text:
Midnight sun
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For other uses, see Midnight sun (disambiguation).
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The Altafjord in Alta, Norway, bathed in the midnight sun.
Midnight sun at the North Cape on the island of Magerøya in Norway
The midnight sun is a natural phenomenon that occurs in the summer months in places north of the Arctic Circle or south of the Antarctic Circle, when the sun remains visible at the local midnight.
Contents
[hide]
1 Details
2 Time zones and daylight saving time
3 White nights
4 Duration
5 See also
6 References
7 Further reading
8 External links
Details[edit]
Around the summer solstice (approximately 21 June in the Northern Hemisphere and 22 December in the Southern Hemisphere), the sun is visible for the full 24 hours, given fair weather. The number of days per year with potential midnight sun increases the closer towards either pole one goes. Although approximately defined by the polar circles, in practice the midnight sun can be seen as much as 55 miles (90 km) outside the polar circle, as described below, and the exact latitudes of the farthest reaches of midnight sun depend on topography and vary slightly year-to-year.
Midnight Sun seen from airplane while passing Greenland
Because there are no permanent human settlements south of the Antarctic Circle, apart from research stations, the countries and territories whose populations experience the midnight sun are limited to those crossed by the Arctic Circle: the Canadian Yukon, Nunavut, and Northwest Territories, and the nations of Iceland, Finland, Norway, Sweden, Denmark (Greenland), Russia, and the State of Alaska in the United States. A quarter of Finland's territory lies north of the Arctic Circle, and at the country's northernmost point the sun does not set at all for 60 days during summer. In Svalbard, Norway, the northernmost inhabited region of Europe, there is no sunset from approximately 19 April to 23 August. The extreme sites are the poles, where the sun can be continuously visible for half the year. The North Pole has midnight sun for 6 months from late March to late September.
The opposite phenomenon, polar night, occurs in winter, when the sun stays below the horizon throughout the day.
Since the axial tilt of the Earth is considerable (approximately 23 degrees 27 minutes), the sun does not set at high latitudes in local summer.[1] The sun remains continuously visible for one day during the summer solstice at the polar circle, for several weeks only 100 km (62 mi) closer to the pole, and for six months at the pole. At extreme latitudes, the midnight sun is usually referred to as polar day.
At the poles themselves, the sun rises and sets only once each year on the equinox. During the six months that the sun is above the horizon, it spends the days continuously moving in circles around the observer, gradually spiralling higher and reaching its highest circuit of the sky at the summer solstice.
Because of atmospheric refraction, and also because the sun is a disc rather than a point, the midnight sun may be experienced at latitudes slightly below the polar circle, though not exceeding one degree (depending on local conditions). For example, Iceland is known for its midnight sun, even though most of it (Grímsey is the exception) is slightly south of the Arctic Circle. For the same reasons, the period of sunlight at the poles is slightly longer than six months. Even the northern extremities of Scotland (and places at similar latitudes, such as St. Petersburg) experience twilight throughout the night in the northern sky at around the summer solstice.
Observers at heights appreciably above sea level can experience extended periods of midnight sun as a result of the "dip" of the horizon viewed from altitude.
Time zones and daylight saving time[edit]
The term "midnight sun" refers to the consecutive 24-hour periods of sunlight experienced north of the Arctic Circle and south of the Antarctic Circle. Other phenomena are sometimes referred to as "midnight sun", but they are caused by time zones and the observance of daylight saving time. For instance, in Fairbanks, Alaska, which is south of the Arctic Circle, the sun sets at 12:47 am at the summer solstice. This is because Fairbanks is 51 minutes ahead of its idealized time zone (as most of the state is in one time zone) and Alaska observes daylight saving time. (Fairbanks is at about 147.72 degrees west, corresponding to UTC−9 hours 51 minutes, and is on UTC−9 in winter.) This means that solar culmination occurs at about 12:51 pm instead of at 12 noon.
If a precise moment for the genuine "midnight sun" is required, the observer's longitude, the local civil time and the equation of time must be taken into account. The moment of the sun's closest approach to the horizon coincides with its passing due north at the observer's position, which occurs only approximately at midnight in general. Each degree of longitude east of the Greenwich meridian makes the vital moment exactly 4 minutes earlier than midnight as shown on the clock, while each hour that the local civil time is ahead of coordinated universal time (UTC, also known as GMT) makes the moment an hour later. These two effects must be added. Furthermore, the equation of time (which depends on the date) must be added: a positive value on a given date means that the sun is running slightly ahead of its average position, so the value must be subtracted.[2]
As an example, at the North Cape of Norway at midnight on June 21/22, the longitude of 25.9 degrees east makes the moment 103.2 minutes earlier by clock time; but the local time, 2 hours ahead of GMT in the summer, makes it 120 minutes later by clock time. The equation of time at that date is -2.0 minutes. Therefore, the sun's lowest elevation occurs 120 - 103.2 + 2.0 minutes after midnight: at 00.19 Central European Summer time. On other nearby dates the only thing different is the equation of time, so this remains a reasonable estimate for a considerable period. The sun's altitude remains within half a degree of the minimum of about 5 degrees for about 45 minutes either side of this time.
White nights[edit]
Main article: White Night festivals
