foaming Over Guinness in a Bottle". Wired. Cork, Ireland. Archived from the original on 29 June 2013.
Jump up ^ Fitzpatrick, Nicholas; John Kuzniarski (3 August 1993). "CA Patent 1320934 - Gas Dissolving Method". Retrieved 2009-11-15.
Jump up ^ "Widget Glass". www.probertencyclopaedia.com. Retrieved 2009-08-28.
Bibliography[edit]
Carey & Hildebrand,Improved method of and means for dispensing carbonated liquids from containers, UK Patent 1266351, published 8 March 1972 — the original invention behind the modern widget.
Forage, et al., "Beverage package and a method of packaging a beverage containing gas in solution". United States Patent 4,832,968. 23 May 1989.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Widget_(beer)&oldid=811290957"
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Transfer RNA
From Wikipedia, the free encyclopedia
Jump to: navigation, search
The interaction of tRNA and mRNA in protein synthesis.
tRNA
Identifiers
Symbol
tRNA
Rfam
RF00005
Other data
RNA type
gene, tRNA
SO
{{{SO}}}
Available PDB structures
3icq, 1asy, 1asz, 1il2, 2tra, 3tra, 486d, 1fir, 1yfg, 3eph, 3epj, 3epk, 3epl, 1efw, 1c0a, 2ake, 2azx, 2dr2, 1f7u, 1f7v, 3foz, 2hgp, 2j00 , 2j02, 2ow8, 2v46, 2v48, 2wdg, 2wdh, 2wdk, 2wdm, 2wh1
A transfer RNA (abbreviated tRNA and formerly referred to as sRNA, for soluble RNA[1]) is an adaptor molecule composed of RNA, typically 76 to 90 nucleotides in length,[2] that serves as the physical link between the mRNA and the amino acid sequence of proteins. tRNA does this by carrying an amino acid to the protein synthetic machinery of a cell (ribosome) as directed by a three-nucleotide sequence (codon) in a messenger RNA (mRNA). As such, tRNAs are a necessary component of translation, the biological synthesis of new proteins in accordance with the genetic code.
Contents
[hide]
1 Overview
2 Structure
3 Anticodon
4 Aminoacylation
5 Binding to ribosome
6 tRNA genes
6.1 Evolution
6.2 tRNA-derived fragments
7 tRNA biogenesis
8 History
9 See also
10 References
11 External links
Overview[edit]
While the specific nucleotide sequence of a mRNA specifies which amino acids are incorporated into the protein product of the gene from which the mRNA is transcribed, the role of tRNA is to specify which sequence from the genetic code corresponds to which amino acid.[3] The mRNA encodes a protein as a series of contiguous codons, each of which is recognized by a particular tRNA. One end of the tRNA matches the genetic code in a three-nucleotide sequence called the anticodon. The anticodon forms three base pairs with a codon in mRNA during protein biosynthesis. On the other end of the tRNA is a covalent attachment to the amino acid that corresponds to the anticodon sequence. Each type of tRNA molecule can be attached to only one type of amino acid, so each organism has many types of tRNA. Because the genetic code contains multiple codons that specify the same amino acid, there are several tRNA molecules bearing different anticodons which carry the same amino acid.
The covalent attachment to the tRNA 3’ end is catalyzed by enzymes called aminoacyl tRNA synthetases. During protein synthesis, tRNAs with attached amino acids are delivered to the ribosome by proteins called elongation factors, which aid in association of the tRNA with the ribosome, synthesis of the new polypeptide and translocation (movement) of the ribosome along the mRNA. If the tRNA's anticodon matches the mRNA, another tRNA already bound to the ribosome transfers the growing polypeptide chain from its 3’ end to the amino acid attached to the 3’ end of the newly delivered tRNA, a reaction catalyzed by the ribosome.
A large number of the individual nucleotides in a tRNA molecule may be chemically modified, often by methylation or deamidation. These unusual bases sometimes affect the tRNA's interaction with ribosomes and sometimes occur in the anticodon to alter base-pairing properties.[4]:29.1.2
Structure[edit]
Secondary cloverleaf structure of tRNAPhe from yeast.
Tertiary structure of tRNA. CCA tail in yellow, Acceptor stem in purple, Variable loop in orange, D arm in red, Anticodon arm in blue with Anticodon in black, T arm in green.
The structure of tRNA can be decomposed into its primary structure, its secondary structure (usually visualized as the cloverleaf structure), and its tertiary structure[5] (all tRNAs have a similar L-shaped 3D structure that allows them to fit into the P and A sites of the ribosome). The cloverleaf structure becomes the 3D L-shaped structure through coaxial stacking of the helices, which is a common RNA tertiary structure motif.
The lengths of each arm, as well as the loop 'diameter', in a tRNA molecule vary from species to species.[5][6]
The tRNA structure consists of the following:
A 5'-terminal phosphate group.
The acceptor stem is a 7- to 9-base pair (bp) stem made by the base pairing of the 5'-terminal nucleotide with the 3'-terminal nucleotide (which contains the CCA 3'-terminal group used to attach the amino acid). In general, such 3'-terminal tRNA-like structures are referred to as 'genomic tags'. The acceptor stem may contain non-Watson-Crick base pairs.[5][7]
