text:
Thursday
From Wikipedia, the free encyclopedia
Jump to: navigation, search
For other uses, see Thursday (disambiguation).
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. (November 2014) (Learn how and when to remove this template message)
Thursday is the day of the week between Wednesday and Friday. According to the ISO 8601 international standard, it is the fourth day of the week.
Contents
[hide]
1 Name
1.1 Thor's (Jupiter's) day
1.2 Fourth day
1.3 Fifth day
2 Cultural and religious practices
2.1 Christian holidays
2.2 Judaism
2.3 Thailand
2.4 Conventional weekly events
2.5 Elections in the United Kingdom
3 Astrology
4 Popular culture
4.1 Literature
4.2 Cinema
4.3 Music
5 References
Name[edit]
See Names of the days of the week for more on naming conventions.
Thor's (Jupiter's) day[edit]
Painting depicting the Norse god Thunor (the Norse Thor), after whom Thursday is named, by Mårten Eskil Winge, 1872
The name is derived from Old English Þūnresdæg and Middle English Thuresday (with loss of -n-, first in northern dialects, from influence of Old Norse Þorsdagr) meaning "Thor's Day". It was named after the Norse god of Thunder, Thor. [1][2][3] Thunor, Donar (German, Donnerstag) and Thor are derived from the name of the Germanic god of thunder, Thunraz, equivalent to Jupiter in the interpretatio romana.
Estonians did not work on Thursdays (days of Thor) and Thursday nights were called "evenings of Tooru".[citation needed] Some sources say Estonians used to gather in holy woods (Hiis) on Thursday evenings, where a bagpipe player sat on a stone and played while people danced and sang until the dawn.
In most Romance languages, the day is named after the Roman god Jupiter, who was the god of sky and thunder. In Latin, the day was known as Iovis Dies, "Jupiter's Day". In Latin, the genitive or possessive case of Jupiter was Iovis/Jovis and thus in most Romance languages it became the word for Thursday: Italian giovedì, Spanish jueves, French jeudi, Sardinian jòvia, Catalan dijous, Galician xoves and Romanian joi. This is also reflected in the p-Celtic Welsh dydd Iau.
The astrological and astronomical sign of the planet Jupiter (♃ ) is sometimes used to represent Thursday.
Since the Roman god Jupiter was identified with Thunor (Norse Thor in northern Europe), most Germanic languages name the day after this god: Torsdag in Danish, Norwegian, and Swedish, Hósdagur/Tórsdagur in Faroese, Donnerstag in German or Donderdag in Dutch. Finnish and Northern Sami, both non-Germanic (Uralic) languages, uses the borrowing "Torstai" and "Duorastat". In the extinct Polabian Slavic language, it was perundan, Perun being the Slavic equivalent of Thor.[4]
There are a number of modern names imitating the naming of Thursday after an equivalent of "Jupiter" in local tradition. In most of the languages of India, the word for Thursday is Guruvāra – vāra meaning day and Guru being the style for Bṛhaspati, guru to the gods and regent of the planet Jupiter. In Thai, the word is Wan Pharuehatsabodi – referring to the Hindu deity Bṛhaspati, also associated with Jupiter. En was an old Illyrian deity and in his honor in the Albanian language Thursday is called "Enjte".[5] In the Nahuatl language, Thursday is Tezcatlipotōnal (Nahuatl pronunciation: [teskat͡ɬipoˈtoːnaɬ]) meaning "day of Tezcatlipoca".
Fourth day[edit]
This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. (January 2008) (Learn how and when to remove this template message)
In Slavic languages and in Chinese, this day's name is "fourth" (Slovak štvrtok, Czech čtvrtek, Slovene četrtek, Croatian and Bosnian četvrtak, Polish czwartek, Russian "четверг" četverg, Bulgarian "четвъртък", Serbian "четвртак", Macedonian "четврток", Ukrainian "четвер" chetver.). Hungarian uses a Slavic loanword "csütörtök". In Chinese, it's 星期四 xīngqīsì ("fourth solar day"). In ancient Chinese, it is 木曜日. In Estonian it's "neljapäev", meaning fourth day or fourth day in a week. The Baltic languages also use the term "fourth day" (Latvian ceturtdiena, Lithuanian ketvirtadienis).
Fifth day[edit]
This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. (January 2008) (Learn how and when to remove this template message)
Greek uses a number for this day: Πέμπτη Pémpti "fifth," as does Portuguese: quinta-feira "fifth day," Hebrew: "יום חמישי" ("Yom Hamishi" – day fifth) often written 'יום ה ("Yom Hey" – 5th letter Hey day), and Arabic: "يوم الخميس" ("Yaum al-Khamīs" – fifth day). In Catholic liturgy, Thursday is referred to in Latin as feria quinta. Portuguese, unlike other Romance languages, uses the word quinta-feira, meaning "fifth day of liturgical celebration", that comes from the Latin "feria quinta" used in religious texts where it was not allowed to consecrate days to pagan gods.
Icelandic also uses the term fifth day (Fimmtudagur).
Quakers traditionally referred to Thursday as "Fifth Day" eschewing the pagan origin of the English name "Thursday".[6]
Cultural and religious practices[edit]
Christian holidays[edit]
In the Christian tradition, Maundy Thursday or Holy Thursday is the Thursday before Easter — the day on which the Last Supper occurred. Also known as Sheer Thursday in the United Kingdom, it is traditionally a day of cleaning and giving out Maundy money there. Holy Thursday is part of Holy Week.
Named after Thursday.
In the Eastern Orthodox Church. Thursdays are dedicated to the Apostles and Saint Nicholas. The Octoechos contains hymns on these themes, arranged in an eight-week cycle, that are chanted on Thursdays throughout the year. At the end of Divine Services on Thursday, the dismissal begins with the words: "May Christ our True God, through the intercessions of his most-pure Mother, of the holy, glorious and all-laudable Apostles, of our Father among the saints Nicholas, Archbishop of Myra in Lycia, the Wonder-worker…"
Ascension Thursday is 40 days after Easter, when Christ ascended into Heaven.
In the United States, Thanksgiving Day is an annual festival celebrated on the fourth Thursday in November.
Judaism[edit]
In Judaism, Thursdays are considered auspicious days for fasting. The Didache warned early Christians not to fast on Thursdays to avoid Judaizing, and suggested Fridays instead.
In Judaism the Torah is read in public on Thursday mornings, and special penitential prayers are said on Thursday, unless there is a special occasion for happiness which cancels them.
Thailand[edit]
This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. (January 2008) (Learn how and when to remove this template message)
In Buddhist Thailand Thursday is considered the "Teacher's Day", and it is believed that one should begin one's education on this auspicious day. Thai students still pay homages to their teachers in specific ceremony always held on a selected Thursday. And graduation day in Thai universities, which can vary depending on each university, almost always will be held on a Thursday.
In the Thai solar calendar, the colour associated with Thursday is orange.
Conventional weekly events[edit]
In Australia, most cinema movies premieres are held on Thursdays. Also, most Australians are paid on a Thursday, either weekly or fortnightly. Shopping malls see this as an opportunity to open longer than usual, generally until 9 pm, as most pay cheques are cleared by Thursday morning.
In Norway, Thursday has also traditionally been the day when most shops and malls are open later than on the other weekdays, although the majority of shopping malls now are open until 8 pm or 9 pm every weekday.
In the USSR of the 1970s and 1980s Thursday was the "Fish Day" (Russian: Рыбный день, Rybny den), when the nation's foodservice establishments were supposed to serve fish (rather than meat) dishes.[7]
For college and university students, Thursday is sometimes referred to as the new Friday. There are often fewer or sometimes no classes on Fridays and more opportunities to hold parties on Thursday night and sleep in on Friday. As a consequence, some call Thursday "thirstday" or "thirsty Thursday".[8]
Elections in the United Kingdom[edit]
In the United Kingdom, all general elections since 1935 have been held on a Thursday, and this has become a tradition, although not a requirement of the law — which merely states that an election may be held on any day "except Saturdays, Sundays, Christmas Eve, Christmas Day, Good Friday, bank holidays in any part of the United Kingdom and any day appointed for public thanksgiving and mourning."[9]
Additionally, local elections are usually held on the first Thursday in May.[citation needed]
The Electoral Administration Act 2006 removed Maundy Thursday as an excluded day on the electoral timetable, therefore an election can now be held on Maundy Thursday; prior to this elections were sometimes scheduled on the Tuesday before as an alternative.
Astrology[edit]
Thursday is aligned by the planet Jupiter and the astrological signs of Pisces and Sagittarius.
Popular culture[edit]
In the nursery rhyme, "Monday's Child", "Thursday's Child has far to go".
In some high schools in the United States during the 1950s and the 1960s, rumours said that if someone wore green on Thursdays, it meant that he or she was gay.[10]
Thursday is the day of the Second Round draw in the English League Cup.
Super Thursday is an annual promotional event in the publishing industry as well as an important day in UK elections (see above).
Literature[edit]
Gabriel Syme, the main character, was given the title of Thursday in G. K. Chesterton's novel The Man Who Was Thursday (1908).
The titular day in Sweet Thursday (1954) (the sequel to John Steinbeck's novel Cannery Row (1945)), the author explains, is the day after Lousy Wednesday and the day before Waiting Friday.
In The Hitchhiker's Guide to the Galaxy by Douglas Adams, the character Arthur Dent says: "This must be Thursday. I never could get the hang of Thursdays". A few minutes later the planet Earth is destroyed. In another Douglas Adams book, The Long Dark Tea-time of the Soul (1988), one of the characters says to the character Thor, after whom the day was named: "I'm not used to spending the evening with someone who's got a whole day named after them".
In the cross media work Thursday's Fictions by Richard James Allen and Karen Pearlman, Thursday is the title character, a woman who tries to cheat the cycle of reincarnation to get a form of eternal life. Thursday's Fictions has been a stage production, a book, a film and an 3D online immersive world in Second Life.[11]
Thursday Next is the central character in a series of novels by Jasper Fforde.
In Garth Nix's popular The Keys to the Kingdom series, Thursday is an antagonist, a violent general who is a personification of the actual day and the Sin of Wrath.
According to Nostradamus' prediction (Century 1, Quatrain 50), a powerful (but otherwise unidentified) leader who will threaten "the East" will be born of three water signs and takes Thursday as his feast day.[12]
Cinema[edit]
Thursday (1998 film) is a movie starring Thomas Jane, about the day of a drug dealer gone straight, who gets pulled back into his old lifestyle.
The Thursday (1963), is an Italian film.
Music[edit]
Thursday Afternoon is a 1985 album by the British ambient musician Brian Eno consisting of one 60-minute-long composition. It is the rearranged soundtrack to a video production of the same title made in 1984.
Donnerstag aus Licht (Thursday from Light) is an opera by Karlheinz Stockhausen.
Thursday is a post-hardcore band from New Brunswick, New Jersey, formed in 1997.
"Thursday's Child" is a David Bowie song from the album hours...(1999).
"Thursday's Child" is a song by The Chameleons on Script of the Bridge (1983).
Outlook for Thursday was a hit in New Zealand for Dave Dobbyn.
In the Placebo song "Evil Dildo", the obscene telephone message is left on Thursday the 23rd of an unknown month and year. The day Thursday 23rd is often celebrated as Evil Dildo day by Placebo fans.[citation needed]
Thursday (mixtape)" is the name of a mixtape by R&B artist The Weeknd released in 2011.
References[edit]
Wikimedia Commons has media related to Thursday.
Wikiquote has quotations related to: Thursday
Look up Thursday in Wiktionary, the free dictionary.
Jump up ^ "Anglo-Saxon Week". English Heathenism. Archived from the original on September 24, 2009. Retrieved July 30, 2015.
Jump up ^ Stone, John Robert (1997). "Observing Bede's Anglo-Saxon Calendar". The English Companions. Archived from the original on May 4, 2010. Retrieved July 30, 2015.
Jump up ^ "Online Etymology Dictionary". Etymonline.com. Retrieved 2012-08-06.
Jump up ^ Selected writings: Comparative Slavic studies – Roman Jakobson – Google Books. Books.google.com. Retrieved 2012-08-06.
Jump up ^ Lurker, Manfred. The Routledge Dictionary of Gods and Goddesses, Devils and Demons, Taylor & Francis e-Library, 2005. p.57
Jump up ^ "Guide to Quaker Calendar Names". Iowa Yearly Meeting (Conservative) Religious Society of Friends (Quakers). Retrieved 30 March 2017. In the 20th Century, many Friends began accepting use of the common date names, feeling that any pagan meaning has been forgotten. The numerical names continue to be used, however, in many documents and more formal situations."
Jump up ^ Petrosian, Irina; Underwood, David (2006), Armenian Food: Fact, Fiction & Folklore, Armenian Research Center collection (2 ed.), p. 115, ISBN 1411698657
Jump up ^ Hafner, Katie (November 6, 2005). "How Thursday Became the New Friday". The New York Times. Retrieved March 28, 2010.
Jump up ^ Schedule 1, Representation of the People Act 1983, 1996-11-01. Retrieved on 2016-11-03.
Jump up ^ Grahn, Judy (1990). Another Mother Tongue: Gay Words, Gay Worlds (updated and expanded ed.). Boston: Beacon Press. pp. 76–81. ISBN 0-8070-7911-1.
Jump up ^ "Magazine – issue 80 – dance film: spiritual odyssey". RealTime Arts. Retrieved 2012-08-06.
Jump up ^ Nostradamus. "Century 1 – Quatrain 50". Nostradamus Quatrains. Retrieved 2012-08-06.
[hide]
v
t
e
Days of the week
Monday
Tuesday
Wednesday
Thursday
Friday
Saturday
Sunday
Authority control
GND: 4275828-2
Retrieved from "https://en.wikipedia.org/w/index.php?title=Thursday&oldid=837030100"
Categories:
Days of the week
Thursday
Hidden categories:
Articles needing additional references from November 2014
All articles needing additional references
All articles with unsourced statements
Articles with unsourced statements from November 2017
Articles containing Classical Nahuatl-language text
Articles needing additional references from January 2008
Articles containing Portuguese-language text
Articles containing Russian-language text
Articles with unsourced statements from November 2016
Articles with unsourced statements from January 2012
Wikipedia articles with GND identifiers
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
In other projects
Wikimedia Commons
Wikiquote
Languages
Afrikaans
Alemannisch
አማርኛ
Ænglisc
Аҧсшәа
العربية
Aragonés
ܐܪܡܝܐ
Armãneashti
Arpetan
Asturianu
Avañe'ẽ
Авар
Azərbaycanca
تۆرکجه
বাংলা
Bahasa Banjar
Bân-lâm-gú
Башҡортса
Беларуская
Беларуская (тарашкевіца)
भोजपुरी
Bikol Central
Български
Boarisch
བོད་ཡིག
Bosanski
Brezhoneg
Буряад
Català
Чӑвашла
Cebuano
Čeština
Chavacano de Zamboanga
ChiShona
Corsu
Cymraeg
Dansk
Davvisámegiella
Deitsch
Deutsch
ދިވެހިބަސް
Dolnoserbski
Eesti
Ελληνικά
Emiliàn e rumagnòl
Эрзянь
Español
Esperanto
Estremeñu
Euskara
Eʋegbe
فارسی
Føroyskt
Français
Frysk
Furlan
Gaeilge
Gaelg
Gagauz
Gàidhlig
Galego
贛語
گیلکی
ગુજરાતી
𐌲𐌿𐍄𐌹𐍃𐌺
客家語/Hak-kâ-ngî
한국어
Hausa
Hawaiʻi
Հայերեն
हिन्दी
Hornjoserbsce
Hrvatski
Ido
Ilokano
Bahasa Indonesia
Interlingua
Ирон
Íslenska
Italiano
עברית
Basa Jawa
Kalaallisut
ಕನ್ನಡ
Къарачай-малкъар
ქართული
Kaszëbsczi
Қазақша
Kernowek
Kinyarwanda
Kiswahili
Коми
Kreyòl ayisyen
Kurdî
Кыргызча
Ladino
Лакку
Лезги
ລາວ
Latina
Latviešu
Lëtzebuergesch
Lietuvių
Lingála
Livvinkarjala
Lumbaart
Magyar
Македонски
മലയാളം
मराठी
მარგალური
مصرى
مازِرونی
Bahasa Melayu
Baso Minangkabau
Mìng-dĕ̤ng-ngṳ̄
Мокшень
Монгол
မြန်မာဘာသာ
Nāhuatl
Nederlands
Nedersaksies
नेपाली
日本語
Napulitano
Нохчийн
Norsk
Norsk nynorsk
Nouormand
Occitan
Олык марий
ଓଡ଼ିଆ
Oʻzbekcha/ўзбекча
ਪੰਜਾਬੀ
Pangasinan
پنجابی
پښتو
Перем Коми
ភាសាខ្មែរ
Tok Pisin
Plattdüütsch
Polski
Ποντιακά
Português
Ripoarisch
Română
Runa Simi
Русиньскый
Русский
Gagana Samoa
संस्कृतम्
Sardu
Scots
Sesotho sa Leboa
Shqip
සිංහල
Simple English
سنڌي
Slovenčina
Slovenščina
Словѣньскъ / ⰔⰎⰑⰂⰡⰐⰠⰔⰍⰟ
Ślůnski
Soomaaliga
کوردی
Српски / srpski
Srpskohrvatski / српскохрватски
Basa Sunda
Suomi
Svenska
Tagalog
தமிழ்
Taqbaylit
Tarandíne
Татарча/tatarça
తెలుగు
ไทย
ትግርኛ
Тоҷикӣ
ᏣᎳᎩ
Türkçe
Türkmençe
Twi
Тыва дыл
ᨅᨔ ᨕᨘᨁᨗ
Українська
اردو
ئۇيغۇرچە / Uyghurche
Vahcuengh
Vèneto
Vepsän kel’
Tiếng Việt
Volapük
Võro
Walon
West-Vlams
Winaray
Wolof
吴语
ייִדיש
Yorùbá
粵語
Zazaki
Žemaitėška
中文
डोटेली
Kabɩyɛ
198 more
Edit links
This page was last edited on 18 April 2018, at 08:34.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