Locations where the sun remains less than 6 (or 7[3]) degrees below the horizon—above 60° 34’ (or 59° 34’) latitude south of the Arctic Circle or north of the Antarctic Circle—experience midnight twilight instead of midnight sun, so that daytime activities, such as reading, are still possible without artificial light on a clear night.
White Nights have become a common symbol of Saint Petersburg, Russia, where they occur from about 11 June to 2 July,[3] and the last 10 days of June are celebrated with cultural events known as the White Nights Festival.
Duration[edit]
Map showing the dates of midnight sun at various latitudes (left) and the total number of nights.
The midnight sun is visible at the Arctic Circle from 12 June until 1 July. This period extends as one travels north: At Cape Nordkinn, Norway, the northernmost point of Continental Europe, the midnight sun lasts approximately from 14 May to 29 July. On the Svalbard archipelago farther north, it lasts from 20 April to 22 August.[4]
See also[edit]
Environment portal
Pytheas, ancient Greek geographer from Massalia and first person to describe the midnight sun
Eagle Summit, which, because of its altitude, experiences the midnight sun despite being south of the Arctic Circle
Polar night, the opposite phenomenon experienced in winter: a day without sunrise.
Midnight Sun Solar Race Team: With the midnight sun phenomenon, a solar-powered vehicle can continue driving 24 hours a day.
References[edit]
Jump up ^ "What is the Midnight Sun Phenomenon? | Earth Phenomena | Planetary Science". Scribd. Retrieved 2017-08-25.
Jump up ^ H. Spencer Jones, General Astronomy (Edward Arnold, London, 1922), Chapters I-III
^ Jump up to: a b Great Soviet Encyclopedia
Jump up ^ Trygve B. Haugan, ed. Det Nordlige Norge Fra Trondheim Til Midnattssolens Land (Trondheim: Reisetrafikkforeningen for Trondheim og Trøndelag. 1940)
Further reading[edit]
Lutgens F.K., Tarbuck E.J. (2007) The Atmosphere, Tenth Edition, page 39, PEARSON, Prentice Hall, NJ.
External links[edit]
Wikimedia Commons has media related to Midnight sun.
Midnight sun seen from Fjellheisen Tromsø - 360 panorama
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Parts of a day
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Midnight sun
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Gotham by Gaslight
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Gotham by Gaslight
Gotham by Gaslight (February 1989). Art by Mike Mignola.
Publication information
Publisher
DC Comics
Format
One-shot
Genre
Steampunk, superhero
Publication date
February 1989
No. of issues
1
Main character(s)
Batman
Jack the Ripper
Creative team
Written by
Brian Augustyn
Artist(s)
Mike Mignola
Inker(s)
P. Craig Russell
Editor(s)
Mark Waid
Collected editions
Gotham by Gaslight
ISBN 0-930289-67-6
Gotham by Gaslight (inc. Master of the Future)
ISBN 1-4012-1153-4
Gotham by Gaslight is a DC Comics one-shot by Brian Augustyn and Mike Mignola, with inks by P. Craig Russell. It spawned a sequel, Master of the Future (1991), also written by Augustyn, with art by Eduardo Barreto.
Gotham by Gaslight is considered to be the first Elseworlds story, where DC Comics heroes are taken out of their usual setting and put into alternate timelines or realities. And although it was not originally labelled as such, subsequent printings include the "Elseworlds" logo.
Contents
[hide]
1 Plot Summaries
1.1 Gotham by Gaslight
1.2 Master of the Future
2 Publication history
2.1 Collected editions
3 Reception
4 Spin-offs
5 In other media
5.1 Film
5.2 Video games
6 See also
7 References
8 External links
Plot Summaries[edit]
Gotham by Gaslight[edit]
In 1889, Bruce Wayne is on a tour of Europe which ends in Vienna. There he is studying under Dr. Freud. Bruce tells Freud that he has a recurring dream in which he recalls the murder of his parents and decides he must return to Gotham City. On the return trip, Bruce meets Jacob Packer, an old family friend whom he calls Uncle Jake, who also returned from a trip to Europe.
Shortly after arriving, Inspector Gordon informs Wayne about criminal gangs currently operating in Gotham. Gordon also shows Bruce the case of a man who poisoned his wife and tried to commit suicide with the poison, which left him alive with a permanent grin. Bruce takes up the mantle of Batman to fight criminals on the street.
At the same time, a series of murders of women take place and some people begin to suspect that Batman is the murderer. It is soon discovered that Jack the Ripper has come to Gotham, as the murders in Gotham City seem to resemble the Ripper murders.
After a search of Wayne Manor, a bloody knife is found under Bruce's bed and Bruce is arrested. A trial is held in which Uncle Jake is Bruce's defense attorney. After the trial, Bruce is convicted of being the Ripper and sentenced to be hanged for his crimes. Bruce is imprisoned in Arkham Asylum.
Once Bruce is in prison, Gordon gives him all the documentation on the crimes. Bruce toils day and night to try to figure out how he can get the Ripper. Just one day before the execution, Bruce learns the identity of the Ripper by discovering he had the skill of a surgeon and used a knife that belonged to the medical group who worked with his father. Bruce escapes from prison with the help of Alfred and heads straight for the Ripper.
Batman interrupts the Ripper as he is about to claim his next victim. Batman chases the Ripper throughout Gotham and the two eventually come to a stop at the grave of Thomas and Martha Wayne, where it is revealed that Jacob Packer is the Ripper. Packer had been trained in medicine and law with the money of Thomas Wayne, but he was driven insane by Martha Wayne's rejection of his advances. Since then he was killing women who resemble Martha to silence the laughter of Martha he heard in his head.
Packer reveals that he hired an assassin to kill Bruce's parents. Gordon appears at this time with the police and Batman tells them to arrest Packer. Packer confesses that he is the Ripper and tries to kill Batman, but Gordon shoots Packer dead at the last minute. Batman disappears into the shadows, leaving Gordon to take in the body of Jack the Ripper.
Master of the Future[edit]