The CCA tail is a cytosine-cytosine-adenine sequence at the 3' end of the tRNA molecule. The amino acid loaded onto the tRNA by aminoacyl tRNA synthetases, to form aminoacyl-tRNA, is covalently bonded to the 3'-hydroxyl group on the CCA tail.[8] This sequence is important for the recognition of tRNA by enzymes and critical in translation.[9][10] In prokaryotes, the CCA sequence is transcribed in some tRNA sequences. In most prokaryotic tRNAs and eukaryotic tRNAs, the CCA sequence is added during processing and therefore does not appear in the tRNA gene.[11]
The D arm is a 4- to 6-bp stem ending in a loop that often contains dihydrouridine.[5]
The anticodon arm is a 5-bp stem whose loop contains the anticodon.[5] The tRNA 5'-to-3' primary structure contains the anticodon but in reverse order, since 3'-to-5' directionality is required to read the mRNA from 5'-to-3'.
The T arm is a 4- to 5- bp stem containing the sequence TΨC where Ψ is pseudouridine, a modified uridine.[5]
Bases that have been modified, especially by methylation (e.g. tRNA (guanine-N7-)-methyltransferase), occur in several positions throughout the tRNA. The first anticodon base, or wobble-position, is sometimes modified to inosine (derived from adenine), pseudouridine or lysidine (derived from cytosine).[12]
Anticodon[edit]
An anticodon[13] is a unit made up of three nucleotides that correspond to the three bases of the codon on the mRNA. Each tRNA contains a distinct anticodon triplet sequence that can base-pair to one or more codons for an amino acid. Some anticodons can pair with more than one codon due to a phenomenon known as wobble base pairing. Frequently, the first nucleotide of the anticodon is one not found on mRNA: inosine, which can hydrogen bond to more than one base in the corresponding codon position.[4]:29.3.9 In the genetic code, it is common for a single amino acid to be specified by all four third-position possibilities, or at least by both pyrimidines and purines; for example, the amino acid glycine is coded for by the codon sequences GGU, GGC, GGA, and GGG. Other modified nucleotides may also appear at the first anticodon position—sometimes known as the "wobble position"—resulting in subtle changes to the genetic code, as for example in mitochondria.[14]
To provide a one-to-one correspondence between tRNA molecules and codons that specify amino acids, 61 types of tRNA molecules would be required per cell, as there are 61 sense codons of the standard genetic code. However, many cells contain fewer than 61 types of tRNAs because the wobble base is capable of binding to several, though not necessarily all, of the codons that specify a particular amino acid. A minimum of 31 tRNAs are required to translate, unambiguously, all 61 sense codons. The maximum observed is 41.[3][15]
Aminoacylation[edit]
See also: Aminoacyl-tRNA
Aminoacylation is the process of adding an aminoacyl group to a compound. It covalently links an amino acid to the CCA 3' end of a tRNA molecule.
Each tRNA is aminoacylated (or charged) with a specific amino acid by an aminoacyl tRNA synthetase. There is normally a single aminoacyl tRNA synthetase for each amino acid, despite the fact that there can be more than one tRNA, and more than one anticodon, for an amino acid. Recognition of the appropriate tRNA by the synthetases is not mediated solely by the anticodon, and the acceptor stem often plays a prominent role.[16]
Reaction:
amino acid + ATP → aminoacyl-AMP + PPi
aminoacyl-AMP + tRNA → aminoacyl-tRNA + AMP
Certain organisms can have one or more aminoacyl tRNA synthetases missing. This leads to charging of the tRNA by a chemically related amino acid. An enzyme or enzymes modify the charged amino acid to the final one. For example, Helicobacter pylori has glutaminyl tRNA synthetase missing. Thus, glutamate tRNA synthetase charges tRNA-glutamine(tRNA-Gln) with glutamate. An amidotransferase then converts the acid side chain of the glutamate to the amide, forming the correctly charged gln-tRNA-Gln.
Binding to ribosome[edit]
Play media
The range of conformations adopted by tRNA as it transits the A/T through P/E sites on the ribosome. The Protein Data Bank (PDB) codes for the structural models used as end points of the animation are given. Both tRNAs are modeled as phenylalanine-specific tRNA from Escherichia coli, with the A/T tRNA as a homology model of the deposited coordinates. Color coding as shown for tRNA tertiary structure. Adapted from.[17]
The ribosome has three binding sites for tRNA molecules that span the space between the two ribosomal subunits: the A (aminoacyl),[18] P (peptidyl), and E (exit) sites. In addition, the ribosome has two other sites for tRNA binding that are used during mRNA decoding or during the initiation of protein synthesis. These are the T site (named elongation factor Tu) and I site (initiation).[19][20] By convention, the tRNA binding sites are denoted with the site on the small ribosomal subunit listed first and the site on the large ribosomal subunit listed second. For example, the A site is often written A/A, the P site, P/P, and the E site, E/E.[19] The binding proteins like L27, L2, L14, L15, L16 at the A- and P- sites have been determined by affinity labeling by A.P. Czernilofsky et al. (Proc. Natl. Acad. Sci, USA, pp 230–234, 1974).