key: html

type: local

value: div

attrs: id class

values: mw-page-base noprint

values: mw-head-base noprint

value: a

attrs: id class lang

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

attrs: href class title

values:
/wiki/Thursday_(disambiguation)
mw-disambig
Thursday (disambiguation)

attrs: role class

values:
note
hatnote navigation-not-searchable

value: img

attrs: alt src width height srcset data-file-width data-file-height

attrs: href class

value: td

attrs: href title

value: b

attrs: class href

values: Help:Introduction to referencing with Wiki Markup/1

value: span

value: i

value: small

values: plainlinks metadata ambox ambox-content ambox-Refimprove

values:
/wiki/Week-day_names
mw-redirect
Week-day names

attrs: alt src width height class srcset data-file-width data-file-height

values:
/wiki/Interpretatio_romana
mw-redirect
Interpretatio romana

values: This claim needs references to reliable sources. (November 2017)

values:
noprint Inline-Template Template-Fact
white-space:nowrap;

values:
/wiki/Jupiter_(astrology)
mw-redirect
Jupiter (astrology)

values:
/wiki/En_(Illyrian_god)
mw-redirect
En (Illyrian god)

values:
/wiki/Nahuatl_language
mw-redirect
Nahuatl language

values: Representation in the International Phonetic Alphabet (IPA)

values: plainlinks metadata ambox ambox-content ambox-Unreferenced

values:
/wiki/Hebrew_Language
mw-redirect
Hebrew Language

values:
/wiki/Arabic_Language
mw-redirect
Arabic Language

values:
/wiki/Religious_Society_of_Friends
mw-redirect
Religious Society of Friends

values: Edit section: Cultural and religious practices

values:
/wiki/Divine_Services
mw-redirect
Divine Services

values:
/wiki/Holy_Fathers
mw-redirect
Holy Fathers

values:
/wiki/Ascension_of_Jesus_Christ
mw-redirect
Ascension of Jesus Christ

values: Edit section: Conventional weekly events

values: ru:Рыбный день

values: Edit section: Elections in the United Kingdom

values: This claim needs references to reliable sources. (November 2016)

values:
/wiki/Cross_media
mw-redirect
Cross media

values: Thursday's Fictions (page does not exist)

values:
/wiki/Seven_Deadly_Sins
mw-redirect
Seven Deadly Sins

values:
/wiki/Water_signs
mw-redirect
Water signs

values:
/wiki/Thursday_(1998_film)
mw-redirect
Thursday (1998 film)

values:
/wiki/Placebo_band
mw-redirect
Placebo band

values: This claim needs references to reliable sources. (January 2012)

values:
/wiki/Thursday_(mixtape)
mw-redirect
Thursday (mixtape)