In 1892, three years later, Bruce has "retired" his alter-ego and is engaged to be married. Having brought his parents' murderer to justice, he has come to regard being Batman as a childish fancy, and intends to devote his life to more serious pursuits. Others believe that Gotham City needs Batman more than ever, namely now-Commissioner Gordon and Bruce's own fiancée, Julie (whom Batman rescued from a late-night assault).
At a City Council meeting, Mayor Tolliver (the former Police Commissioner who aggressively prosecuted Bruce as the suspected Ripper) is promoting Gotham's hosting of the "American Discovery Exposition" to market Gotham as the "City of the Future". The meeting is interrupted by the flamboyant Alexandre LeRoi, who demands that he be proclaimed master of the city, or else he will burn it to the ground. He leaps out the window before he can be arrested, and Tolliver insists that the fair proceed.
While the fairgrounds are being erected, a Maxim Gun on an automated carriage rolls in and opens fire. Bruce, in attendance, pushes Tolliver out of the line of fire and disables the machine.
On the opening day of the fair, LeRoi kidnaps Tolliver from his mansion and takes him aboard an airship, forcing him to watch as LeRoi focuses a giant burning glass on the fair's main pavilion, before LeRoi throws Tolliver over the side to his death. Bruce is desperate to act, when Alfred arrives with his costume.
With the police fully occupied, Julie runs into the burning pavilion to rescue a small girl, but both of them are trapped by falling debris. Batman saves them, then fashions an improvised glider and takes off after LeRoi's airship. The two men engage in a brutal rapier duel that disables LeRoi's robotic pilot "Antonio", causing the airship to drift off course and become unstable. Noticing this, Batman urges LeRoi to surrender and abandon ship, but LeRoi refuses to believe him. Batman jumps overboard, but LeRoi is trapped as the airship crashes into the ocean and explodes. More than a hundred people are killed by LeRoi's fires, but further fatalities and destruction are prevented by the Gotham Police and firefighters. LeRoi is presumed dead, although he is shown to have survived the crash.
Batman confronts Councilman Franklin Claypool, who confesses to being LeRoi's accomplice and providing him with information for his attack. Claypool secretly owns a large portion of Gotham City's slum housing, which he expected to sell profitably through his proposed redevelopment program, but the program was pushed aside by Tolliver in favor of the fair. LeRoi agreed to target Claypool's properties, allowing Claypool to collect a lucrative insurance payout, but Claypool never intended the deaths LeRoi's actions caused, and so does not resist when Gordon arrives to arrest him.
Bruce and Julie are walking along a hilltop overlooking the city, when she reveals that she recognized him at the fair, even through his mask. Apprehensively, he asks her what she plans to do, and she replies, what she has always done: continue to love and encourage him, especially now that they both agree that Gotham still needs Batman.
Publication history[edit]
Gotham by Gaslight, which retroactively became the first official Elseworlds publication, was initially released as a 52-page one-shot in February 1989. It was written by Brian Augustyn and pencilled by Mike Mignola, with inks by P. Craig Russell. Two years later, the 68-page sequel Batman: Master of the Future — also written by Augustyn, but with art by Eduardo Barreto — was released, this time officially labelled an Elseworlds publication.
In 2006, in the wake of the DC event Infinite Crisis (which reshuffled the DC Multiverse), the Gotham by Gaslight world was identified as "Earth-19".
Prior to its appearance in Countdown Presents: The Search for Ray Palmer: Gotham by Gaslight (January 2008), the two one-shot issues were collected into one 112-page trade paperback volume (which was released under the same name, and with the same Mignola cover as the one-shot release).
Collected editions[edit]
The two stories have been collected into a trade paperback.
Gotham by Gaslight, August 2006, DC Comics, ISBN 1-4012-1153-4 (Titan Books, ISBN 1-84576-403-X), collects:
Gotham by Gaslight, 48 pages, DC Comics, November 1989, ISBN 0-930289-67-6 (Titan Books, December 1989, ISBN 1-85286-265-3)
Master of the Future, 63 pages, DC Comics, December 1991, ISBN 1-56389-015-1
Reception[edit]
IGN Comics ranked Gotham by Gaslight #11 on a list of the 25 greatest Batman graphic novels, saying that the comic is "as taut and well-conceived a graphic novel as you can find — Elseworlds or otherwise," they add, "quite simply, no other Elseworlds tale has managed such a brilliant concept nor executed it so perfectly."
Gotham by Gaslight was not the highest-ranking Elseworlds story on the list, as Batman & Dracula: Red Rain ranked higher at #9.[1]
Spin-offs[edit]
Main article: Countdown to Final Crisis
After the events of Identity Crisis, the Atom spent DC's missing year out of the limelight. In the weekly-sequel to the weekly event 52, Countdown, the Source Wall names Palmer as a key figure in impending DC Universe events. The search for Ray Palmer began in the Wildstorm Universe, and soon crossed onto the new 52 worlds created during the course of Infinite Crisis.
The first of these was called Countdown Presents: The Search for Ray Palmer: Gotham by Gaslight (January 2008) and was written by Gotham by Gaslight's original author, Brian Augustyn, relishing his chance to return to the world he had created.[2] The story saw Bob the Monitor accompany Jason Todd, Donna Troy, and Kyle Rayner (the latter three inhabitants of "New Earth") through the multiverse in search of multiuniversal counterparts to Ray Palmer. The group arrived on Earth-19, the Earth where the Batman of the earlier two Elseworlds storylines acted, there, they also encountered this Earth's counterpart of Blue Beetle and Man-Bat before they went to the next Earth for their continuing search. Their search would take them to worlds populated by characters from other Elseworlds stories, including those seen in the Batman & Dracula trilogy (Earth-43) and Superman: Red Son (Earth-30).
In other media[edit]
Film[edit]