Once translation initiation is complete, the first aminoacyl tRNA is located in the P/P site, ready for the elongation cycle described below. During translation elongation, tRNA first binds to the ribosome as part of a complex with elongation factor Tu (EF-Tu) or its eukaryotic (eEF-1) or archaeal counterpart. This initial tRNA binding site is called the A/T site. In the A/T site, the A-site half resides in the small ribosomal subunit where the mRNA decoding site is located. The mRNA decoding site is where the mRNA codon is read out during translation. The T-site half resides mainly on the large ribosomal subunit where EF-Tu or eEF-1 interacts with the ribosome. Once mRNA decoding is complete, the aminoacyl-tRNA is bound in the A/A site and is ready for the next peptide bond to be formed to its attached amino acid. The peptidyl-tRNA, which transfers the growing polypeptide to the aminoacyl-tRNA bound in the A/A site, is bound in the P/P site. Once the peptide bond is formed, the tRNA in the P/P site is deacylated, or has a free 3’ end, and the tRNA in the A/A site carries the growing polypeptide chain. To allow for the next elongation cycle, the tRNAs then move through hybrid A/P and P/E binding sites, before completing the cycle and residing in the P/P and E/E sites. Once the A/A and P/P tRNAs have moved to the P/P and E/E sites, the mRNA has also moved over by one codon and the A/T site is vacant, ready for the next round of mRNA decoding. The tRNA bound in the E/E site then leaves the ribosome.
The P/I site is actually the first to bind to aminoacyl tRNA, which is delivered by an initiation factor called IF2 in bacteria.[20] However, the existence of the P/I site in eukaryotic or archaeal ribosomes has not yet been confirmed. The P-site protein L27 has been determined by affinity labeling by E. Collatz and A.P. Czernilofsky (FEBS Lett., Vol. 63, pp 283–286, 1976).
tRNA genes[edit]
Organisms vary in the number of tRNA genes in their genome. The nematode worm C. elegans, a commonly used model organism in genetics studies, has 29,647 [21] genes in its nuclear genome, of which 620 code for tRNA.[22][23] The budding yeast Saccharomyces cerevisiae has 275 tRNA genes in its genome.
In the human genome, which, according to January 2013 estimates, has about 20,848 protein coding genes [24] in total, there are 497 nuclear genes encoding cytoplasmic tRNA molecules, and 324 tRNA-derived pseudogenes—tRNA genes thought to be no longer functional[25] (although pseudo tRNAs have been shown to be involved in antibiotic resistance in bacteria ).[26] Regions in nuclear chromosomes, very similar in sequence to mitochondrial tRNA genes, have also been identified (tRNA-lookalikes).[27] These tRNA-lookalikes are also considered part of the nuclear mitochondrial DNA (genes transferred from the mitochondria to the nucleus).[27][28]
As with all eukaryotes, there are 22 mitochondrial tRNA genes[29] in humans. Mutations in some of these genes have been associated with severe diseases like the MELAS syndrome.
Cytoplasmic tRNA genes can be grouped into 49 families according to their anticodon features. These genes are found on all chromosomes, except 22 and Y chromosome. High clustering on 6p is observed (140 tRNA genes), as well on 1 chromosome.[25]
The HGNC, in collaboration with the Genomic tRNA Database (GtRNAdb) and experts in the field, has approved unique names for human genes that encode tRNAs.
Evolution[edit]
The top half of tRNA (consisting of the D arm and the acceptor stem with 5'-terminal phosphate group and 3'-terminal CCA group) and the bottom half (consisting of the T arm and the anticodon arm) are independent units in structure as well as in function. The top half may have evolved first including the 3'-terminal genomic tag which originally may have marked tRNA-like molecules for replication in early RNA world. The bottom half may have evolved later as an expansion, e. g. as protein synthesis started in RNA world and turned it into a (ribonucleoprotein world (RNP world). This proposed scenario is called genomic tag hypothesis. In fact, tRNA and tRNA-like aggregates have an important catalytic influence (i. e. as ribozymes) on replication still today. These roles may be regarded as 'molecular (or chemical) fossiles' of RNA world.[30]
Genomic tRNA content is a differentiating feature of genomes among biological domains of life: Archaea present the simplest situation in terms of genomic tRNA content with a uniform number of gene copies, Bacteria have an intermediate situation and Eukarya present the most complex situation.[31] Eukarya present not only more tRNA gene content than the other two kingdoms but also a high variation in gene copy number among different isoacceptors, and this complexity seem to be due to duplications of tRNA genes and changes in anticodon specificity[citation needed].
