document_title: The Motorcycle Diaries (film)

question_text: where do fuser and alberto meet the indigenous couple who were traveling to look for work

text: The Motorcycle Diaries (film)

text: Plot[edit]

text: "The Che of The Motorcycle Diaries is more akin to Jack Kerouac or Neal Cassady than Marx or Lenin."

text: In 1952, a semester before Ernesto "Fuser" Guevara is due to complete his medical degree, he and his older friend Alberto Granado, a biochemist, leave Buenos Aires in order to travel across South America. While there is a goal at the end of their journey - they intend to work in a leper colony in Peru - the main purpose is initially fun and adventure. They desire to see as much of Latin America as they can, more than 14,000 kilometres (8,700 mi) in just four and a half months, while Granado's purpose is also to bed as many women as will fall for his pick-up lines. Their initial method of transport is Granado's dilapidated Norton 500 motorcycle christened La Poderosa ("The Mighty One").

text: Their planned route is ambitious, bringing them north across the Andes, along the coast of Chile, through the Atacama Desert and into the Peruvian Amazon in order to reach Venezuela just in time for Granado's 30th birthday on 2 April. However, due to La Poderosa's breakdown, they are forced to travel at a much slower pace, often walking, and don't make it to Caracas until July.

text: During their expedition, Guevara and Granado encounter the poverty of the indigenous peasants, and the movie assumes a greater seriousness once the men gain a better sense of the disparity between the "haves" (to which they belong) and the obviously exploited "have-nots" (who make up the majority of those they encounter) by traveling on foot. In Chile, for instance, they encounter a penniless and persecuted couple forced onto the road because of their communist beliefs. In a fire-lit scene, Guevara and Granado ashamedly admit to the couple that they are not out looking for work as well. The duo then accompanies the couple to the Chuquicamata copper mine, where Guevara becomes angry at the treatment of the workers.

text: However, it is a visit to the ancient Incan ruins of Machu Picchu in Peru that solidifies something in Guevara. His musings are then somberly refocused to how an indigenous civilization capable of building such beauty could be destroyed by the creators of the eventual polluted urban decay of nearby Lima.[5] His reflections are interrupted by Granado, who shares with him a dream to peacefully revolutionize and transform modern South America, to which Guevara quickly retorts: "A revolution without guns? It will never work."

text: Later, in Peru, they volunteer for three weeks at the San Pablo leper colony. There, Guevara observes both literally and metaphorically the division of society, as the staff live on the north side of a river, separated from the deprived lepers living across the river to the south. To demonstrate his solidarity, and his medical belief that leprosy is not contagious, Guevara refuses to wear rubber gloves during his visit as the head nun requires, choosing instead to shake bare hands and interact normally with the surprised leper patients.

text:
"Every generation needs a journey story; every generation needs a story about what it is to be transformed by geography, what it is to be transformed by encounters with cultures and people that are alien from yourself, and you know that age group 15 to 25, that’s the perfect generation to get on a motorcycle, to hit the road, to put on your backpack and just go out."

— José Rivera, screenwriter, NPR[6]

tag: div

text: At the end of the film, after his sojourn at the leper colony, Guevara confirms his nascent egalitarian, anti-authoritarian impulses, while making a birthday toast, which is also his first political speech. In it he invokes a pan-Latin American identity that transcends both the arbitrary boundaries of nation and race. These encounters with social injustice transform the way Guevara sees the world and his purposes in it, and by implication motivates his later political activities as a Marxist revolutionary.

text: Guevara makes his symbolic "final journey" at night when, despite the danger and his asthma, he swims across the river that separates the two societies of the leper colony, to spend the night in a leper shack, instead of in the doctors' cabins. Later, as they bid each other farewell at an airport, Granado reveals that his birthday was not in fact 2 April, but rather 8 August, and that the aforementioned goal was simply a motivator: Guevara replies that he knew all along. The film closes with an appearance by the real 82-year-old Alberto Granado, along with pictures from the actual journey and a brief mention of Che Guevara's eventual 1967 CIA-assisted execution in the Bolivian jungle.