Batman: Gotham by Gaslight is an animated adaptation released in 2018.[3] It features Bruce Greenwood as Batman and Jennifer Carpenter as Selina Kyle. The adaptation differs significantly from the original comic, and includes elements from both books.
Video games[edit]
A video game based on the comic book was planned for release by Day 1 Studios (who also developed Fear 3). After THQ failed to secure the rights to the property, the game was canceled.[4] The developers released some concept illustration which revealed the game had a steampunk feel to it while being set in Victorian England and also test footage was released which revealed the misty Gothic settings and the character model for Batman. Reception to the leaked footage and concept illustration was positive, with viewers praising the setting, the cape physics and the different approach the developers were trying to achieve.[5]
Season pass holders for Batman: Arkham Origins, released in 2013, were awarded a special costume for Batman that was based on Gotham By Gaslight.[6]
The Gotham by Gaslight version of Batman was also introduced as a playable character in an update to the mobile version of Injustice: Gods Among Us in early 2018.[7]
See also[edit]
List of Elseworlds publications
List of steampunk works
From Hell, Alan Moore's Ripper-based graphic novel.
The League of Extraordinary Gentlemen, Moore's Victorian-era tales of derring-do.
Robur the Conqueror, a character created by Jules Verne that LeRoi appears to have been based on.
Other Elseworlds that involve Jack the Ripper:
Batman: Two Faces
JLA: The Island of Dr. Moreau
Wonder Woman: Amazonia
References[edit]
Jump up ^ Yehl, Josuha; Goldstein, Hilary (9 April 2014). "The 25 Greatest Batman Graphic Novels". IGN. Retrieved 25 August 2017.
Jump up ^ Contino, Jennifer M. (November 12, 2007). "Augustyn by Gaslight! Brian returns to Gotham". ComicCon.com. Archived from the original on March 2, 2008. Retrieved 2008-09-03.
Jump up ^ Lovett, Jamie (7 August 2017). "DC Animation's Next Movie Revealed". Comic Book.com. Retrieved 25 August 2017.
Jump up ^ "Gotham By Gaslight Video Shows Canceled Elseworlds Batman Game". February 27, 2012. Retrieved February 27, 2012.
Jump up ^ Cummins, Chris (February 29, 2012). "This Gotham by Gaslight Game Looks Great, Doesn't Exist". Toplessrobot.com. Retrieved March 17, 2012.
Jump up ^ Sarkar, Samit. "Batman: Arkham Origins season pass available for $19.99". Polygon. Retrieved 1 February 2018.
Jump up ^ Musgrave, Shawn. "Best iPhone Game Updates: 'Vainglory', 'Marvel Avengers Academy', 'Portal Knights', 'Dashy Crashy', and More". TouchArcade. Retrieved 25 March 2018.
External links[edit]
Gotham by Gaslight trade paperback details at DC Comics.com
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[show]
v
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Jack the Ripper in fiction
Seminal works
The Lodger
Jack the Ripper: The Final Solution
Letters
"Dear Boss" letter
"From Hell" letter
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Film
Waxworks (1924)
The Lodger: A Story of the London Fog (1926)
Pandora's Box (1929)
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Room to Let (1950)
Man in the Attic (1953)
Jack the Ripper (1959)
Lulu (1962)
A Study in Terror (1965)
Hands of the Ripper (1971)
Dr. Jekyll and Sister Hyde (1971)
The Ruling Class (1972)
What the Swedish Butler Saw (1975)
Jack the Ripper (1976)
Murder by Decree (1979)
Time After Time (1979)
Jack's Back (1988)
Edge of Sanity (1989)
Deadly Advice (1994)
Ripper (2001)
From Hell (2001)
Bad Karma (2002)
Case Closed: The Phantom of Baker Street (2002)
The Lodger (2009)
Holmes & Watson. Madrid Days (2012)
Batman: Gotham by Gaslight (2018)
Parody
Bizarre, Bizarre (1937)
Amazon Women on the Moon (1987)
Music
"Jack the Ripper" (1963)
"The Ripper" (1976)
"Killer on the Loose" (1980)
The Somatic Defilement (2007)
Stage
Earth Spirit (1895 play)
Pandora's Box (1904 play)
Lulu (1937 opera)
The Lodger (1960 opera)
The Ruling Class (1968 play)
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From Hell
Blood of the Innocent
Gotham by Gaslight
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Martin Mystère
The League of Extraordinary Gentlemen, Volume III: Century
Literature
Sherlock Holmes
The Last Sherlock Holmes Story
The Whitechapel Horrors
Sherlock Holmes: The Unauthorized Biography
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The Ripper Legacy
Short stories
"A Toy for Juliette"
"The Prowler in the City at the Edge of the World"
Other
A Feast Unknown
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I, Ripper
Matrix
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Jack the Ripper, Light-Hearted Friend
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Television
Series
Jack the Ripper (1973)
The Phantom Raspberry Blower of Old London Town (1976)
Jack the Ripper (1988)
Sanctuary (2007–2011)
Whitechapel (2009–2013)
Ripper Street (2012–2017)
Time After Time (2017)
Episodes
"Wolf in the Fold" (1967)
"Comes the Inquisitor" (1995)
"Ripper" (1999)
"Sanctuary for All" (2008)
Other
Bridge Across Time (1985 TV film)
Video games
Jack the Ripper (1987)
Ripper (1996)
Duke Nukem: Zero Hour (1999)
Shadow Man (1999)
MediEvil 2 (2000)
Jack the Ripper (2003)
Mystery in London: On the Trail of Jack the Ripper (2007)
Sherlock Holmes Versus Jack the Ripper (2009)
The Ripper (canceled)
Jack the Ripper (2015)
Other
Casebook: Jack the Ripper
Blood!: The Life and Future Times of Jack the Ripper
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TATA box
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Figure 1. TATA box structural elements. The TATA box consensus sequence is TATAWAW, where W is either A or T.
In molecular biology, the TATA box (also called the Goldberg-Hogness box)[1] is a sequence of DNA found in the core promotor region of genes in archaea and eukaryotes.[2] The prokaryotic homolog of the TATA box is called the Pribnow box which has a shorter consensus sequence.
The TATA box is considered a non-coding DNA sequence (also known as a cis-regulatory element). It was termed the "TATA box" as it contains a consensus sequence characterized by repeating T and A base pairs.[3] How the term "box" originated is unclear. In the 1980s, while investigating nucleotide sequences in mouse genome loci, the Hogness box sequence was found and "boxed in" at the -31 position.[4] When consensus nucleotides and alternative ones were compared, homologous regions were "boxed" by the researchers.[4] The boxing in of sequences sheds light on the origin of the term "box".