values:
nofollow
external text
http://iymc.org/calendarnames.html

values: nowraplinks collapsible autocollapse navbox-inner

values:
nofollow
external text
https://d-nb.info/gnd/4275828-2

values: Category:Articles needing additional references from November 2014

values: Category:All articles needing additional references

values: Category:All articles with unsourced statements

values: Category:Articles with unsourced statements from November 2017

values: Category:Articles containing Classical Nahuatl-language text

values: Category:Articles needing additional references from January 2008

values: Category:Articles containing Portuguese-language text

values: Category:Articles containing Russian-language text

values: Category:Articles with unsourced statements from November 2016

values: Category:Articles with unsourced statements from January 2012

values: Category:Wikipedia articles with GND identifiers

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

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

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

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

values:
/wiki/Thursday
View the content page [ctrl-alt-c]
c

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

values: p-namespaces navigation vectorTabs p-namespaces-label

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

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

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

values: p-views navigation vectorTabs p-views-label

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

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

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

values: Featured content – the best of Wikipedia

values: Find background information on current events

values:
/wiki/Special:Random
Load a random article [ctrl-alt-x]
x

values: portal navigation p-navigation p-navigation-label

values: Guidance on how to use and edit Wikipedia

values: About the project, what you can do, where to find things

values: A list of recent changes in the wiki [ctrl-alt-r]

values: portal navigation p-interaction p-interaction-label

values: List of all English Wikipedia pages containing links to this page [ctrl-alt-j]

values: Recent changes in pages linked from this page [ctrl-alt-k]

values:
/wiki/Wikipedia:File_Upload_Wizard
Upload files [ctrl-alt-u]
u

values: A list of all special pages [ctrl-alt-q]

values: Permanent link to this revision of the page

values:
https://af.wikipedia.org/wiki/Donderdag
Donderdag – Afrikaans
af
af
interlanguage-link-target

values:
interlanguage-link interwiki-af
display: none;

values:
interlanguage-link interwiki-als
display: none;

values:
interlanguage-link interwiki-am
display: none;

values:
interlanguage-link interwiki-ang
display: none;

values:
interlanguage-link interwiki-ab
display: none;

values: الخميس – Arabic

values:
https://an.wikipedia.org/wiki/Chueves
Chueves – Aragonese
an
an
interlanguage-link-target

values:
interlanguage-link interwiki-an
display: none;

values: ܚܡܫܒܫܒܐ – Aramaic

values:
interlanguage-link interwiki-arc
display: none;

values:
https://roa-rup.wikipedia.org/wiki/Gioi
Gioi – Aromanian
roa-rup
roa-rup
interlanguage-link-target

values:
interlanguage-link interwiki-roa-rup
display: none;

values:
interlanguage-link interwiki-frp
display: none;

values:
https://ast.wikipedia.org/wiki/Xueves
Xueves – Asturian
ast
ast
interlanguage-link-target

values:
interlanguage-link interwiki-ast
display: none;

values:
https://gn.wikipedia.org/wiki/Arapo
Arapo – Guarani
gn
gn
interlanguage-link-target

values:
interlanguage-link interwiki-gn
display: none;

values: Хамиз – Avaric

values:
interlanguage-link interwiki-av
display: none;

values:
interlanguage-link interwiki-az
display: none;

values:
interlanguage-link interwiki-azb
display: none;

values: বৃহস্পতিবার – Bangla

values:
interlanguage-link interwiki-bn
display: none;

values:
interlanguage-link interwiki-bjn
display: none;

values:
interlanguage-link interwiki-zh-min-nan
display: none;

values: Кесаҙна – Bashkir

values:
interlanguage-link interwiki-ba
display: none;

values:
interlanguage-link interwiki-be
display: none;

values: Чацьвер – Belarusian (Taraškievica orthography)

values:
interlanguage-link interwiki-be-x-old
display: none;

values:
interlanguage-link interwiki-bh
display: none;

values:
https://bcl.wikipedia.org/wiki/Huwebes
Huwebes – Central Bikol
bcl
bcl
interlanguage-link-target

values:
interlanguage-link interwiki-bcl
display: none;

values: Четвъртък – Bulgarian

values:
interlanguage-link interwiki-bg
display: none;

values:
https://bar.wikipedia.org/wiki/Pfinzda
Pfinzda – Bavarian
bar
bar
interlanguage-link-target

values:
interlanguage-link interwiki-bar
display: none;

values: གཟའ་ཕུར་བུ། – Tibetan

values:
interlanguage-link interwiki-bo
display: none;

values:
interlanguage-link interwiki-bs
display: none;

values:
interlanguage-link interwiki-br
display: none;

values: Гарагай табан – Russia Buriat

values:
interlanguage-link interwiki-bxr
display: none;

values:
https://ca.wikipedia.org/wiki/Dijous
Dijous – Catalan
ca
ca
interlanguage-link-target

values:
interlanguage-link interwiki-ca
display: none;

values: Кĕçнерникун – Chuvash

values:
interlanguage-link interwiki-cv
display: none;

values:
https://ceb.wikipedia.org/wiki/Huwebes
Huwebes – Cebuano
ceb
ceb
interlanguage-link-target

values:
interlanguage-link interwiki-ceb
display: none;

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

values:
interlanguage-link interwiki-cbk-zam
display: none;

values:
interlanguage-link interwiki-sn
display: none;

values:
https://co.wikipedia.org/wiki/Ghjovi
Ghjovi – Corsican
co
co
interlanguage-link-target

values:
interlanguage-link interwiki-co
display: none;

values:
https://cy.wikipedia.org/wiki/Dydd_Iau
Dydd Iau – Welsh
cy
cy
interlanguage-link-target

values:
interlanguage-link interwiki-cy
display: none;

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

values:
interlanguage-link interwiki-da
display: none;

values:
https://se.wikipedia.org/wiki/Duorastat
Duorastat – Northern Sami
se
se
interlanguage-link-target

values:
interlanguage-link interwiki-se
display: none;

values:
interlanguage-link interwiki-pdc
display: none;

values:
interlanguage-link interwiki-de
display: none;

values: ބުރާސްފަތި ދުވަސް – Divehi

values:
interlanguage-link interwiki-dv
display: none;

values:
interlanguage-link interwiki-dsb
display: none;

values:
interlanguage-link interwiki-et
display: none;

values: Πέμπτη – Greek

values:
interlanguage-link interwiki-el
display: none;

values:
interlanguage-link interwiki-eml
display: none;

values: Калоньчи – Erzya

values:
interlanguage-link interwiki-myv
display: none;

values:
https://es.wikipedia.org/wiki/Jueves
Jueves – Spanish
es
es
interlanguage-link-target

values:
interlanguage-link interwiki-eo
display: none;

values:
https://ext.wikipedia.org/wiki/Juevis
Juevis – Extremaduran
ext
ext
interlanguage-link-target

values:
interlanguage-link interwiki-ext
display: none;

values:
https://eu.wikipedia.org/wiki/Ostegun
Ostegun – Basque
eu
eu
interlanguage-link-target

values:
interlanguage-link interwiki-eu
display: none;

values:
interlanguage-link interwiki-ee
display: none;

values: پنجشنبه – Persian

values:
interlanguage-link interwiki-fa
display: none;

values:
interlanguage-link interwiki-fo
display: none;

values:
https://fr.wikipedia.org/wiki/Jeudi
Jeudi – French
fr
fr
interlanguage-link-target

values:
interlanguage-link interwiki-fr
display: none;

values:
interlanguage-link interwiki-fy
display: none;

values:
https://fur.wikipedia.org/wiki/Joibe
Joibe – Friulian
fur
fur
interlanguage-link-target

values:
interlanguage-link interwiki-fur
display: none;

values:
interlanguage-link interwiki-ga
display: none;

values:
https://gv.wikipedia.org/wiki/Jerdein
Jerdein – Manx
gv
gv
interlanguage-link-target

values:
interlanguage-link interwiki-gv
display: none;

values:
interlanguage-link interwiki-gag
display: none;

values:
interlanguage-link interwiki-gd
display: none;

values:
https://gl.wikipedia.org/wiki/Xoves
Xoves – Galician
gl
gl
interlanguage-link-target

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

values:
interlanguage-link interwiki-gan
display: none;

values: پئنشمبه – Gilaki

values:
interlanguage-link interwiki-glk
display: none;

values: ગુરુવાર – Gujarati

values:
interlanguage-link interwiki-gu
display: none;

values: 𐍀𐌰𐌹𐌽𐍄𐌴𐌳𐌰𐌲𐍃 – Gothic

values:
interlanguage-link interwiki-got
display: none;

values:
interlanguage-link interwiki-hak
display: none;

values:
interlanguage-link interwiki-ko
display: none;

values:
https://ha.wikipedia.org/wiki/Alhamis
Alhamis – Hausa
ha
ha
interlanguage-link-target

values:
interlanguage-link interwiki-ha
display: none;

values:
interlanguage-link interwiki-haw
display: none;

values: Հինգշաբթի – Armenian

values:
interlanguage-link interwiki-hy
display: none;

values: गुरुवार – Hindi

values:
interlanguage-link interwiki-hsb
display: none;

values:
interlanguage-link interwiki-hr
display: none;

values:
https://io.wikipedia.org/wiki/Jovdio
Jovdio – Ido
io
io
interlanguage-link-target

values:
interlanguage-link interwiki-io
display: none;

values:
https://ilo.wikipedia.org/wiki/Huebes
Huebes – Iloko
ilo
ilo
interlanguage-link-target

values:
interlanguage-link interwiki-ilo
display: none;

values:
https://id.wikipedia.org/wiki/Kamis
Kamis – Indonesian
id
id
interlanguage-link-target

values:
https://ia.wikipedia.org/wiki/Jovedi
Jovedi – Interlingua
ia
ia
interlanguage-link-target

values:
interlanguage-link interwiki-ia
display: none;

values: Цыппæрæм – Ossetic

values:
interlanguage-link interwiki-os
display: none;

values:
interlanguage-link interwiki-is
display: none;

values:
interlanguage-link interwiki-it
display: none;

values: יום חמישי – Hebrew

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

values:
https://jv.wikipedia.org/wiki/Kemis
Kemis – Javanese
jv
jv
interlanguage-link-target

values:
interlanguage-link interwiki-jv
display: none;

values:
interlanguage-link interwiki-kl
display: none;

values: ಗುರುವಾರ – Kannada

values:
interlanguage-link interwiki-kn
display: none;

values:
interlanguage-link interwiki-krc
display: none;

values: ხუთშაბათი – Georgian

values:
interlanguage-link interwiki-ka
display: none;

values:
interlanguage-link interwiki-csb
display: none;

values: Бейсенбі – Kazakh

values:
interlanguage-link interwiki-kk
display: none;

values:
https://kw.wikipedia.org/wiki/Dy%27_Yow
Dy' Yow – Cornish
kw
kw
interlanguage-link-target

values:
interlanguage-link interwiki-kw
display: none;

values:
interlanguage-link interwiki-rw
display: none;

values:
https://sw.wikipedia.org/wiki/Alhamisi
Alhamisi – Swahili
sw
sw
interlanguage-link-target