text: Cast[edit]

text:
Gael García Bernal as Che Guevara [7]
Rodrigo de la Serna as Alberto Granado
Mercedes Morán as Celia de la Serna
Jean Pierre Noher as Ernesto Guevara Lynch
Facundo Espinosa as Tomás Granado
Mía Maestro as Chichina
Lucas Oro as Roberto Guevara
Marina Glezer as Celita Guevara
Sofia Bertolotto as Ana María Guevara
Franco Solazzi as Juan Martín Guevara
Ricardo Díaz Mourelle as Uncle Jorge
Sergio Boris as Young Traveler
Daniel Cargieman as Young Traveler
Diego Giorzi as Rodolfo
Gustavo Bueno as Doctor Hugo Pesce
Alberto Granado as himself (cameo at the end of the film)
Matías Strafe as Chichina's friend

text: Development[edit]

text:
“
The film shows what we were, which was two young men - boys, really - who went looking for adventure and found the truth and tragedy of our homeland.
”

—  Alberto Granado, 2004[8]

text: To prepare for the role of the young Che Guevara, Gael García Bernal went through six months of intense preparation. This groundwork included reading "every biography" about Guevara, traveling to Cuba to speak with Guevara's family, and consulting with Guevara's then still living travel partner Alberto Granado.[9] Despite being in his eighties, Granado was also taken on as an adviser by Salles, and enthusiastically followed the film crew as they retraced his former journey.[10]

text: Moreover, García Bernal (who is Mexican) adopted an Argentine accent and spent 14 weeks reading the works of José Martí,[11] Karl Marx and Pablo Neruda (Guevara's favorite poet). García Bernal told reporters "I feel a lot of responsibility. I want to do it well because of what Che represents to the world. He is a romantic. He had a political consciousness that changed Latin America."[12] According to García Bernal, the role crystallized his "own sense of duty" because Guevara "decided to live on the side of the mistreated, to live on the side of the people who have no justice - and no voice." In surmising the similarities between his own personal transformation and Guevara's, García Bernal posits that "my generation is awakening, and we're discovering a world full of incredible injustice."[11]

text: Granado later stated that he appreciated the film's effort "to dig beneath the "mythical Che", whose defiant image appears on T-shirts and posters around the world, "to reveal the flawed, flesh-and-blood Ernesto beneath."[8]

text: Film locales[edit]

text: "We were re-enacting a journey that was done 50 years ago, and what's surprising is that the social problems of Latin America are the same. Which is heartbreaking in a way, but it also makes you feel how important it is to tell the story."

text: In a journey that lasts eight months, the partners travel over 14,000 kilometers, from Argentina through Chile, Peru, and Colombia to Venezuela. Key locations along the journey described in the film include: In Argentina: Buenos Aires, Miramar, Villa Gesell, San Martín de los Andes, Lago Frías, Patagonia and Nahuel Huapi Lake ; In Chile: Temuco, Los Angeles, Valparaiso, Atacama desert, and Chuquicamata; In Peru: Cuzco, Machu Picchu, Lima; The San Pablo Leper Colony; Plus Leticia, Colombia and Caracas, Venezuela.

text: Reviewer Nick Cowen of The Daily Telegraph described the scenery as "visually stunning" while remarking that "the cinematography of fog-cloaked mountains, lush, green forests and sun burnt deserts is breathtakingly beautiful enough to serve as a travel advert for the entire continent."[13]

text: Tourist trails[edit]

text: "The film The Motorcycle Diaries showed the world Che's journey through Latin America and we have even designed our own tour following in his footsteps, it's great to know that the countries are collaborating to ensure his memory too."