The TATA box was first identified in 1978[1] as a component of eukaryotic promoters. Transcription is initiated at the TATA box in TATA-containing genes. The TATA box is the binding site of the TATA-binding protein (TBP) and transcription factors in some eukaryotic genes. Gene transcription by RNA polymerase II depends on the regulation of the core promoter by long-range regulatory elements such as enhancers and silencers.[5] Without proper regulation of transcription, eukaryotic organisms would not be able to properly respond to their environment.
Based on the sequence and mechanism of TATA box initiation, mutations such as insertions, deletions, and point mutations to this consensus sequence can result in phenotypic changes. These phenotypic changes can then turn into a disease phenotype. Some diseases associated with mutations in the TATA box include gastric cancer, spinocerebellar ataxia, Huntington's disease, blindness, B-thalassemia, immunosuppression, Gilbert's syndrome, and HIV-1. The TATA-binding protein (TBP) could also be targeted by viruses as a means of viral transcription.[6]
Contents
[hide]
1 History
1.1 Discovery
1.2 Evolutionary History
2 Features
2.1 Location
2.2 Analogous Sequences
3 Structure
3.1 Sequence and Prevalence
4 Function
4.1 Role in Transcription Initiation
4.2 Interactions
5 Mutations
5.1 Insertions or Deletions
5.2 Point Mutations
5.3 Diseases
6 Clinical Significance
6.1 Technology
6.2 Cancer Therapy
7 Genetic engineering
7.1 TATA box modification
8 See also
9 References
History[edit]
Discovery[edit]
The TATA box was the first eukaryotic core promoter motif to be identified in 1978 by American biochemist David Hogness[1] while he and his graduate student, Michael Goldberg where on sabbatical at the University of Basel in Switzerland.[7] They first discovered the TATA sequence while analyzing 5' DNA promoter sequences in Drosophila, mammalian, and viral genes.[8][2] The TATA box was found in protein coding genes transcribed by RNA polymerase II.[2]
Evolutionary History[edit]
Most research on the TATA box has been conducted on yeast, human, and Drosophila genomes, however, similar elements have been found in archaea and ancient eukaryotes.[2] In archaea species, the promoter contains an 8 bp AT-rich sequence located ~24 bp upstream of the transcription start site. This sequence was originally called Box A, which is now known to be the sequence that interacts with the homologue of the archaeal TATA-binding protein (TBP). Also, even though some studies have uncovered several similarities, there are others that have detected notable differences between archaeal and eukaryotic TBP. The archaea protein exhibits a greater symmetry in its primary sequence and in the distribution of electrostatic charge, which is important because the higher symmetry lowers the protein's ability to bind the TATA box in a polar manner.[2]
Even though the TATA box is present in many eukaryotic promoters, is important to note that is not contained in the majority of promoters. One study found less than 30% of 1031 potential promoter regions contain a putative TATA box motif in humans.[9] In Drosophila, less than 40% of 205 core promoters contain a TATA box.[8] When there is an absence of the TATA box and TBP is not present, the downstream promoter element (DPE) in cooperation with the initiator element (Inr) bind to the transcription factor II D (TFIID), initiating transcription in TATA-less promoters. The DPE has been identified in three Drosophila TATA-less promoters and in the TATA-less human IRF-1 promoter.[10]
Features[edit]
Location[edit]
Promoter sequences vary between prokaryotes and eukaryotes. In eukaryotes, the TATA box is located 25 base pairs upstream of the start site that Rpb4/Rbp7 use to initiate transcription. In metazoans, the TATA box is located 30 base pairs upstream of the transcription start site.[5] While in yeast, S. cerevisiae, the TATA box has a variable position which can range from 40 to 100 bp upstream of the start site. The TATA box is also found in 40% of the core promoters of genes that code for the actin cytoskeleton and contractile apparatus in cells.[5]
The type of core promoter affects the level of transcription and expression of a gene. TATA-binding protein (TBP) can be recruited in two ways, by SAGA, a cofactor for RNA polymerase II, or by TFIID.[11] When promoters use the SAGA/TATA box complex to recruit RNA polymerase II, they are more highly regulated and display higher expression levels than promoters using the TFIID/TBP mode of recruitment.[11]
Analogous Sequences[edit]
In prokaryotes, promoter regions may contain a Pribnow box, which serves an analogous purpose to the eukaryotic TATA box. The Pribnow box has a 6 bp region centered around the -10 position and a 8-12 bp sequence around the -35 region that are both conserved.[10]
A CAAT box (also CAT box) is a region of nucleotides with the following consensus sequence: 5’ GGCCAATCT 3’. The CAAT box is located about 75-80 bases upstream of the transcription initiation site and about 150 bases upstream of the TATA box. It binds transcription factors (CAAT TF or CTFs) and thereby stabilizes the nearby preinitiation complex for easier binding of RNA polymerases. CAAT boxes are rarely found in genes that express proteins ubiquitous in all cell types.[10]
Structure[edit]
Sequence and Prevalence[edit]
Figure 2. Mechanism for transcription initiation at the TATA box. Transcription factors, TATA binding protein (TBP), and RNA polymerase II are all recruited to begin transcription.
The TATA box is a component of the eukaryotic core promoter and generally contains the consensus sequence 5'-TATA(A/T)A(A/T)-3'.[3] In yeast, for example, one study found that various Saccharomyces genomes had the consensus sequence 5'-TATA(A/T)A(A/T)(A/G)-3', yet only about 20% of yeast genes even contained the TATA sequence.[12] Similarly, in humans only 24% of genes have promoter regions containing the TATA box.[13] Genes containing the TATA-box tend to be involved in stress-responses and certain types of metabolism and are more highly regulated when compared to TATA-less genes.[12][14] Generally, TATA-containing genes are not involved in essential cellular functions such as cell growth, DNA replication, transcription, and translation because of their highly regulated nature.[14]