values:
interlanguage-link interwiki-sw
display: none;

values: Четверг – Komi

values:
interlanguage-link interwiki-kv
display: none;

values:
https://ht.wikipedia.org/wiki/Jedi
Jedi – Haitian Creole
ht
ht
interlanguage-link-target

values:
interlanguage-link interwiki-ht
display: none;

values:
interlanguage-link interwiki-ku
display: none;

values: Бейшемби – Kyrgyz

values:
interlanguage-link interwiki-ky
display: none;

values:
https://lad.wikipedia.org/wiki/Djueves
Djueves – Ladino
lad
lad
interlanguage-link-target

values:
interlanguage-link interwiki-lad
display: none;

values:
interlanguage-link interwiki-lbe
display: none;

values:
interlanguage-link interwiki-lez
display: none;

values: ວັນພະຫັດ – Lao

values:
interlanguage-link interwiki-lo
display: none;

values:
interlanguage-link interwiki-la
display: none;

values:
interlanguage-link interwiki-lv
display: none;

values:
interlanguage-link interwiki-lb
display: none;

values:
interlanguage-link interwiki-lt
display: none;

values:
interlanguage-link interwiki-ln
display: none;

values:
interlanguage-link interwiki-olo
display: none;

values:
interlanguage-link interwiki-lmo
display: none;

values:
interlanguage-link interwiki-hu
display: none;

values:
interlanguage-link interwiki-mk
display: none;

values: വ്യാഴം (ദിവസം) – Malayalam

values:
interlanguage-link interwiki-ml
display: none;

values: गुरुवार – Marathi

values:
interlanguage-link interwiki-mr
display: none;

values:
interlanguage-link interwiki-xmf
display: none;

values:
interlanguage-link interwiki-arz
display: none;

values:
interlanguage-link interwiki-mzn
display: none;

values:
https://ms.wikipedia.org/wiki/Khamis
Khamis – Malay
ms
ms
interlanguage-link-target

values:
https://min.wikipedia.org/wiki/Kamih
Kamih – Minangkabau
min
min
interlanguage-link-target

values:
interlanguage-link interwiki-min
display: none;

values:
interlanguage-link interwiki-cdo
display: none;

values: Шуваланя – Moksha

values:
interlanguage-link interwiki-mdf
display: none;

values: Пүрэв гараг – Mongolian

values:
interlanguage-link interwiki-mn
display: none;

values: ကြာသပတေး – Burmese

values:
interlanguage-link interwiki-my
display: none;

values:
interlanguage-link interwiki-nah
display: none;

values:
https://nl.wikipedia.org/wiki/Donderdag
Donderdag – Dutch
nl
nl
interlanguage-link-target

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

values:
interlanguage-link interwiki-nds-nl
display: none;

values: बिहीवार – Nepali

values:
interlanguage-link interwiki-ne
display: none;

values:
interlanguage-link interwiki-ja
display: none;

values:
interlanguage-link interwiki-nap
display: none;

values:
interlanguage-link interwiki-ce
display: none;

values:
https://no.wikipedia.org/wiki/Torsdag
Torsdag – Norwegian
no
no
interlanguage-link-target

values:
interlanguage-link interwiki-no
display: none;

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

values:
interlanguage-link interwiki-nn
display: none;

values:
https://nrm.wikipedia.org/wiki/Jeudi
Jeudi – Norman
nrm
nrm
interlanguage-link-target

values:
interlanguage-link interwiki-nrm
display: none;

values:
interlanguage-link interwiki-oc
display: none;

values:
interlanguage-link interwiki-mhr
display: none;

values: ଗୁରୁବାର – Odia

values:
interlanguage-link interwiki-or
display: none;

values:
https://uz.wikipedia.org/wiki/Payshanba
Payshanba – Uzbek
uz
uz
interlanguage-link-target

values:
interlanguage-link interwiki-uz
display: none;

values: ਵੀਰਵਾਰ – Punjabi

values:
interlanguage-link interwiki-pa
display: none;

values:
https://pag.wikipedia.org/wiki/Huwebes
Huwebes – Pangasinan
pag
pag
interlanguage-link-target

values:
interlanguage-link interwiki-pag
display: none;

values:
interlanguage-link interwiki-pnb
display: none;

values: زيارت – Pashto

values:
interlanguage-link interwiki-ps
display: none;

values:
interlanguage-link interwiki-koi
display: none;

values: ថ្ងៃព្រហស្បតិ៍ – Khmer

values:
interlanguage-link interwiki-km
display: none;

values:
https://tpi.wikipedia.org/wiki/Fonde
Fonde – Tok Pisin
tpi
tpi
interlanguage-link-target

values:
interlanguage-link interwiki-tpi
display: none;

values:
interlanguage-link interwiki-nds
display: none;

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

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

values:
interlanguage-link interwiki-pnt
display: none;

values:
interlanguage-link interwiki-ksh
display: none;

values:
https://ro.wikipedia.org/wiki/Joi
Joi – Romanian
ro
ro
interlanguage-link-target

values:
interlanguage-link interwiki-ro
display: none;

values:
interlanguage-link interwiki-qu
display: none;

values: Четверь – Rusyn

values:
interlanguage-link interwiki-rue
display: none;

values: Четверг – Russian

values:
https://sm.wikipedia.org/wiki/Aso_Tofi
Aso Tofi – Samoan
sm
sm
interlanguage-link-target

values:
interlanguage-link interwiki-sm
display: none;

values: गुरुवासरः – Sanskrit

values:
interlanguage-link interwiki-sa
display: none;

values:
https://sc.wikipedia.org/wiki/Gioja
Gioja – Sardinian
sc
sc
interlanguage-link-target

values:
interlanguage-link interwiki-sc
display: none;

values:
https://sco.wikipedia.org/wiki/Fuirsday
Fuirsday – Scots
sco
sco
interlanguage-link-target

values:
interlanguage-link interwiki-sco
display: none;

values:
https://nso.wikipedia.org/wiki/Labone
Labone – Northern Sotho
nso
nso
interlanguage-link-target

values:
interlanguage-link interwiki-nso
display: none;

values:
https://sq.wikipedia.org/wiki/E_enjtja
E enjtja – Albanian
sq
sq
interlanguage-link-target

values:
interlanguage-link interwiki-sq
display: none;

values: බ්‍රහස්පතින්දා – Sinhala

values:
interlanguage-link interwiki-si
display: none;

values:
interlanguage-link interwiki-simple
display: none;

values:
interlanguage-link interwiki-sd
display: none;

values:
interlanguage-link interwiki-sk
display: none;

values:
interlanguage-link interwiki-sl
display: none;

values:
interlanguage-link interwiki-cu
display: none;

values:
interlanguage-link interwiki-szl
display: none;

values:
https://so.wikipedia.org/wiki/Khamiis
Khamiis – Somali
so
so
interlanguage-link-target

values:
interlanguage-link interwiki-so
display: none;

values:
interlanguage-link interwiki-ckb
display: none;

values: Четвртак – Serbian

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

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

values:
https://su.wikipedia.org/wiki/Kemis
Kemis – Sundanese
su
su
interlanguage-link-target

values:
interlanguage-link interwiki-su
display: none;

values:
https://fi.wikipedia.org/wiki/Torstai
Torstai – Finnish
fi
fi
interlanguage-link-target

values:
interlanguage-link interwiki-fi
display: none;

values:
https://sv.wikipedia.org/wiki/Torsdag
Torsdag – Swedish
sv
sv
interlanguage-link-target

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

values:
https://tl.wikipedia.org/wiki/Huwebes
Huwebes – Tagalog
tl
tl
interlanguage-link-target

values:
interlanguage-link interwiki-tl
display: none;

values: வியாழன் (கிழமை) – Tamil

values:
interlanguage-link interwiki-ta
display: none;

values:
https://kab.wikipedia.org/wiki/Amhad
Amhad – Kabyle
kab
kab
interlanguage-link-target

values:
interlanguage-link interwiki-kab
display: none;

values:
interlanguage-link interwiki-roa-tara
display: none;

values: Пәнҗешәмбе – Tatar

values:
interlanguage-link interwiki-tt
display: none;

values: గురువారము – Telugu

values:
interlanguage-link interwiki-te
display: none;

values: วันพฤหัสบดี – Thai

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

values:
interlanguage-link interwiki-ti
display: none;

values: Панҷшанбе – Tajik

values:
interlanguage-link interwiki-tg
display: none;

values:
interlanguage-link interwiki-chr
display: none;

values:
interlanguage-link interwiki-tr
display: none;

values:
interlanguage-link interwiki-tk
display: none;

values:
https://tw.wikipedia.org/wiki/Yawoada
Yawoada – Twi
tw
tw
interlanguage-link-target

values:
interlanguage-link interwiki-tw
display: none;

values: Дөрткү хүн – Tuvinian

values:
interlanguage-link interwiki-tyv
display: none;

values:
https://bug.wikipedia.org/wiki/Kamisi
Kamisi – Buginese
bug
bug
interlanguage-link-target

values:
interlanguage-link interwiki-bug
display: none;

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

values: جمعرات – Urdu

values: پەيشەنبە – Uyghur

values:
interlanguage-link interwiki-ug
display: none;

values:
interlanguage-link interwiki-za
display: none;

values:
https://vec.wikipedia.org/wiki/Zioba
Zioba – Venetian
vec
vec
interlanguage-link-target

values:
interlanguage-link interwiki-vec
display: none;

values:
interlanguage-link interwiki-vep
display: none;

values:
interlanguage-link interwiki-vi
display: none;

values:
interlanguage-link interwiki-vo
display: none;

values:
interlanguage-link interwiki-fiu-vro
display: none;

values:
https://wa.wikipedia.org/wiki/Djudi
Djudi – Walloon
wa
wa
interlanguage-link-target

values:
interlanguage-link interwiki-wa
display: none;

values:
interlanguage-link interwiki-vls
display: none;

values:
https://war.wikipedia.org/wiki/Huwebes
Huwebes – Waray
war
war
interlanguage-link-target

values:
interlanguage-link interwiki-war
display: none;

values:
https://wo.wikipedia.org/wiki/Alxames
Alxames – Wolof
wo
wo
interlanguage-link-target

values:
interlanguage-link interwiki-wo
display: none;

values:
interlanguage-link interwiki-wuu
display: none;

values: דאנערשטיק – Yiddish

values:
interlanguage-link interwiki-yi
display: none;

values:
interlanguage-link interwiki-yo
display: none;

values:
interlanguage-link interwiki-zh-yue
display: none;

values:
interlanguage-link interwiki-diq
display: none;

values:
interlanguage-link interwiki-bat-smg
display: none;

values: बिहीवार – Doteli

values:
interlanguage-link interwiki-dty
display: none;

values:
interlanguage-link interwiki-kbp
display: none;