Evolution of the tRNA gene copy number across different species has been linked to the appearance of specific tRNA modification enzymes (uridine methyltransferases in Bacteria, and adenosine deaminases in Eukarya), which increase the decoding capacity of a given tRNA.[31] As an example, tRNAAla encodes four different tRNA isoacceptors (AGC, UGC, GGC and CGC). In Eukarya, AGC isoacceptors are extremely enriched in gene copy number in comparison to the rest of isoacceptors, and this has been correlated with its A-to-I modification of its wobble base. This same trend has been shown for most amino acids of eukaryal species. Indeed, the effect of these two tRNA modifications is also seen in codon usage bias. Highly expressed genes seem to be enriched in codons that are exclusively using codons that will be decoded by these modified tRNAs, which suggests a possible role of these codons—and consequently of these tRNA modifications—in translation efficiency.[31]
tRNA-derived fragments[edit]
tRNA-derived fragments (or tRFs) are short molecules that emerge after cleavage of the mature tRNAs or the precursor transcript.[32][33][34][35] Both cytoplasmic and mitochondrial tRNAs can produce fragments.[36] There are at least four structural types of tRFs believed to originate from mature tRNAs, including the relatively long tRNA halves and short 5’-tRFs, 3’-tRFs and i-tRFs.[32][36][37] The precursor tRNA can be cleaved to produce molecules from the 5’ leader or 3’ trail sequences. Cleavage enzymes include Angiogenin, Dicer, RNase Z and RNase P.[32][33] Especially in the case of Angiogenin, the tRFs have a characteristically unusual cyclic phosphate at their 3’ end and a hydroxyl group at the 5’ end.[38]
tRFs have multiple dependencies and roles. They exhibit significant changes between sexes, among races and disease status.[36] Functionally, they can be loaded on Ago and act through RNAi pathways,[34][36][37][39] participate in the formation of stress granules,[40] displace mRNAs from RNA-binding proteins[41] or inhibit translation.[42] At the system or the organismal level, the four types of tRFs have a diverse spectrum of activities. Functionally, tRFs are associated with viral infection,[43] cancer,[36][37] cell proliferation [38] and also with epigenetic transgenerational regulation of metabolism.[44]
tRFs are not restricted to humans but have been shown to exist in multiple organisms.[37][45][46][47]
Two online tools are available for those wishing to learn more about tRFs: the framework for the interactive exploration of mitochondrial and nuclear tRNA fragments (MINTbase)[48] and the relational database of Transfer RNA related Fragments(tRFdb).[49] MINTbase also provides a naming scheme for the naming of tRFs called tRF-license plates that is genome independent.
tRNA biogenesis[edit]
In eukaryotic cells, tRNAs are transcribed by RNA polymerase III as pre-tRNAs in the nucleus.[50] RNA polymerase III recognizes two highly conserved downstream promoter sequences: the 5' intragenic control region (5'-ICR, D-control region, or A box), and the 3'-ICR (T-control region or B box) inside tRNA genes.[2][51][52] The first promoter begins at +8 of mature tRNAs and the second promoter is located 30-60 nucleotides downstream of the first promoter. The transcription terminates after a stretch of four or more thymidines.[2][52]
Bulge-helix-bulge motive of tRNA intron
Pre-tRNAs undergo extensive modifications inside the nucleus. Some pre-tRNAs contain introns that are spliced, or cut, to form the functional tRNA molecule;[53] in bacteria these self-splice, whereas in eukaryotes and archaea they are removed by tRNA-splicing endonucleases.[54] Eukaryotic pre-tRNA contains bulge-helix-bulge (BHB) structure motif that is important for recognition and precise splicing of tRNA intron by endonucleases.[55] This motif position and structure are evolutionary conserved. However, some organisms, such as unicellular algae have a non-canonical position of BHB-motif as well as 5'- and 3'-ends of the spliced intron sequence.[55] The 5' sequence is removed by RNase P,[56] whereas the 3' end is removed by the tRNase Z enzyme.[57] A notable exception is in the archaeon Nanoarchaeum equitans, which does not possess an RNase P enzyme and has a promoter placed such that transcription starts at the 5' end of the mature tRNA.[58] The non-templated 3' CCA tail is added by a nucleotidyl transferase.[59] Before tRNAs are exported into the cytoplasm by Los1/Xpo-t,[60][61] tRNAs are aminoacylated.[62] The order of the processing events is not conserved. For example, in yeast, the splicing is not carried out in the nucleus but at the cytoplasmic side of mitochondrial membranes.[63]
History[edit]
The existence of tRNA was first hypothesized by Francis Crick, based on the assumption that there must exist an adapter molecule capable of mediating the translation of the RNA alphabet into the protein alphabet. Significant research on structure was conducted in the early 1960s by Alex Rich and Don Caspar, two researchers in Boston, the Jacques Fresco group in Princeton University and a United Kingdom group at King's College London.[64] In 1965, Robert W. Holley of Cornell University reported the primary structure and suggested three secondary structures.[65] tRNA was first crystallized in Madison, Wisconsin, by Robert M. Bock.[66] The cloverleaf structure was ascertained by several other studies in the following years[67] and was finally confirmed using X-ray crystallography studies in 1974. Two independent groups, Kim Sung-Hou working under Alexander Rich and a British group headed by Aaron Klug, published the same crystallography findings within a year.[68][69]