The TATA box is usually located 25-35 base pairs upstream of the transcription start site. Genes containing the TATA box usually require additional promoter elements, including an initiator site located just upstream of the transcription start site and a downstream core element (DCE).[3] These additional promoter regions work in conjunction with the TATA box to regulate initiation of transcription in eukaryotes.
Function[edit]
Role in Transcription Initiation[edit]
The TATA-box is the site of preinitiation complex formation, which is the first step in transcription initiation in eukaryotes. Formation of the preinitiation complex begins when the multi-subunit transcription factor II D (TFIID) binds to the TATA box at its TATA-binding protein (TBP) subunit.[3] TBP binds to the minor groove[15] of the TATA box via a region of antiparallel β sheets in the protein.[16] Three types of molecular interactions contribute to TBP binding to the TATA box:
Two phenylalanine residues on insert at the ends of the TATA box and form kinks in the DNA.[16][17][18]
Four hydrogen bonds form between polar side chains on TBP amino acid residues and bases in the minor groove.[16]
Numerous hydrophobic interactions form between TBP residues and DNA bases, including van der Waals forces.[16][17][18]
Additionally, binding of TBP is facilitated by stabilizing interactions with DNA flanking the TATA box, which consists of G-C rich sequences.[19] These secondary interactions induce bending of the DNA and helical unwinding.[20] The degree of DNA bending is species and sequence dependent. For example, one study used the adenovirus TATA promoter sequence (5'-CGCTATAAAAGGGC-3') as a model binding sequence and found that human TBP binding to the TATA box induced a 97° bend toward the major groove while the yeast TBP protein only induced an 82° bend.[21] X-ray crystallography studies of TBP/TATA-box complexes generally agree that the DNA goes through an ~80° bend during the process of TBP-binding.[16][17][18]
The conformational changes induced by TBP binding to the TATA box allows for additional transcription factors and RNA polymerase II to bind to the promoter region. TFIID first binds to the TATA box, facilitated by TFIIA binding to the upstream part of the TFIID complex.[22][23] TFIIB then binds to the TFIID-TFIIA-DNA complex through interactions both upstream and downstream of the TATA box.[24] RNA polymerase II is then recruited to this multi-protein complex with the help of TFIIF.[24] Additional transcription factors then bind, first TFIIE and then TFIIH.[24] This completes the assembly of the preinitiation complex for eukaryotic transcription.[3] Generally, the TATA box is found at RNA polymerase II promoter regions, although some in vitro studies have demonstrated that RNA polymerase III can recognize TATA sequences.[25]
This cluster of RNA polymerase II and various transcription factors is known as the basal transcriptional complex (BTC). In this state, it only gives a low level of transcription. Other factors must stimulate the BTC to increase transcription levels.[2] One such example of a BTC stimulating region of DNA is the CAAT box. Additional factors, including the Mediator complex, transcriptional regulatory proteins, and nucleosome-modifying enzymes also enhance transcription in vivo.[3]
Interactions[edit]
Interaction of TATA boxes with a variety of activators or repressors can influence the transcription of genes in many ways. Enhancers are long-range regulatory elements that increase promotor activity while silencers repress promoter activity.
Mutations[edit]
Figure 3. Effects on TBP binding to the TATA box from mutations. Wildtype shows transcription done normally. An insertion or deletion shifts the TATA box recognition site which results in a shifted transcription site.[26] Point mutations risk the TBP being unable to bind for initiation.[27]
Mutations to the TATA box can range from a deletion or insertion to a point mutation with varying effects based on the gene that has been mutated. The mutations change the binding of the TATA-binding protein (TBP) for transcription initiation. Thus, there is a resulting change in phenotype based on the gene that is not being expressed (Figure 3).
Insertions or Deletions[edit]
One of the first studies of TATA box mutations looked at a sequence of DNA from Agrobacterium tumefaciens for the octopine type cytokinin gene.[26] This specific gene has three TATA boxes. A phenotype change was only observed when all three TATA boxes were deleted. An insertion of extra base pairs between the last TATA box and the transcription start site resulted in a shift in the start site; thus, resulting in a phenotypic change. From this original mutation study, a change in transcription can be seen when there is no TATA box to promote transcription, but transcription of a gene will occur when there is an insertion to the sequence. The nature of the resulting phenotype may be affected due to the insertion.
Mutations in maize promoters affect the expression of the promoter genes in a plant-organ-specific manner.[28] A duplication of the TATA box leads to a significant decrease in enzymatic activity in the scutellum and roots, leaving pollen enzymatic levels unaffected. A deletion of the TATA box leads to a small decrease in enzymatic activity in the scutellum and roots, but a large decrease in enzymatic levels in pollen.[28]
Point Mutations[edit]
Point mutations to the TATA box have similar varying phenotypic changes depending on the gene that is being affected. Studies also show that the placement of the mutation in the TATA box sequence hinders the binding of TBP.[27] For example, a mutation from TATAAAA to CATAAAA does completely hinder the binding sufficiently to change transcription, the neighboring sequences can affect if there is a change or not.[29] However, a change can be seen in HeLa cells with a TATAAAA to TATACAA which leads to a 20 fold decrease in transcription.[30] Some diseases that can be caused due to this insufficiency by specific gene transcription are: Thalassemia,[31] lung cancer,[32] chronic hemolytic anemia,[33] immunosuppression,[34] hemophilia B Leyden,[35] and thrombophlebitis and myocardial infarction.[36]