text: The Observer reported that shortly after the film's release, tour operators in the region received a surge of inquiries, with some of them even offering Che Guevara-themed trips, where travelers could "follow in the footsteps of the revolutionary icon."[16]

text: In May 2010, tourism chiefs in Argentina, Bolivia and Cuba announced that they were working together on an international tourist route that will trace the life of Che Guevara.[15] Although Guevara is already the focus of tourist sites in his home country of Argentina, where visitors can visit his birthplace and his family's mate plantation, this trail entitled the "Caminos de Che", aims to allow tourists to venture to the three countries that most influenced the young Guevara's life.[15] Diego Conca, coordinator for the Che trail in Argentina, remarked that "people all over the world ask us for more information, each month there's more interest, and now with Bolivia, we think there will be even more."[15]

text: The crew filmed in the same San Pablo Leper Colony that Guevara himself had visited. According to Bernal, 85% of the people suffering leprosy in the film were actual lepers, with some of them having lived there when Che and Granado worked at the colony.[17] In fact, when Granado returned with the film crew to the leprosarium of San Pablo, he found some of the people he had treated half a century earlier, remarking that "It was wonderful and amazing that they could still remember me."[10] Granado was also pleased that buildings constructed for the scenes shot at the leprosarium were afterwards used by the patients themselves.[18]

text: The scene which features Guevara's character swimming across to the other side of the river was filmed during three nights in which Bernal swam across the actual Amazon River.[17]

text: Soundtrack[edit]

text: The score for The Motorcycle Diaries was composed by Gustavo Santaolalla. The film's soundtrack was released on the Deutsche Grammophon label in 2004.

text: Distribution[edit]

text: The film was first presented at the Sundance Film Festival on 15 January 2004. Granado had an invitation to the Sundance premiere, but he was refused an entry Visa by the United States.[10] Later it was featured at the 2004 Cannes Film Festival on 19 May, and Granado was able to attend.[citation needed]

text: The film later screened at many other film festivals, including: the Auckland International Film Festival, New Zealand; the Copenhagen International Film Festival, Denmark; the Espoo Film Festival, Finland; the Telluride Film Festival, United States; the Toronto International Film Festival, Canada; the Vancouver International Film Festival, Canada; the Celebrating Literature in Cinema Film Festival Frankfurt, Germany; the Morelia Film Festival, Mexico; and others.[citation needed]

text:
United States: 15 January 2004 (premiere at Sundance Film Festival)
France: 7 July 2004
Argentina: 29 July 2004
United Kingdom: 27 August 2004
United States: 24 September 2004
Chile: 21 October 2004
Germany: 28 October 2004

text: Critical reception[edit]

text: "The Motorcycle Diaries may not provide any satisfactory answers as to how a 23-year-old medical student went on to become arguably the most famous revolutionary of the latter half of the 20th Century, but it has an undeniable charm in that it imbues the memories of youth with a sense of altruism and purity – which are complemented by the scenery. It's an incomplete portrait to be sure, but it's a gorgeous depiction of two best friends riding unknowingly into the history books."

text: The Motorcycle Diaries was released to very positive reviews by critics, and received a standing ovation at the 2004 Sundance Film Festival.[19] The New York Times film critic, A.O. Scott, wrote that "in Mr. Salles's hands what might have been a schematic story of political awakening becomes a lyrical exploration of the sensations and perceptions from which a political understanding of the world emerges."[20] Gregory Weinkauf of the Dallas Observer espoused that the film "delivers as both biography and road movie, and proves itself a deceptively humble epic, an illuminating part of the Che legacy."[21] Claudia Puig of USA Today postulated that "the movie achieves an impressive blend of emotional resonance and light entertainment" while describing it as "more coming-of-age story than biopic" and "a transformative adventure well worth watching."[22] Keri Petersen of The Gainesville Sun referred to the film as "a gorgeous, poetic adventure."[23]

text: Paula Nechak of the Seattle Post-Intelligencer praised director Salles by remarking that he "presents the evolutionary course of a young man who coincidentally became the dorm-room poster boy for an idealistic generation, and captures the lovely, heart-and-eye-opening ode to youthful possibility with affection and compassion."[24] While Washington Post critic Desson Thomson lent praise for the films starring actor by observing that "what Bernal and this well-wrought movie convey so well is the charisma that would soon become a part of human history, and yes, T-shirts."[25]