See also[edit]
Cloverleaf model of tRNA
Kim Sung-Hou
Kissing stem-loop
mRNA
non-coding RNA and introns
Slippery sequence
tmRNA
Transfer RNA-like structures
Translation
tRNADB
Wobble hypothesis
Aminoacyl-tRNA
References[edit]
Jump up ^ Plescia, O J; Palczuk, N C; Cora-Figueroa, E; Mukherjee, A; Braun, W (October 1965). "Production of antibodies to soluble RNA (sRNA)". Proc. Natl. Acad. Sci. USA. 54 (4): 1281–1285. doi:10.1073/pnas.54.4.1281. PMC 219862  . PMID 5219832.
^ Jump up to: a b c Sharp, Stephen J; Schaack, Jerome; Cooley, Lynn; Burke, Deborah J; Soll, Dieter (1985). "Structure and Transcription of Eukaryotic tRNA Genes". CRC Critical Reviews in Biochemistry. 19 (2): 107–144. doi:10.3109/10409238509082541. PMID 3905254.
^ Jump up to: a b Crick F (1968). "The origin of the genetic code". J Mol Biol. 38 (3): 367–379. doi:10.1016/0022-2836(68)90392-6. PMID 4887876.
^ Jump up to: a b Stryer L, Berg JM, Tymoczko JL (2002). Biochemistry (5th ed.). San Francisco: W.H. Freeman. ISBN 0-7167-4955-6.
^ Jump up to: a b c d e f Itoh, Yuzuru; Sekine, Shun-ichi Sekine; Suetsugu, Shiro; Yokoyama, Shigeyuki (6 May 2013). "Tertiary structure of bacterial selenocysteine tRNA". Nucleic Acids Research. 41 (13): 6729–6738. doi:10.1093/nar/gkt321. PMC 3711452  . PMID 23649835. Retrieved 23 November 2014.
Jump up ^ Goodenbour, J. M.; Pan, T. (29 October 2006). "Diversity of tRNA genes in eukaryotes". Nucleic Acids Research. 34 (21): 6137–6146. doi:10.1093/nar/gkl725. PMC 1693877  . PMID 17088292. Retrieved 23 November 2014.
Jump up ^ Jahn, Martina; Rogers, M. John; Söll, Dieter (18 July 1991). "Anticodon and acceptor stem nucleotides in tRNAGln are major recognition elements for E. coli glutaminyl-tRNA synthetase". Nature. 352 (6332): 258–260. doi:10.1038/352258a0. PMID 1857423. Retrieved 23 November 2014.
Jump up ^ Ibba, Michael; Söll, Dieter (June 2000). "Aminoacyl-tRNA Synthesis". Annual Review of Biochemistry. 69 (1): 617–650. doi:10.1146/annurev.biochem.69.1.617. PMID 10966471. Retrieved 23 November 2014.
Jump up ^ Sprinzl, M., and Cramer, F. (1979) Prog. Nucleic Acids Res. Mol. Biol. 22, 1–16
Jump up ^ Green, R., and Noller, H. F. (1997) Annu. Rev. Biochem. 66, 679–716
Jump up ^ Aebi M, Kirchner G, Chen JY, et al. (September 1990). "Isolation of a temperature-sensitive mutant with an altered tRNA nucleotidyltransferase and cloning of the gene encoding tRNA nucleotidyltransferase in the yeast Saccharomyces cerevisiae". J. Biol. Chem. 265 (27): 16216–16220. PMID 2204621.
Jump up ^ McCloskey, James A.; Nishimura, Susumu (November 1977). "Modified nucleosides in transfer RNA". Accounts of Chemical Research. 10 (11): 403–410. doi:10.1021/ar50119a004. Retrieved 23 November 2014.
Jump up ^ Felsenfeld G, Cantoni G; Cantoni (1964). "Use of thermal denaturation studies to investigate the base sequence of yeast serine sRNA". Proc Natl Acad Sci USA. 51 (5): 818–26. Bibcode:1964PNAS...51..818F. doi:10.1073/pnas.51.5.818. PMC 300168  . PMID 14172997.
Jump up ^ Suzuki, T; Suzuki, T (June 2014). "A complete landscape of post-transcriptional modifications in mammalian mitochondrial tRNAs". Nucleic Acids Research. 42 (11): 7346–57. doi:10.1093/nar/gku390. PMC 4066797  . PMID 24831542.
Jump up ^ Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipursky SL, Darnell J. (2004). Molecular Biology of the Cell. WH Freeman: New York, NY. 5th ed.
Jump up ^ Schimmel P, Giege R, Moras D, Yokoyama S; Giege; Moras; Yokoyama (1993). "An operational RNA code for amino acids and possible relationship to genetic code". Proc. Natl. Acad. Sci. USA. 90 (19): 8763–876. Bibcode:1993PNAS...90.8763S. doi:10.1073/pnas.90.19.8763. CS1 maint: Multiple names: authors list (link)
Jump up ^ Dunkle JA, Wang L, Feldman MB, Pulk A, Chen VB, Kapral GJ, Noeske J, Richardson JS, Blanchard SC, Cate JH; Wang; Feldman; Pulk; Chen; Kapral; Noeske; Richardson; Blanchard; Cate (2011). "Structures of the bacterial ribosome in classical and hybrid states of tRNA binding". Science. 332 (6032): 981–984. Bibcode:2011Sci...332..981D. doi:10.1126/science.1202692. PMC 3176341  . PMID 21596992. CS1 maint: Multiple names: authors list (link)
Jump up ^ Konevega, AL; Soboleva, NG; Makhno, VI; Semenkov, YP; Wintermeyer, W; Rodnina, MV; Katunin, VI (Jan 2004). "Purine bases at position 37 of tRNA stabilize codon-anticodon interaction in the ribosomal A site by stacking and Mg2+-dependent interactions". RNA. 10 (1): 90–101. doi:10.1261/rna.5142404. PMC 1370521  . PMID 14681588.
^ Jump up to: a b Agirrezabala X, Frank J; Frank (2009). "Elongation in translation as a dynamic interaction among the ribosome, tRNA, and elongation factors EF-G and EF-Tu". Q Rev Biophys. 42 (3): 159–200. doi:10.1017/S0033583509990060. PMC 2832932  . PMID 20025795.
^ Jump up to: a b Allen GS, Zavialov A, Gursky R, Ehrenberg M, Frank J; Zavialov; Gursky; Ehrenberg; Frank (2005). "The cryo-EM structure of a translation initiation complex from Escherichia coli". Cell. 121 (5): 703–712. doi:10.1016/j.cell.2005.03.023. PMID 15935757. CS1 maint: Multiple names: authors list (link)
Jump up ^ WormBase web site, http://www.wormbase.org, release WS187, date 25-Jan-2008.
Jump up ^ Spieth, J; Lawson, D (Jan 2006). "Overview of gene structure". WormBook: 1–10. doi:10.1895/wormbook.1.65.1. PMID 18023127.