Savinkova et al. has written a simulation to predict the KD value for a selected TATA box sequence and TBP.[37] This can be used to directly predict the phenotypic traits resulting from a selected mutation based on how tightly TBP is binding to the TATA box.
Diseases[edit]
Mutations in the TATA box region affects the binding of the TATA-binding protein (TBP) for transcription initiation, which may cause carriers to have a disease phenotype.
Gastric cancer is correlated with TATA box polymorphism.[38] The TATA box has a binding site for the transcription factor of the PG2 gene. This gene produces PG2 serum, which is used as a biomarker for tumours in gastric cancer. Longer TATA box sequences correlates with higher levels of PG2 serum indicating gastric cancer conditions. Carriers with shorter TATA box sequences may produce lower levels of PG2 serum.
Several neurodegenerative disorders are associated TATA box mutations.[39] Two disorders have been highlighted, spinocerebellar ataxia and Huntington's disease. In spinocerebellar ataxia, the disease phenotype is caused by expansion of the polyglutamine repeat in the TATA-binding protein (TBP). An accumulation of these polyglutamine-TBP cells will occur, as shown by protein aggregates in brain sections of patients, resulting in a loss of neuronal cells.
Blindness can be caused by excessive cataract formation when the TATA box is targeted by microRNAs to increase the level of oxidative stress genes.[40] MicroRNAs can target the 3'-untranslated region and bind to the TATA box to activate the transcription of oxidative stress related genes.
SNPs in TATA boxes are associated with B-thalassemia, immunosuppression, and other neurological disorders.[41] SNPs destabilize the TBP/TATA complex which significantly decreases the rate at which TATA-binding proteins (TBP) will bind to the TATA box. This leads to lower levels of transcription affecting the severity of the disease. Results from studies have shown the interaction in vitro so far, but results may be comparable to that in vivo.
Gilbert's syndrome is correlated with UTG1A1 TATA box polymorphism.[42] This poses a risk for developing jaundice in newborns.
MicroRNAs also play a role in replicating viruses such as HIV-1.[43] Novel HIV-1-encoded microRNA have been found to enhance the production of the virus as well as activating HIV-1 latency by targeting the TATA box region.
Clinical Significance[edit]
Technology[edit]
Many of the studies so far have been performed in vitro, providing only a prediction of what may happen not a real-time representation of what is happening in the cells. Recent studies in 2016 have been done to demonstrate TATA-binding activity in vivo. Core promoter-specific mechanisms for transcription initiation by the canonical TBP/TFIID-dependent basal transcription machinery has recently been documented in vivo showing the activation by SRF-dependent upstream activating sequence (UAS) of the human ACTB gene involved in TATA-binding.[5]
Cancer Therapy[edit]
Pharmaceutical companies have been designing cancer therapy drugs to target DNA in traditional methods over the years, and have proven to be successful.[44] However, the toxicity of these drugs have pushed scientists to explore other processes related to DNA that could be targeted instead. In recent years, a collective effort has been made to find cancer-specific molecular targets, such as protein-DNA complexes, which include the TATA binding motif. Compounds that trap the protein-DNA intermediate could result in it being toxic to the cell once they encounter a DNA processing event. Example of drugs that contain such compounds include topotecan, SN-38 (topoisomerase I), doxorubicin, and mitoxantrone (topoisomerase II).[44] Cisplatin is a compound that binds covalently to adjacent guanines in the major groove of DNA, which distorts DNA to allow access of DNA-binding proteins in the minor groove.[44] This will destabilize the interaction between the TATA-binding protein (TBP) to the TATA box. The end result is to immobilize the TATA-binding protein (TBP) on DNA in order to down-regulate transcription initiation.
Genetic engineering[edit]
TATA box modification[edit]
Evolutionary changes have pushed plants to adapt to the changing environmental conditions. In the history of Earth, the development of Earth's aerobic atmosphere resulted in an iron deficiency in plants.[45] Modifications to the iron-regulated transporter 1 (IRT1) promoter region of apple trees, genus Malus, have been done recently to increase iron uptake. By inserting a TATA box in the promoter upstream of the iron-regulated transporter 1 (IRT1) promoter, the promoter activity levels are enhanced, which causes TFIID activity to increase, which helps to up-regulate transcription initiation.[45]
See also[edit]
Pribnow box
Initiator element
References[edit]
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[hide]
v
t
e
Transcription (Bacterial, Eukaryotic)
Transcriptional regulation
prokaryotic
Operon
lac operon
trp operon
gab operon
ara operon
gal operon
Repressor
lac repressor
trp repressor
eukaryotic
Histone-modifying enzymes
(histone/nucleosome):
Histone methylation/Histone methyltransferase
EZH2
Histone demethylase
Histone acetylation and deacetylation
Histone deacetylase HDAC1
Histone acetyltransferase
DNA methylation:
DNA methyltransferase
Chromatin remodeling:
CHD7
both
Transcription coregulator
Coactivator
Corepressor
Inducer
Promotion
Promoter
Pribnow box
TATA box
BRE
CAAT box
Response element
Enhancer
E-box
Response element
Insulator
Silencer
Internal control region
Initiation (bacterial,
eukaryotic
Transcription start site
Elongation
bacterial RNA polymerase: rpoB
eukaryotic RNA polymerase: RNA polymerase II
Termination
(bacterial,
eukaryotic)
Terminator
Intrinsic termination
Rho factor
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catlinks mw-normal-catlinks
catlinks mw-normal-catlinks
interface