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

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

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-Thursday rootpage-Thursday skin-vector action-view


text:
DNA replication
From Wikipedia, the free encyclopedia
Jump to: navigation, search
DNA replication: The double helix is un'zipped' and unwound, then each separated strand (turquoise) acts as a template for replicating a new partner strand (green). Nucleotides (bases) are matched to synthesize the new partner strands into two new double helices.
In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. This process occurs in all living organisms and is the basis for biological inheritance. The cell possesses the distinctive property of division, which makes replication of DNA essential.
DNA is made up of a double helix of two complementary strands. During replication, these strands are separated. Each strand of the original DNA molecule then serves as a template for the production of its counterpart, a process referred to as semiconservative replication. Cellular proofreading and error-checking mechanisms ensure near perfect fidelity for DNA replication.[1][2]
In a cell, DNA replication begins at specific locations, or origins of replication, in the genome.[3] Unwinding of DNA at the origin and synthesis of new strands results in replication forks growing bi-directionally from the origin. A number of proteins are associated with the replication fork to help in the initiation and continuation of DNA synthesis. Most prominently, DNA polymerase synthesizes the new strands by adding nucleotides that complement each (template) strand. DNA replication occurs during the S-stage of interphase.
DNA replication can also be performed in vitro (artificially, outside a cell). DNA polymerases isolated from cells and artificial DNA primers can be used to initiate DNA synthesis at known sequences in a template DNA molecule. The polymerase chain reaction (PCR), a common laboratory technique, cyclically applies such artificial synthesis to amplify a specific target DNA fragment from a pool of DNA.
Contents
[hide]
1 DNA structures
2 DNA polymerase
3 Replication process
3.1 Initiation
3.2 Elongation
3.3 Replication fork
3.3.1 Leading strand
3.3.2 Lagging strand
3.3.3 Dynamics at the replication fork
3.4 DNA replication proteins
3.5 Replication machinery
3.6 Termination
4 Regulation
4.1 Eukaryotes
4.1.1 Replication focus
4.2 Bacteria
5 Polymerase chain reaction
6 See also
7 Notes
8 References
DNA structures[edit]
DNA usually exists as a double-stranded structure, with both strands coiled together to form the characteristic double-helix. Each single strand of DNA is a chain of four types of nucleotides. Nucleotides in DNA contain a deoxyribose sugar, a phosphate, and a nucleobase. The four types of nucleotide correspond to the four nucleobases adenine, cytosine, guanine, and thymine, commonly abbreviated as A, C, G and T. Adenine and guanine are purine bases, while cytosine and thymine are pyrimidines. These nucleotides form phosphodiester bonds, creating the phosphate-deoxyribose backbone of the DNA double helix with the nuclei bases pointing inward (i.e., toward the opposing strand). Nucleotides (bases) are matched between strands through hydrogen bonds to form base pairs. Adenine pairs with thymine (two hydrogen bonds), and guanine pairs with cytosine (stronger: three hydrogen bonds).
DNA strands have a directionality, and the different ends of a single strand are called the "3' (three-prime) end" and the "5' (five-prime) end". By convention, if the base sequence of a single strand of DNA is given, the left end of the sequence is the 5' end, while the right end of the sequence is the 3' end. The strands of the double helix are anti-parallel with one being 5' to 3', and the opposite strand 3' to 5'. These terms refer to the carbon atom in deoxyribose to which the next phosphate in the chain attaches. Directionality has consequences in DNA synthesis, because DNA polymerase can synthesize DNA in only one direction by adding nucleotides to the 3' end of a DNA strand.
The pairing of complementary bases in DNA (through hydrogen bonding) means that the information contained within each strand is redundant. Phosphodiester (intra-strand) bonds are stronger than hydrogen (inter-strand) bonds. This allows the strands to be separated from one another. The nucleotides on a single strand can therefore be used to reconstruct nucleotides on a newly synthesized partner strand.[4]
DNA polymerase[edit]
Main article: DNA polymerase
DNA polymerases adds nucleotides to the 3' end of a strand of DNA.[5] If a mismatch is accidentally incorporated, the polymerase is inhibited from further extension. Proofreading removes the mismatched nucleotide and extension continues.
DNA polymerases are a family of enzymes that carry out all forms of DNA replication.[6] DNA polymerases in general cannot initiate synthesis of new strands, but can only extend an existing DNA or RNA strand paired with a template strand. To begin synthesis, a short fragment of RNA, called a primer, must be created and paired with the template DNA strand.
DNA polymerase adds a new strand of DNA by extending the 3' end of an existing nucleotide chain, adding new nucleotides matched to the template strand one at a time via the creation of phosphodiester bonds. The energy for this process of DNA polymerization comes from hydrolysis of the high-energy phosphate (phosphoanhydride) bonds between the three phosphates attached to each unincorporated base. Free bases with their attached phosphate groups are called nucleotides; in particular, bases with three attached phosphate groups are called nucleoside triphosphates. When a nucleotide is being added to a growing DNA strand, the formation of a phosphodiester bond between the proximal phosphate of the nucleotide to the growing chain is accompanied by hydrolysis of a high-energy phosphate bond with release of the two distal phosphates as a pyrophosphate. Enzymatic hydrolysis of the resulting pyrophosphate into inorganic phosphate consumes a second high-energy phosphate bond and renders the reaction effectively irreversible.[Note 1]
In general, DNA polymerases are highly accurate, with an intrinsic error rate of less than one mistake for every 107 nucleotides added.[7] In addition, some DNA polymerases also have proofreading ability; they can remove nucleotides from the end of a growing strand in order to correct mismatched bases. Finally, post-replication mismatch repair mechanisms monitor the DNA for errors, being capable of distinguishing mismatches in the newly synthesized DNA strand from the original strand sequence. Together, these three discrimination steps enable replication fidelity of less than one mistake for every 109 nucleotides added.[7]
The rate of DNA replication in a living cell was first measured as the rate of phage T4 DNA elongation in phage-infected E. coli.[8] During the period of exponential DNA increase at 37 °C, the rate was 749 nucleotides per second. The mutation rate per base pair per replication during phage T4 DNA synthesis is 1.7 per 108.[9]
Replication process[edit]
Main articles: Prokaryotic DNA replication and Eukaryotic DNA replication
DNA replication, like all biological polymerization processes, proceeds in three enzymatically catalyzed and coordinated steps: initiation, elongation and termination.
Initiation[edit]
Role of initiators for initiation of DNA replication.
Formation of pre-replication complex.
For a cell to divide, it must first replicate its DNA.[10] This process is initiated at particular points in the DNA, known as "origins", which are targeted by initiator proteins.[3] In E. coli this protein is DnaA; in yeast, this is the origin recognition complex.[11] Sequences used by initiator proteins tend to be "AT-rich" (rich in adenine and thymine bases), because A-T base pairs have two hydrogen bonds (rather than the three formed in a C-G pair) and thus are easier to strand separate.[12] Once the origin has been located, these initiators recruit other proteins and form the pre-replication complex, which unzips the double-stranded DNA.
Elongation[edit]
DNA polymerase has 5'-3' activity. All known DNA replication systems require a free 3' hydroxyl group before synthesis can be initiated (note: the DNA template is read in 3' to 5' direction whereas a new strand is synthesized in the 5' to 3' direction—this is often confused). Four distinct mechanisms for DNA synthesis are recognized:
All cellular life forms and many DNA viruses, phages and plasmids use a primase to synthesize a short RNA primer with a free 3' OH group which is subsequently elongated by a DNA polymerase.
The retroelements (including retroviruses) employ a transfer RNA that primes DNA replication by providing a free 3′ OH that is used for elongation by the reverse transcriptase.
In the adenoviruses and the φ29 family of bacteriophages, the 3' OH group is provided by the side chain of an amino acid of the genome attached protein (the terminal protein) to which nucleotides are added by the DNA polymerase to form a new strand.
In the single stranded DNA viruses — a group that includes the circoviruses, the geminiviruses, the parvoviruses and others — and also the many phages and plasmids that use the rolling circle replication (RCR) mechanism, the RCR endonuclease creates a nick in the genome strand (single stranded viruses) or one of the DNA strands (plasmids). The 5′ end of the nicked strand is transferred to a tyrosine residue on the nuclease and the free 3′ OH group is then used by the DNA polymerase to synthesize the new strand.
The first is the best known of these mechanisms and is used by the cellular organisms. In this mechanism, once the two strands are separated, primase adds RNA primers to the template strands. The leading strand receives one RNA primer while the lagging strand receives several. The leading strand is continuously extended from the primer by a DNA polymerase with high processivity, while the lagging strand is extended discontinuously from each primer forming Okazaki fragments. RNase removes the primer RNA fragments, and a low processivity DNA polymerase distinct from the replicative polymerase enters to fill the gaps. When this is complete, a single nick on the leading strand and several nicks on the lagging strand can be found. Ligase works to fill these nicks in, thus completing the newly replicated DNA molecule.
The primase used in this process differs significantly between bacteria and archaea/eukaryotes. Bacteria use a primase belonging to the DnaG protein superfamily which contains a catalytic domain of the TOPRIM fold type.[13] The TOPRIM fold contains an α/β core with four conserved strands in a Rossmann-like topology. This structure is also found in the catalytic domains of topoisomerase Ia, topoisomerase II, the OLD-family nucleases and DNA repair proteins related to the RecR protein.
The primase used by archaea and eukaryotes, in contrast, contains a highly derived version of the RNA recognition motif (RRM). This primase is structurally similar to many viral RNA-dependent RNA polymerases, reverse transcriptases, cyclic nucleotide generating cyclases and DNA polymerases of the A/B/Y families that are involved in DNA replication and repair. In eukaryotic replication, the primase forms a complex with Pol α.[14]
Multiple DNA polymerases take on different roles in the DNA replication process. In E. coli, DNA Pol III is the polymerase enzyme primarily responsible for DNA replication. It assembles into a replication complex at the replication fork that exhibits extremely high processivity, remaining intact for the entire replication cycle. In contrast, DNA Pol I is the enzyme responsible for replacing RNA primers with DNA. DNA Pol I has a 5' to 3' exonuclease activity in addition to its polymerase activity, and uses its exonuclease activity to degrade the RNA primers ahead of it as it extends the DNA strand behind it, in a process called nick translation. Pol I is much less processive than Pol III because its primary function in DNA replication is to create many short DNA regions rather than a few very long regions.
In eukaryotes, the low-processivity enzyme, Pol α, helps to initiate replication because it forms a complex with primase.[15] In eukaryotes, leading strand synthesis is thought to be conducted by Pol ε; however, this view has recently been challenged, suggesting a role for Pol δ.[16] Primer removal is completed Pol δ[17] while repair of DNA during replication is completed by Pol ε.
As DNA synthesis continues, the original DNA strands continue to unwind on each side of the bubble, forming a replication fork with two prongs. In bacteria, which have a single origin of replication on their circular chromosome, this process creates a "theta structure" (resembling the Greek letter theta: θ). In contrast, eukaryotes have longer linear chromosomes and initiate replication at multiple origins within these.[18]
Replication fork[edit]
Scheme of the replication fork.
a: template, b: leading strand, c: lagging strand, d: replication fork, e: primer, f: Okazaki fragments