Jump up ^ Hartwell LH, Hood L, Goldberg ML, Reynolds AE, Silver LM, Veres RC. (2004). Genetics: From Genes to Genomes 2nd ed. McGraw-Hill: New York, NY. p 264.
Jump up ^ Ensembl release 70 - Jan 2013 http://www.ensembl.org/Homo_sapiens/Info/StatsTable?db=core
^ Jump up to: a b Lander E.; et al. (2001). "Initial sequencing and analysis of the human genome". Nature. 409 (6822): 860–921. doi:10.1038/35057062. PMID 11237011.
Jump up ^ Rogers Theresa E.; et al. (2012). "A Pseudo-tRNA Modulates Antibiotic Resistance in Bacillus cereus". PLoS ONE. 7 (7): e41248. doi:10.1371/journal.pone.0041248. PMC 3399842  . PMID 22815980.
^ Jump up to: a b Telonis Aristeidis G.; et al. (2014). "Nuclear and Mitochondrial tRNA-lookalikes in the Human Genome". Frontiers in Genetics. 5: 00344. doi:10.3389/fgene.2014.00344. PMC 4189335  . PMID 25339973.
Jump up ^ Ramos A.; et al. (2011). "Nuclear Insertions of Mitochondrial Origin: Database Updating and Usefulness in Cancer Studies". Mitochondrion. 11 (6): 946–53. doi:10.1016/j.mito.2011.08.009. PMID 21907832.
Jump up ^ Ibid. p 529.
Jump up ^ Nancy Maizels and Alan M. Weiner: The Genomic Tag Hypothesis - What Molecular Fossils Tell Us about the Evolution of tRNA, in: The RNA World, Second Edition © 1999 Cold Spring Harbor Laboratory Press ISBN 0-87969-561-7/99, PDF
^ Jump up to: a b c Novoa, Eva Maria; Pavon-Eternod, Mariana; Pan, Tao; Ribas de Pouplana, Lluís (March 2012). "A Role for tRNA Modifications in Genome Structure and Codon Usage". Cell. 149 (1): 202–213. doi:10.1016/j.cell.2012.01.050. PMID 22464330. Retrieved 23 November 2014.
^ Jump up to: a b c Gebetsberger Jennifer; et al. (2013). "Slicing tRNAs to boost functional ncRNA diversity". RNA biology. 10: 1798–1806. doi:10.4161/rna.27177. PMC 3917982  . PMID 24351723.
^ Jump up to: a b Shigematsu Megumi; et al. (2014). "Transfer RNA as a source of small functional RNA". Journal of Molecular Biology and Molecular Imaging. 1: 8.
^ Jump up to: a b Sobala Andrew; et al. (2011). "Transfer RNA-derived fragments: origins, processing, and functions". Wiley interdisciplinary reviews. 2: 853–862. doi:10.1002/wrna.96. PMID 21976287.
Jump up ^ Keam Simon P; et al. (2015). "tRNA-Derived Fragments (tRFs): Emerging New Roles for an Ancient RNA in the Regulation of Gene Expression". Life (Basel). 5: 1638–1651. doi:10.3390/life5041638. PMC 4695841  . PMID 26703738.
^ Jump up to: a b c d e Telonis Aristeidis G; et al. (2015). "Dissecting tRNA-derived fragment complexities using personalized transcriptomes reveals novel fragment classes and unexpected dependencies". Oncotarget. 6: 24797–822. doi:10.18632/oncotarget.4695. PMC 4694795  . PMID 26325506.
^ Jump up to: a b c d Kumar Pankaj; et al. (2014). "Meta-analysis of tRNA derived RNA fragments reveals that they are evolutionarily conserved and associate with AGO proteins to recognize specific RNA targets". BMC Biol. 12: 78. doi:10.1186/s12915-014-0078-0. PMC 4203973  . PMID 25270025.
^ Jump up to: a b Honda Shozo; et al. (2015). "Sex hormone-dependent tRNA halves enhance cell proliferation in breast and prostate cancers". Proc Natl Acad Sci USA. 112: E3816–25. doi:10.1073/pnas.1510077112. PMC 4517238  . PMID 26124144.
Jump up ^ Shigematsu Megumi; et al. (2015). "tRNA-Derived Short Non-coding RNA as Interacting Partners of Argonaute Proteins". Gene Regul Syst Bio. 9: 27–33. doi:10.4137/GRSB.S29411. PMC 4567038  . PMID 26401098.
Jump up ^ Emara Mohamed M; et al. (2010). "Angiogenin-induced tRNA-derived stress-induced RNAs promote stress-induced stress granule assembly". J Biol Chem. 285: 10959–68. doi:10.1074/jbc.M109.077560. PMC 2856301  . PMID 20129916.
Jump up ^ Goodarzi Hani; et al. (2015). "Endogenous tRNA-derived fragments suppress breast cancer progression via YBX1 displacement". Cell. 161: 790–802. doi:10.1016/j.cell.2015.02.053. PMC 4457382  . PMID 25957686.
Jump up ^ Ivanov Pavel; et al. (2011). "Angiogenin-induced tRNA fragments inhibit translation initiation". Mol Cell. 43: 613–23. doi:10.1016/j.molcel.2011.06.022. PMC 3160621  . PMID 21855800.
Jump up ^ Selitsky Sara R; et al. (2015). "Small tRNA-derived RNAs are increased and more abundant than microRNAs in chronic hepatitis B and C". Sci Rep. 5: 7675. doi:10.1038/srep07675. PMC 4286764  . PMID 25567797.
Jump up ^ Sharma Upasna; et al. (2016). "Biogenesis and function of tRNA fragments during sperm maturation and fertilization in mammals". Science. 351: 391–6. doi:10.1126/science.aad6780. PMC 4888079  . PMID 26721685.
Jump up ^ Casas Eduardo; et al. (2015). "Characterization of circulating transfer RNA-derived RNA fragments in cattle". Front Genet. 6: 271. doi:10.3389/fgene.2015.00271. PMC 4547532  . PMID 26379699.
Jump up ^ Hirose Yuka; et al. (2015). "Precise mapping and dynamics of tRNA-derived fragments (tRFs) in the development of Triops cancriformis (tadpole shrimp)". BMC Genet. 16: 83. doi:10.1186/s12863-015-0245-5. PMC 4501094  . PMID 26168920.
Jump up ^ Karaiskos Spyros; et al. (2015). "Age-driven modulation of tRNA-derived fragments in Drosophila and their potential targets". Biol Direct. 10: 51. doi:10.1186/s13062-015-0081-6. PMC 4572633  . PMID 26374501.