value: Discussion about edits from this IP address [ctrl-alt-n]

value: A list of edits made from this IP address [ctrl-alt-y]

value: You are encouraged to create an account and log in; however, it is not mandatory

value: You're encouraged to log in; however, it's not mandatory. [ctrl-alt-o]

value:
/wiki/TATA_box
View the content page [ctrl-alt-c]
c

value: Discussion about the content page [ctrl-alt-t]

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value:
p-variants
navigation
vectorMenu emptyPortlet
p-variants-label

value:
/w/index.php?title=TATA_box&action=edit
Edit this page [ctrl-alt-e]
e

value: Past revisions of this page [ctrl-alt-h]

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value:
p-cactions
navigation
vectorMenu emptyPortlet
p-cactions-label

value:
mw-wiki-logo
/wiki/Main_Page
Visit the main page

value:
/wiki/Main_Page
Visit the main page [ctrl-alt-z]
z

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value:
https://cs.wikipedia.org/wiki/TATA_box
TATA box – Czech
cs
cs
interlanguage-link-target

value:
interlanguage-link interwiki-cs
display: none;

value:
https://de.wikipedia.org/wiki/TATA-Box
TATA-Box – German
de
de
interlanguage-link-target

value:
https://es.wikipedia.org/wiki/Caja_TATA
Caja TATA – Spanish
es
es
interlanguage-link-target

value:
interlanguage-link interwiki-gl
display: none;

value:
https://it.wikipedia.org/wiki/TATA_box
TATA box – Italian
it
it
interlanguage-link-target

value:
interlanguage-link interwiki-he
display: none;

value:
https://nl.wikipedia.org/wiki/TATA-box
TATA-box – Dutch
nl
nl
interlanguage-link-target

value:
interlanguage-link interwiki-nl
display: none;

value:
interlanguage-link interwiki-pl
display: none;

value: ТАТА-бокс – Russian

value: interlanguage-link interwiki-ru badge-Q17559452 badge-recommendedarticle

value: ТАТА Кутија – Serbian

value:
interlanguage-link interwiki-sr
display: none;

value:
interlanguage-link interwiki-sh
display: none;

value:
https://sv.wikipedia.org/wiki/TATA-box
TATA-box – Swedish
sv
sv
interlanguage-link-target

value:
interlanguage-link interwiki-sv
display: none;

value:
https://th.wikipedia.org/wiki/TATA_box
TATA box – Thai
th
th
interlanguage-link-target

value:
interlanguage-link interwiki-th
display: none;

value:
interlanguage-link interwiki-uk
display: none;

value: All languages (initial selection from common choices by you and others)

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value:
suggestions
display: none; font-size: 13px;

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