Many enzymes are involved in the DNA replication fork.
The replication fork is a structure that forms within the nucleus during DNA replication. It is created by helicases, which break the hydrogen bonds holding the two DNA strands together. The resulting structure has two branching "prongs", each one made up of a single strand of DNA. These two strands serve as the template for the leading and lagging strands, which will be created as DNA polymerase matches complementary nucleotides to the templates; the templates may be properly referred to as the leading strand template and the lagging strand template.
DNA is always synthesized in the 5' to 3' direction. Since the leading and lagging strand templates are oriented in opposite directions at the replication fork, a major issue is how to achieve synthesis of nascent (new) lagging strand DNA, whose direction of synthesis is opposite to the direction of the growing replication fork.
Leading strand[edit]
The leading strand is the strand of nascent DNA which is being synthesized in the same direction as the growing replication fork. A polymerase "reads" the leading strand template and adds complementary nucleotides to the nascent leading strand on a continuous basis.
Lagging strand[edit]
The lagging strand is the strand of nascent DNA whose direction of synthesis is opposite to the direction of the growing replication fork. Because of its orientation, replication of the lagging strand is more complicated as compared to that of the leading strand. As a consequence, the DNA polymerase on this strand is seen to "lag behind" the other strand.
The lagging strand is synthesized in short, separated segments. On the lagging strand template, a primase "reads" the template DNA and initiates synthesis of a short complementary RNA primer. A DNA polymerase extends the primed segments, forming Okazaki fragments. The RNA primers are then removed and replaced with DNA, and the fragments of DNA are joined together by DNA ligase.
Dynamics at the replication fork[edit]
The assembled human DNA clamp, a trimer of the protein PCNA.
As helicase unwinds DNA at the replication fork, the DNA ahead is forced to rotate. This process results in a build-up of twists in the DNA ahead.[19] This build-up forms a torsional resistance that would eventually halt the progress of the replication fork. Topoisomerases are enzymes that temporarily break the strands of DNA, relieving the tension caused by unwinding the two strands of the DNA helix; topoisomerases (including DNA gyrase) achieve this by adding negative supercoils to the DNA helix.[20]
Bare single-stranded DNA tends to fold back on itself forming secondary structures; these structures can interfere with the movement of DNA polymerase. To prevent this, single-strand binding proteins bind to the DNA until a second strand is synthesized, preventing secondary structure formation.[21]
Clamp proteins form a sliding clamp around DNA, helping the DNA polymerase maintain contact with its template, thereby assisting with processivity. The inner face of the clamp enables DNA to be threaded through it. Once the polymerase reaches the end of the template or detects double-stranded DNA, the sliding clamp undergoes a conformational change that releases the DNA polymerase. Clamp-loading proteins are used to initially load the clamp, recognizing the junction between template and RNA primers.[2]:274-5
DNA replication proteins[edit]
At the replication fork, many replication enzymes assemble on the DNA into a complex molecular machine called the replisome. The following is a list of major DNA replication enzymes that participate in the replisome:[22]
Enzyme
Function in DNA replication
DNA Helicase Also known as helix destabilizing enzyme. Helicase separates the two strands of DNA at the Replication Fork behind the topoisomerase.
DNA Polymerase The enzyme responsible for catalyzing the addition of nucleotide substrates to DNA in the 5' to 3' direction during DNA replication. Also performs proof-reading and error correction. There exist many different types of DNA Polymerase, (such as the DNA Polymerase III )each of which perform different functions in different types of cells.
DNA clamp A protein which prevents elongating DNA polymerases from dissociating from the DNA parent strand.
Single-Strand Binding (SSB) Proteins Bind to ssDNA and prevent the DNA double helix from re-annealing after DNA helicase unwinds it, thus maintaining the strand separation, and facilitating the synthesis of the nascent strand.
Topoisomerase Relaxes the DNA from its super-coiled nature.
DNA Gyrase Relieves strain of unwinding by DNA helicase; this is a specific type of topoisomerase
DNA Ligase Re-anneals the semi-conservative strands and joins Okazaki Fragments of the lagging strand.
Primase Provides a starting point of RNA (or DNA) for DNA polymerase to begin synthesis of the new DNA strand.
Telomerase Lengthens telomeric DNA by adding repetitive nucleotide sequences to the ends of eukaryotic chromosomes. This allows germ cells and stem cells to avoid the Hayflick limit on cell division.[23]
Replication machinery[edit]
Replication machineries consist of factors involved in DNA replication and appearing on template ssDNAs. Replication machineries include primosotors are replication enzymes; DNA polymerase, DNA helicases, DNA clamps and DNA topoisomerases, and replication proteins; e.g. single-stranded DNA binding proteins (SSB). In the replication machineries these components coordinate. In most of the bacteria, all of the factors involved in DNA replication are located on replication forks and the complexes stay on the forks during DNA replication. These replication machineries are called replisomes or DNA replicase systems. These terms are generic terms for proteins located on replication forks. In eukaryotic and some bacterial cells the replisomes are not formed.
Since replication machineries do not move relatively to template DNAs such as factories, they are called a replication factory.[24] In an alternative figure, DNA factories are similar to projectors and DNAs are like as cinematic films passing constantly into the projectors. In the replication factory model, after both DNA helicases for leading strands and lagging strands are loaded on the template DNAs, the helicases run along the DNAs into each other. The helicases remain associated for the remainder of replication process. Peter Meister et al. observed directly replication sites in budding yeast by monitoring green fluorescent protein(GFP)-tagged DNA polymerases α. They detected DNA replication of pairs of the tagged loci spaced apart symmetrically from a replication origin and found that the distance between the pairs decreased markedly by time.[25] This finding suggests that the mechanism of DNA replication goes with DNA factories. That is, couples of replication factories are loaded on replication origins and the factories associated with each other. Also, template DNAs move into the factories, which bring extrusion of the template ssDNAs and nascent DNAs. Meister’s finding is the first direct evidence of replication factory model. Subsequent research has shown that DNA helicases form dimers in many eukaryotic cells and bacterial replication machineries stay in single intranuclear location during DNA synthesis.[24]
The replication factories perform disentanglement of sister chromatids. The disentanglement is essential for distributing the chromatids into daughter cells after DNA replication. Because sister chromatids after DNA replication hold each other by Cohesin rings, there is the only chance for the disentanglement in DNA replication. Fixing of replication machineries as replication factories can improve the success rate of DNA replication. If replication forks move freely in chromosomes, catenation of nuclei is aggravated and impedes mitotic segregation.[25]
Termination[edit]
Eukaryotes initiate DNA replication at multiple points in the chromosome, so replication forks meet and terminate at many points in the chromosome. Because eukaryotes have linear chromosomes, DNA replication is unable to reach the very end of the chromosomes. Due to this problem, DNA is lost each replication cycle from the end of the chromosome. Telomeres are regions of repetitive DNA close to the ends and help prevent loss of genes due to this shortening. Shortening of the telomeres is a normal process in somatic cells. This shortens the telomeres of the daughter DNA chromosome. As a result, cells can only divide a certain number of times before the DNA loss prevents further division. (This is known as the Hayflick limit.) Within the germ cell line, which passes DNA to the next generation, telomerase extends the repetitive sequences of the telomere region to prevent degradation. Telomerase can become mistakenly active in somatic cells, sometimes leading to cancer formation. Increased telomerase activity is one of the hallmarks of cancer.
Termination requires that the progress of the DNA replication fork must stop or be blocked. Termination at a specific locus, when it occurs, involves the interaction between two components: (1) a termination site sequence in the DNA, and (2) a protein which binds to this sequence to physically stop DNA replication. In various bacterial species, this is named the DNA replication terminus site-binding protein, or Ter protein.
Because bacteria have circular chromosomes, termination of replication occurs when the two replication forks meet each other on the opposite end of the parental chromosome. E. coli regulates this process through the use of termination sequences that, when bound by the Tus protein, enable only one direction of replication fork to pass through. As a result, the replication forks are constrained to always meet within the termination region of the chromosome.[26]
Regulation[edit]
Main articles: Cell division and Cell cycle
The cell cycle of eukaryotic cells.
Eukaryotes[edit]
Within eukaryotes, DNA replication is controlled within the context of the cell cycle. As the cell grows and divides, it progresses through stages in the cell cycle; DNA replication takes place during the S phase (synthesis phase). The progress of the eukaryotic cell through the cycle is controlled by cell cycle checkpoints. Progression through checkpoints is controlled through complex interactions between various proteins, including cyclins and cyclin-dependent kinases.[27] Unlike bacteria, eukaryotic DNA replicates in the confines of the nucleus.[28]
The G1/S checkpoint (or restriction checkpoint) regulates whether eukaryotic cells enter the process of DNA replication and subsequent division. Cells that do not proceed through this checkpoint remain in the G0 stage and do not replicate their DNA.
Replication of chloroplast and mitochondrial genomes occurs independently of the cell cycle, through the process of D-loop replication.
Replication focus[edit]
In vertebrate cells, replication sites concentrate into positions called replication foci.[25] Replication sites can be detected by immunostaining daughter strands and replication enzymes and monitoring GFP-tagged replication factors. By these methods it is found that replication foci of varying size and positions appear in S phase of cell division and their number per nucleus is far smaller than the number of genomic replication forks.
P. Heun et al.(2001) tracked GFP-tagged replication foci in budding yeast cells and revealed that replication origins move constantly in G1 and S phase and the dynamics decreased significantly in S phase.[25] Traditionally, replication sites were fixed on spatial structure of chromosomes by nuclear matrix or lamins. The Heun’s results denied the traditional concepts, budding yeasts don't have lamins, and support that replication origins self-assemble and form replication foci.
By firing of replication origins, controlled spatially and temporally, the formation of replication foci is regulated. D. A. Jackson et al.(1998) revealed that neighboring origins fire simultaneously in mammalian cells.[25] Spatial juxtaposition of replication sites brings clustering of replication forks. The clustering do rescue of stalled replication forks and favors normal progress of replication forks. Progress of replication forks is inhibited by many factors; collision with proteins or with complexes binding strongly on DNA, deficiency of dNTPs, nicks on template DNAs and so on. If replication forks stall and the remaining sequences from the stalled forks are not replicated, the daughter strands have nick obtained un-replicated sites. The un-replicated sites on one parent's strand hold the other strand together but not daughter strands. Therefore, the resulting sister chromatids cannot separate from each other and cannot divide into 2 daughter cells. When neighboring origins fire and a fork from one origin is stalled, fork from other origin access on an opposite direction of the stalled fork and duplicate the un-replicated sites. As other mechanism of the rescue there is application of dormant replication origins that excess origins don't fire in normal DNA replication.
Bacteria[edit]