Jump up ^ Pliatsika Venetia; et al. (2016). "MINTbase: a framework for the interactive exploration of mitochondrial and nuclear tRNA fragments". Bioinformatics. doi:10.1093/bioinformatics/btw194.
Jump up ^ Kumar Panjav; et al. (2014). "tRFdb: a database for transfer RNA fragments". Nucleic Acids Res. 43: D141–D145. doi:10.1093/nar/gku1138. PMC 4383946  . PMID 25392422.
Jump up ^ White RJ (1997). "Regulation of RNA polymerases I and III by the retinoblastoma protein: a mechanism for growth control?". Trends in Biochemical Sciences. 22 (3): 77–80. doi:10.1016/S0968-0004(96)10067-0. PMID 9066256.
Jump up ^ Sharp, Stephen; Dingermann, Theodor; Söll, Dieter (1982). "The minimum intragenic sequences required for promotion of eukaryotic tRNA gene transcription" (PDF). Nucleic Acids Research. 10 (18): 5393–5406. doi:10.1093/nar/10.18.5393. PMC 320884  . PMID 6924209. Retrieved 23 November 2014.
^ Jump up to: a b Dieci G, Fiorino G, Castelnuovo M, Teichmann M, Pagano A; Fiorino; Castelnuovo; Teichmann; Pagano (December 2007). "The expanding RNA polymerase III transcriptome". Trends Genet. 23 (12): 614–22. doi:10.1016/j.tig.2007.09.001. PMID 17977614. CS1 maint: Multiple names: authors list (link)
Jump up ^ Tocchini-Valentini, Giuseppe D.; Fruscoloni, Paolo; Tocchini-Valentini, Glauco P. (12 November 2009). "Processing of multiple-intron-containing pretRNA". Proceedings of the National Academy of Sciences. 106 (48): 20246–20251. doi:10.1073/pnas.0911658106. PMC 2787110  . PMID 19910528.
Jump up ^ Abelson J, Trotta CR, Li H; Trotta; Li (1998). "tRNA Splicing". J Biol Chem. 273 (21): 12685–12688. doi:10.1074/jbc.273.21.12685. PMID 9582290. CS1 maint: Multiple names: authors list (link)
^ Jump up to: a b Soma, Akiko (2014). "Circularly permuted tRNA genes: their expression and implications for their physiological relevance and development". Frontiers in Genetics. 5. doi:10.3389/fgene.2014.00063. ISSN 1664-8021.
Jump up ^ Frank DN, Pace NR; Pace (1998). "Ribonuclease P: unity and diversity in a tRNA processing ribozyme". Annu. Rev. Biochem. 67 (1): 153–80. doi:10.1146/annurev.biochem.67.1.153. PMID 9759486.
Jump up ^ Ceballos M, Vioque A; Vioque (2007). "tRNase Z". Protein Pept. Lett. 14 (2): 137–45. doi:10.2174/092986607779816050. PMID 17305600.
Jump up ^ Randau L, Schröder I, Söll D; Schröder; Söll (May 2008). "Life without RNase P". Nature. 453 (7191): 120–3. Bibcode:2008Natur.453..120R. doi:10.1038/nature06833. PMID 18451863. CS1 maint: Multiple names: authors list (link)
Jump up ^ Weiner AM (October 2004). "tRNA maturation: RNA polymerization without a nucleic acid template". Curr. Biol. 14 (20): R883–5. doi:10.1016/j.cub.2004.09.069. PMID 15498478.
Jump up ^ Kutay, U. .; Lipowsky, G. .; Izaurralde, E. .; Bischoff, F. .; Schwarzmaier, P. .; Hartmann, E. .; Görlich, D. . (1998). "Identification of a tRNA-Specific Nuclear Export Receptor". Molecular Cell. 1 (3): 359–369. doi:10.1016/S1097-2765(00)80036-2. PMID 9660920.
Jump up ^ Arts, G. J.; Fornerod, M. .; Mattaj, L. W. (1998). "Identification of a nuclear export receptor for tRNA". Current Biology. 8 (6): 305–314. doi:10.1016/S0960-9822(98)70130-7. PMID 9512417.
Jump up ^ Arts, G. -J.; Kuersten, S.; Romby, P.; Ehresmann, B.; Mattaj, I. W. (1998). "The role of exportin-t in selective nuclear export of mature tRNAs". The EMBO Journal. 17 (24): 7430–7441. doi:10.1093/emboj/17.24.7430. PMC 1171087  . PMID 9857198.
Jump up ^ Yoshihisa, T.; Yunoki-Esaki, K.; Ohshima, C.; Tanaka, N.; Endo, T. (2003). "Possibility of cytoplasmic pre-tRNA splicing: the yeast tRNA splicing endonuclease mainly localizes on the mitochondria". Molecular Biology of the Cell. 14 (8): 3266–3279. doi:10.1091/mbc.E02-11-0757. PMC 181566  . PMID 12925762.
Jump up ^ Brian F.C. Clark (October 2006). "The crystal structure of tRNA" (PDF). J. Biosci. 31 (4): 453–7. doi:10.1007/BF02705184. PMID 17206065.
Jump up ^ HOLLEY RW; APGAR J; EVERETT GA; et al. (March 1965). "STRUCTURE OF A RIBONUCLEIC ACID". Science. 147 (3664): 1462–5. Bibcode:1965Sci...147.1462H. doi:10.1126/science.147.3664.1462. PMID 14263761. Retrieved 2010-09-03.
Jump up ^ https://www.nytimes.com/1991/07/04/obituaries/robert-m-bock-67-biologist-and-a-dean.html
Jump up ^ "The Nobel Prize in Physiology or Medicine 1968". Nobel Foundation. Retrieved 2007-07-28.
Jump up ^ Ladner JE; Jack A; Robertus JD; et al. (November 1975). "Structure of yeast phenylalanine transfer RNA at 2.5 A resolution". Proc. Natl. Acad. Sci. U.S.A. 72 (11): 4414–8. Bibcode:1975PNAS...72.4414L. doi:10.1073/pnas.72.11.4414. PMC 388732  . PMID 1105583.
Jump up ^ Kim SH; Quigley GJ; Suddath FL; et al. (1973). "Three-dimensional structure of yeast phenylalanine transfer RNA: folding of the polynucleotide chain". Science. 179 (4070): 285–8. Bibcode:1973Sci...179..285K. doi:10.1126/science.179.4070.285. PMID 4566654.
External links[edit]
Wikimedia Commons has media related to TRNA.
tRNAdb (updated and completely restructured version of Spritzls tRNA compilation)
original Sprinzl tRNA compilation
tRNA link to heart disease and stroke
GtRNAdb: Collection of tRNAs identified from complete genomes
HGNC: Gene nomenclature of human tRNAs
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see also mitochondrial diseases
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values: حمض نووي ريبوزي ناقل – Arabic