text: Among the film's few detractors was Roger Ebert of the Chicago Sun-Times, who described the film's positive reviews as "a matter of political correctness, I think; it is uncool to be against Che Guevara." Ebert also criticized the film's characterization: "seen simply as a film, The Motorcycle Diaries is attenuated and tedious. We understand that Ernesto and Alberto are friends, but that's about all we find out about them; they develop none of the complexities of other on-the-road couples... Nothing is startling or poetic."[26] Jessica Winter of The Village Voice also criticized the film's simplistic representation of the peasantry, describing "the young men's encounters with conscience-pricking, generically noble locals" who are occasionally assembled "to face the camera in a still life of heroic, art-directed suffering".[27] The film also received criticism for its positive representation of Guevara as a youthful idealist. Anthony Daniels, an outspoken critic of Guevara's, argued that the film helps to continue his wrongful glorification, noting "The film is thus the cinematic equivalent of the Che Guevara T-shirt; it is morally monstrous and emotionally trivial."[28] While Frans Weiser agreed, saying that the film's narrative is dominated by reductive images of Guevara as an idealistic, loveable rogue.[29]

text: For their part, the online review aggregator Metacritic gives the film a score of 75, indicating generally favorable reviews, while Rotten Tomatoes records 84% favorable reviews among 148 reviews.[30] Furthermore, British historian Alex von Tunzelmann, who reviews films at The Guardian for historical accuracy, graded the film an A- in "History", while giving the film a B in "Entertainment".[31] After comparing scenes from the film to the actual diaries, Tunzelmann posited that "The Motorcycle Diaries gets a lot right, it's an entertaining and accurate portrayal of the formative youth of a revolutionary icon."[31]

text: Awards won[edit]

text:
Cannes Film Festival: François Chalais Award, Walter Salles; Prize of the Ecumenical Jury, Walter Salles; Technical Grand Prize, Eric Gautier; 2004.[32]
Donostia-San Sebastián International Film Festival: Audience Award Walter Salles; 2004.
Academy Awards: Oscar; Best Achievement in Music Written for Motion Pictures, Original Song; Jorge Drexler; for the song "Al otro lado del río"; 2005.
Argentine Film Critics Association Awards: Best Actor, Rodrigo de la Serna; Best Music, Gustavo Santaolalla; Best Adapted Screenplay, Jose Rivera; 2005.
British Academy of Film and Television Arts: BAFTA Film Award Best Film not in the English Language, Michael Nozik, Edgard Tenenbaum, Karen Tenkhoff, Walter Salles;Anthony Asquith Award for Film Music, Gustavo Santaolalla; 2005.
Goya Awards: Goya; Best Adapted Screenplay, José Rivera; 2005.
Independent Spirit Awards: Independent Spirit Award; Best Cinematography, Eric Gautier; Best Debut Performance, Rodrigo de la Serna; 2005.

text: Related films[edit]


document_title: List of country calling codes

question_text: what is the international phone code for canada

root

text: List of country calling codes

text: Tree list[edit]

text: Country calling codes are prefix codes; hence, they can be organized as a tree. In each row of the table below, the country codes given in the left-most column share the same first digit; then subsequent columns give the second digit in ascending order.

tag: p

text:
First
digit
Following digit

0
1
2
3
4
5
6
7
8
9

1

+1: North American Numbering Plan countries and territories
CA, US, AG, AI, AS, BB, BM, BS, DM, DO, GD, GU, JM, KN, KY, LC, MP, MS, PR, SX, TC, TT, VC, VG, VI