Dam methylates adenine of GATC sites after replication.
Most bacteria do not go through a well-defined cell cycle but instead continuously copy their DNA; during rapid growth, this can result in the concurrent occurrence of multiple rounds of replication.[29] In E. coli, the best-characterized bacteria, DNA replication is regulated through several mechanisms, including: the hemimethylation and sequestering of the origin sequence, the ratio of adenosine triphosphate (ATP) to adenosine diphosphate (ADP), and the levels of protein DnaA. All these control the binding of initiator proteins to the origin sequences.
Because E. coli methylates GATC DNA sequences, DNA synthesis results in hemimethylated sequences. This hemimethylated DNA is recognized by the protein SeqA, which binds and sequesters the origin sequence; in addition, DnaA (required for initiation of replication) binds less well to hemimethylated DNA. As a result, newly replicated origins are prevented from immediately initiating another round of DNA replication.[30]
ATP builds up when the cell is in a rich medium, triggering DNA replication once the cell has reached a specific size. ATP competes with ADP to bind to DnaA, and the DnaA-ATP complex is able to initiate replication. A certain number of DnaA proteins are also required for DNA replication — each time the origin is copied, the number of binding sites for DnaA doubles, requiring the synthesis of more DnaA to enable another initiation of replication.
See also: Plasmid, Plasmid copy number, and Plasmid partition system
Polymerase chain reaction[edit]
Main article: Polymerase chain reaction
Researchers commonly replicate DNA in vitro using the polymerase chain reaction (PCR). PCR uses a pair of primers to span a target region in template DNA, and then polymerizes partner strands in each direction from these primers using a thermostable DNA polymerase. Repeating this process through multiple cycles amplifies the targeted DNA region. At the start of each cycle, the mixture of template and primers is heated, separating the newly synthesized molecule and template. Then, as the mixture cools, both of these become templates for annealing of new primers, and the polymerase extends from these. As a result, the number of copies of the target region doubles each round, increasing exponentially.[31]
See also[edit]
Wikimedia Commons has media related to DNA replication.
Autopoiesis
Data storage device
Self-replication
Notes[edit]
Jump up ^ The energetics of this process may also help explain the directionality of synthesis—if DNA were synthesized in the 3' to 5' direction, the energy for the process would come from the 5' end of the growing strand rather than from free nucleotides. The problem is that if the high energy triphosphates were on the growing strand and not on the free nucleotides, proof-reading by removing a mismatched terminal nucleotide would be problematic: Once a nucleotide is added, the triphosphate is lost and a single phosphate remains on the backbone between the new nucleotide and the rest of the strand. If the added nucleotide were mismatched, removal would result in a DNA strand terminated by a monophosphate at the end of the "growing strand" rather than a high energy triphosphate. So strand would be stuck and wouldn't be able to grow anymore. In actuality, the high energy triphosphates hydrolyzed at each step originate from the free nucleotides, not the polymerized strand, so this issue does not exist.
References[edit]
Jump up ^ Imperfect DNA replication results in mutations. Berg JM, Tymoczko JL, Stryer L, Clarke ND (2002). "Chapter 27: DNA Replication, Recombination, and Repair". Biochemistry. W.H. Freeman and Company. ISBN 0-7167-3051-0. External link in |chapter= (help)
^ Jump up to: a b Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "5DNA Replication, Repair, and Recombination". Molecular Biology of the Cell. Garland Science. ISBN 0-8153-3218-1. External link in |chapter= (help)
^ Jump up to: a b Berg JM, Tymoczko JL, Stryer L, Clarke ND (2002). "Chapter 27, Section 4: DNA Replication of Both Strands Proceeds Rapidly from Specific Start Sites". Biochemistry. W.H. Freeman and Company. ISBN 0-7167-3051-0. External link in |chapter= (help)
Jump up ^ Alberts, B., et al., Molecular Biology of the Cell, Garland Science, 4th ed., 2002, pp. 238–240 ISBN 0-8153-3218-1
Jump up ^ Allison, Lizabeth A. Fundamental Molecular Biology. Blackwell Publishing. 2007. p.112 ISBN 978-1-4051-0379-4
Jump up ^ Berg JM, Tymoczko JL, Stryer L, Clarke ND (2002). Biochemistry. W.H. Freeman and Company. ISBN 0-7167-3051-0. Chapter 27, Section 2: DNA Polymerases Require a Template and a Primer
^ Jump up to: a b McCulloch, Scott D; Kunkel, Thomas A (January 2008). "The fidelity of DNA synthesis by eukaryotic replicative and translesion synthesis polymerases". Cell Research. 18 (1): 148–161. PMC 3639319  . PMID 18166979. doi:10.1038/cr.2008.4.
Jump up ^ McCarthy, David; Minner, Charles; Bernstein, Harris; Bernstein, Carol (October 1976). "DNA elongation rates and growing point distributions of wild-type phage T4 and a DNA-delay amber mutant". Journal of Molecular Biology. 106 (4): 963–981. PMID 789903. doi:10.1016/0022-2836(76)90346-6.
Jump up ^ Drake JW (1970) The Molecular Basis of Mutation. Holden-Day, San Francisco ISBN 0816224501 ISBN 978-0816224500
Jump up ^ Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the Cell. Garland Science. ISBN 0-8153-3218-1. Chapter 5: DNA Replication Mechanisms
Jump up ^ Weigel C, Schmidt A, Rückert B, Lurz R, Messer W (November 1997). "DnaA protein binding to individual DnaA boxes in the Escherichia coli replication origin, oriC". The EMBO Journal. 16 (21): 6574–83. PMC 1170261  . PMID 9351837. doi:10.1093/emboj/16.21.6574.
Jump up ^ Lodish H, Berk A, Zipursky LS, Matsudaira P, Baltimore D, Darnell J (2000). Molecular Cell Biology. W. H. Freeman and Company. ISBN 0-7167-3136-3. 12.1. General Features of Chromosomal Replication: Three Common Features of Replication Origins
Jump up ^ Aravind, L.; Leipe, D. D.; Koonin, E. V. (1998). "Toprim—a conserved catalytic domain in type IA and II topoisomerases, DnaG-type primases, OLD family nucleases and RecR proteins". Nucleic Acids Research. 26 (18): 4205–4213. PMC 147817  . PMID 9722641. doi:10.1093/nar/26.18.4205.
Jump up ^ Frick, David; Richardson, Charles (July 2001). "DNA Primases". Annual Review of Biochemistry. 70: 39–80. PMID 11395402. doi:10.1146/annurev.biochem.70.1.39.
Jump up ^ Barry, Elizabeth R.; Bell, Stephen D. (8 December 2006). "DNA Replication in the Archaea". Microbiology and Molecular Biology Reviews. 70 (4): 876–887. PMC 1698513  . PMID 17158702. doi:10.1128/MMBR.00029-06. |access-date= requires |url= (help)
Jump up ^ Stillman, Bruce (July 2015). "Reconsidering DNA Polymerases at the Replication Fork in Eukaryotes". Molecular Cell. 59 (2): 139–141. PMC 4636199  . PMID 26186286. doi:10.1016/j.molcel.2015.07.004. |access-date= requires |url= (help)
Jump up ^ Distinguishing the pathways of primer removal during Eukaryotic Okazaki fragment maturation Contributor Author Rossi, Marie Louise. Date Accessioned: 2009-02-23T17:05:09Z. Date Available: 2009-02-23T17:05:09Z. Date Issued: 2009-02-23T17:05:09Z. Identifier Uri: http://hdl.handle.net/1802/6537. Description: Dr. Robert A. Bambara, Faculty Advisor. Thesis (PhD) – School of Medicine and Dentistry, University of Rochester. UR only until January 2010. UR only until January 2010.
Jump up ^ Huberman, Joel A.; Riggs, Arthur D. (March 1968). "On the mechanism of DNA replication in mammalian chromosomes". Journal of Molecular Biology. 32 (2): 327–341. PMID 5689363. doi:10.1016/0022-2836(68)90013-2.
Jump up ^ Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the Cell. Garland Science. ISBN 0-8153-3218-1. DNA Replication Mechanisms: DNA Topoisomerases Prevent DNA Tangling During Replication
Jump up ^ Reece, Richard J.; Maxwell, Anthony; Wang, James C. (26 September 2008). "DNA Gyrase: Structure and Function". Critical Reviews in Biochemistry and Molecular Biology. 26 (3-4): 335–375. PMID 1657531. doi:10.3109/10409239109114072.
Jump up ^ Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the Cell. Garland Science. ISBN 0-8153-3218-1. DNA Replication Mechanisms: Special Proteins Help to Open Up the DNA Double Helix in Front of the Replication Fork
Jump up ^ Griffiths A.J.F.; Wessler S.R.; Lewontin R.C.; Carroll S.B. (2008). Introduction to Genetic Analysis. W. H. Freeman and Company. ISBN 0-7167-6887-9. [Chapter 7: DNA: Structure and Replication. pg 283–290]
Jump up ^ "Will the Hayflick limit keep us from living forever?". Howstuffworks. Retrieved January 20, 2015.
^ Jump up to: a b James D. Watson et al. (2008), "Molecular Biology of the gene", Pearson Education: 237
^ Jump up to: a b c d e Peter Meister, Angela Taddei1, Susan M. Gasser(June 2006), "In and out of the Replication Factory", Cell 125 (7): 1233–1235
Jump up ^ TA Brown (2002). Genomes. BIOS Scientific Publishers. ISBN 1-85996-228-9. 13.2.3. Termination of replication
Jump up ^ Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the Cell. Garland Science. ISBN 0-8153-3218-1. Intracellular Control of Cell-Cycle Events: S-Phase Cyclin-Cdk Complexes (S-Cdks) Initiate DNA Replication Once Per Cycle
Jump up ^ Brown, TA (2002). "13". Genomes (2nd ed.). Oxford: Wiley-Liss.
Jump up ^ Tobiason DM, Seifert HS (2006). "The Obligate Human Pathogen, Neisseria gonorrhoeae, Is Polyploid". PLoS Biology. 4 (6): e185. PMC 1470461  . PMID 16719561. doi:10.1371/journal.pbio.0040185.
Jump up ^ Slater, Steven; Wold, Sture; Lu, Min; Boye, Erik; Skarstad, Kirsten; Kleckner, Nancy (September 1995). "E. coli SeqA protein binds oriC in two different methyl-modulated reactions appropriate to its roles in DNA replication initiation and origin sequestration". Cell. 82 (6): 927–936. PMID 7553853. doi:10.1016/0092-8674(95)90272-4.
Jump up ^ Saiki, Randall; Gelfand, David H.; Stoffel, Susanne; Scharf, Stephen J.; Higuchi, Russell; Horn, Glenn T.; Mullis, Kary B.; Erlich, Henry A. (29 January 1988). "Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase". Science. 239 (4839): 487–491. Bibcode:1988Sci...239..487S. PMID 2448875. doi:10.1126/science.2448875. Retrieved 7 April 2016.
[hide]
v
t
e
DNA replication (comparing Prokaryotic to Eukaryotic)
Initiation
Prokaryotic
(initiation)
Pre-replication complex
dnaC
Cdc6
Helicase
dnaA
dnaB
T7
Primase
dnaG
Eukaryotic
(preparation in
G1 phase)
Pre-replication complex
Origin recognition complex
ORC1
ORC2
ORC3
ORC4
ORC5
ORC6
Cdc6
Cdt1
Minichromosome maintenance
MCM2
MCM3
MCM4
MCM5
MCM6
MCM7
Licensing factor
Autonomously replicating sequence
Single-strand binding protein
SSBP2
SSBP3
SSBP4
RNase H
RNASEH1
RNASEH2A
Helicase: HFM1
Primase: PRIM1
PRIM2
Both
Origin of replication/Ori/Replicon
Replication fork
Lagging and leading strands
Okazaki fragments
Primer
Replication
Prokaryotic
(elongation)
DNA polymerase III holoenzyme
dnaC
dnaE
dnaH
dnaN
dnaQ
dnaT
dnaX
holA
holB
holC
holD
holE
Replisome
DNA ligase
DNA clamp
Topoisomerase
DNA gyrase
Prokaryotic DNA polymerase: DNA polymerase I
Klenow fragment
Eukaryotic
(synthesis in
S phase)
Replication factor C
RFC1
Flap endonuclease
FEN1
Topoisomerase
Replication protein A
RPA1
Eukaryotic DNA polymerase:
alpha
POLA1
POLA2
PRIM1
PRIM2
delta
POLD1
POLD2
POLD3
POLD4
epsilon
POLE
POLE2
POLE3
POLE4
DNA clamp
PCNA
Control of chromosome duplication
Both
Movement: Processivity
DNA ligase
Termination
Telomere: Telomerase
TERT
TERC
DKC1
Molecular and cellular biology portal
Authority control
GND: 4153174-7
Retrieved from "https://en.wikipedia.org/w/index.php?title=DNA_replication&oldid=804924944"
Categories:
DNA replication
Senescence
Hidden categories:
CS1 errors: external links
Pages using citations with accessdate and no URL
Wikipedia articles with GND identifiers
Navigation menu
Personal tools
Not logged in
Talk
Contributions
Create account
Log in
Namespaces
Article
Talk
Variants
Views
Read
Edit
View history
More
Search
Navigation
Main page
Contents
Featured content
Current events
Random article
Donate to Wikipedia
Wikipedia store
Interaction
Help
About Wikipedia
Community portal
Recent changes
Contact page
Tools
What links here
Related changes
Upload file
Special pages
Permanent link
Page information
Wikidata item
Cite this page
Print/export
Create a book
Download as PDF
Printable version
In other projects
Wikimedia Commons
Languages
العربية
Беларуская
Български
Bosanski
Català
Čeština
Dansk
Deutsch
Eesti
Ελληνικά
Español
Euskara
فارسی
Français
Galego
한국어
Հայերեն
Hrvatski
Bahasa Indonesia
Íslenska
Italiano
עברית
Basa Jawa
ქართული
Қазақша
Kreyòl ayisyen
Кыргызча
Lietuvių
Magyar
Македонски
Bahasa Melayu
Nederlands
日本語
Norsk
Occitan
Polski
Português
Română
Русский
Scots
Simple English
Slovenčina
Slovenščina
کوردی
Српски / srpski
Srpskohrvatski / српскохрватски
Suomi
Svenska
தமிழ்
ไทย
Türkçe
Українська
اردو
Tiếng Việt
中文
Edit links
This page was last edited on 11 October 2017, at 23:35.
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site, you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc., a non-profit organization.
Privacy policy
About Wikipedia
Disclaimers
Contact Wikipedia
Developers
Cookie statement
Mobile view
Enable previews