values: Транспортна РНК – Bulgarian

values:
https://cs.wikipedia.org/wiki/TRNA
TRNA – Czech
cs
cs
interlanguage-link-target

values:
https://da.wikipedia.org/wiki/TRNA
TRNA – Danish
da
da
interlanguage-link-target

values:
https://de.wikipedia.org/wiki/TRNA
TRNA – German
de
de
interlanguage-link-target

values: Μεταφορικό RNA – Greek

values: آران‌ای حامل – Persian

values: Acide ribonucléique de transfert – French

values: Փոխադրող ՌՆԹ – Armenian

values:
https://is.wikipedia.org/wiki/TRNA
TRNA – Icelandic
is
is
interlanguage-link-target

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https://he.wikipedia.org/wiki/TRNA
TRNA – Hebrew
he
he
interlanguage-link-target

values: სატრანსპორტო რიბონუკლეინის მჟავა – Georgian

values: Тасымалдаушы РНҚ – Kazakh

values: Ташуучу РНК – Kyrgyz

values:
https://la.wikipedia.org/wiki/TRNA
TRNA – Latin
la
la
interlanguage-link-target

values:
https://lt.wikipedia.org/wiki/TRNR
TRNR – Lithuanian
lt
lt
interlanguage-link-target

values:
https://nn.wikipedia.org/wiki/TRNA
TRNA – Norwegian Nynorsk
nn
nn
interlanguage-link-target

values: Acid ribonucleïc de transferiment – Occitan

values:
https://pl.wikipedia.org/wiki/TRNA
TRNA – Polish
pl
pl
interlanguage-link-target

values: Transferová ribonukleová kyselina – Slovak

values: ئاڕ ئێن ئەی گوێزەرەوە – Central Kurdish

values: Транспортна РНК – Serbian

values: ทีอาร์เอ็นเอ – Thai

values: Транспортна рибонуклеїнова кислота – Ukrainian

values: 转运核糖核酸 – Chinese

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