values: mw-page-base noprint

values: mw-head-base noprint

values: firstHeading firstHeading en

values: siteSub noprint

values: jump-to-nav mw-jump

values:
/wiki/Double_helix
mw-redirect
Double helix

attrs: class style

value: p

values:
/wiki/Proofreading_(Biology)
mw-redirect
Proofreading (Biology)

values:
/wiki/Replication_fork
mw-redirect
Replication fork

values:
/wiki/Phosphodiester_bonds
mw-redirect
Phosphodiester bonds

values:
/wiki/Hydrogen_bonding
mw-redirect
Hydrogen bonding

values:
/wiki/Okazaki_fragment
mw-redirect
Okazaki fragment

values: Edit section: Dynamics at the replication fork

values:
/wiki/Trimer_(biochemistry)
mw-redirect
Trimer (biochemistry)

values:
/wiki/Single-strand_binding_protein
mw-redirect
Single-strand binding protein

values:
/wiki/DNA_Helicase
mw-redirect
DNA Helicase

values:
/wiki/Replication_Fork
mw-redirect
Replication Fork

values:
/wiki/DNA_Polymerase
mw-redirect
DNA Polymerase

values:
/wiki/Okazaki_Fragments
mw-redirect
Okazaki Fragments

values:
/wiki/Budding_yeast
mw-redirect
Budding yeast

values:
/wiki/Tus_protein
mw-redirect
Tus protein

values:
/wiki/Data_storage_device
mw-redirect
Data storage device

values: -moz-column-width: 30em; -webkit-column-width: 30em; column-width: 30em; list-style-type: decimal;

attrs: alt src title width height srcset data-file-width data-file-height

values:
/wiki/PubMed_Identifier
mw-redirect
PubMed Identifier

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/18166979

values:
nofollow
external text
//doi.org/10.1038%2Fcr.2008.4

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/789903

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9351837

values:
nofollow
external text
//doi.org/10.1093%2Femboj%2F16.21.6574

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/9722641

values:
nofollow
external text
//doi.org/10.1093%2Fnar%2F26.18.4205

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/11395402

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/17158702

values:
nofollow
external text
//doi.org/10.1128%2FMMBR.00029-06

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/26186286

values:
nofollow
external text
http://hdl.handle.net/1802/6537

values:
nofollow
external free
http://hdl.handle.net/1802/6537

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/5689363

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/1657531

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/16719561

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/7553853

values:
nofollow
external text
//www.ncbi.nlm.nih.gov/pubmed/2448875

values:
nofollow
external text
//doi.org/10.1126%2Fscience.2448875

values:
/wiki/T7_DNA_Helicase
mw-redirect
T7 DNA Helicase

values:
/wiki/Leading_strand
mw-redirect
Leading strand

values:
/wiki/Prokaryotic_DNA_polymerase
mw-redirect
Prokaryotic DNA polymerase

values:
/wiki/Eukaryotic_DNA_polymerase
mw-redirect
Eukaryotic DNA polymerase

values: nowraplinks hlist collapsible autocollapse navbox-inner

values:
noprint metadata navbox
navigation
Portals
font-weight:bold;padding:0.4em 2em

values:
nofollow
external text
http://d-nb.info/gnd/4153174-7

values: Category:Pages using citations with accessdate and no URL

values: Category:Wikipedia articles with GND identifiers

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

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

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

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

values:
/wiki/DNA_replication
View the content page [ctrl-alt-c]
c

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

values: p-namespaces navigation vectorTabs p-namespaces-label

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

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

values: p-views navigation vectorTabs p-views-label

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

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

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

values: Featured content – the best of Wikipedia

values: Find background information on current events

values:
/wiki/Special:Random
Load a random article [ctrl-alt-x]
x

values: portal navigation p-navigation p-navigation-label

values: Guidance on how to use and edit Wikipedia

values: About the project, what you can do, where to find things

values: A list of recent changes in the wiki [ctrl-alt-r]

values: portal navigation p-interaction p-interaction-label

values: List of all English Wikipedia pages containing links to this page [ctrl-alt-j]

values: Recent changes in pages linked from this page [ctrl-alt-k]

values:
/wiki/Wikipedia:File_Upload_Wizard
Upload files [ctrl-alt-u]
u

values: A list of all special pages [ctrl-alt-q]

values: Permanent link to this revision of the page

values: Link to connected data repository item [ctrl-alt-g]

values: Printable version of this page [ctrl-alt-p]

values: portal navigation p-wikibase-otherprojects p-wikibase-otherprojects-label

values:
uls-settings-trigger
Language settings
0
button
true

values: تضاعف الحمض النووي الريبوزي منقوص الأكسجين – Arabic

values: Рэплікацыя ДНК – Belarusian

values: Репликация на ДНК – Bulgarian

values: interlanguage-link interwiki-cs badge-Q17437798 badge-goodarticle

values: Αντιγραφή του DNA – Greek

values: interlanguage-link interwiki-eu badge-Q17437796 badge-featuredarticle

values: همانندسازی – Persian

values: ԴՆԹ-ի կրկնապատկում – Armenian

values: დნმ-ის რეპლიკაცია – Georgian

values: ДНҚ репликациясы – Kazakh

values: Репликация – Kyrgyz

values: Репликација на ДНК – Macedonian

values: Репликация ДНК – Russian

values: دووھێندبوونی دی ئێن ئەی – Central Kurdish

values: Репликација ДНК – Serbian

values: டி. என். ஏ இரட்டித்தல் – Tamil

values: การถ่ายแบบดีเอ็นเอ – Thai

values: Реплікація ДНК – Ukrainian

values: ڈی این اے تنسخ – Urdu

values: /static/images/wikimedia-button-1.5x.png 1.5x, /static/images/wikimedia-button-2x.png 2x

values: /static/images/poweredby_mediawiki_132x47.png 1.5x, /static/images/poweredby_mediawiki_176x62.png 2x

values: mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject page-DNA_replication rootpage-DNA_replication vector-experimental-print-styles vector-nav-directionality skin-vector action-view