+1 242: BS
+1 246: BB
+1 264: AI
+1 268: AG
+1 284: VG

+1 340: VI
+1 345: KY

+1 441: BM
+1 473: GD

+1 649: TC
+1 664: MS
+1 670: MP
+1 671: GU
+1 684: AS

+1 721: SX
+1 758: LC
+1 767: DM
+1 784: VC
+1 787: PR

+1 809: DO
+1 829: DO
+1 849: DO
+1 868: TT
+1 869: KN
+1 876: JM

+1 939: PR

2
+20: EG

+210: —
+211: SS
+212: MA
+213: DZ
+214: —
+215: —
+216: TN
+217: —
+218: LY
+219: —

+220: GM
+221: SN
+222: MR
+223: ML
+224: GN
+225: CI
+226: BF
+227: NE
+228: TG
+229: BJ

+230: MU
+231: LR
+232: SL
+233: GH
+234: NG
+235: TD
+236: CF
+237: CM
+238: CV
+239: ST

+240: GQ
+241: GA
+242: CG
+243: CD
+244: AO
+245: GW
+246: IO
+247: AC
+248: SC
+249: SD

+250: RW
+251: ET
+252:SO
+253: DJ
+254: KE
+255: TZ
+256: UG
+257: BI
+258: MZ
+259: —

+260: ZM
+261: MG
+262: RE, YT, TF
+263: ZW
+264: NA
+265: MW
+266: LS
+267: BW
+268: SZ
+269: KM

+27: ZA
+28: —

+290: SH, TA
+291: ER
+292: —
+293: —
+294: —
+295: —
+296: —
+297: AW
+298: FO
+299: GL

3
+30: GR
+31: NL
+32: BE
+33: FR
+34: ES

+350: GI
+351: PT
+352: LU
+353: IE
+354: IS
+355: AL
+356: MT
+357: CY
+358: FI, AX
+359: BG

+36: HU

+370: LT
+371: LV
+372: EE
+373: MD
+374: AM, QN
+375: BY
+376: AD
+377: MC
+378: SM
+379: VA

+380: UA
+381: RS
+382: ME
+383: XK
+384: —
+385: HR
+386: SI
+387: BA
+388: EU
+389: MK

+39: IT, VA

4
+40: RO
+41: CH

+420: CZ
+421: SK
+422: —
+423: LI
+424: —
+425: —
+426: —
+427: —
+428: —
+429: —

+43: AT
+44: GB/UK, GG, IM, JE
+45: DK
+46: SE
+47: NO, SJ
+48: PL
+49: DE

5

+500: FK, GS
+501: BZ
+502: GT
+503: SV
+504: HN
+505: NI
+506: CR
+507: PA
+508: PM
+509: HT

+51: PE
+52: MX
+53: CU
+54: AR
+55: BR
+56: CL
+57: CO
+58: VE

+590: GP, BL, MF
+591: BO
+592: GY
+593: EC
+594: GF
+595: PY
+596: MQ
+597: SR
+598: UY
+599: BQ, CW

6
+60: MY
+61: AU, CX, CC
+62: ID
+63: PH
+64: NZ, PN
+65: SG
+66: TH

+670: TL
+671: —
+672: NF, AQ
+673: BN
+674: NR
+675: PG
+676: TO
+677: SB
+678: VU
+679: FJ

+680: PW
+681: WF
+682: CK
+683: NU
+684: —
+685: WS
+686: KI
+687: NC
+688: TV
+689: PF

+690: TK
+691: FM
+692: MH
+693: —
+694: —
+695: —
+696: —
+697: —
+698: —
+699: —

7
+7: RU, KZ

+7 3: RU
+7 4: RU

+7 6: KZ
+7 7: KZ
+7 8: RU
+7 9: RU

8

+800: XT
+801: —
+802: —
+803: —
+804: —
+805: —
+806: —
+807: —
+808: XS
+809: —

+81: JP
+82: KR
+83: —
+84: VN

+850: KP
+851: —
+852: HK
+853: MO
+854: —
+855: KH
+856: LA
+857: —
+858: —
+859: —

+86: CN

+870: XN
+871: —
+872: —
+873: —
+874: —
+875: —
+876: —
+877: —
+878: XP
+879: —

+880: BD
+881: XG
+882: XV
+883: XV
+884: —
+885: —
+886: TW
+887: —
+888: XD
+889: —

+89: —

9
+90: TR, CT
+91: IN
+92: PK
+93: AF
+94: LK
+95: MM

+960: MV
+961: LB
+962: JO
+963: SY
+964: IQ
+965: KW
+966: SA
+967: YE
+968: OM
+969: —

+970: PS
+971: AE
+972: IL
+973: BH
+974: QA
+975: BT
+976: MN
+977: NP
+978: —
+979: XR

+98: IR

+990: —
+991: XC
+992: TJ
+993: TM
+994: AZ
+995: GE
+996: KG
+997: —
+998: UZ
+999: —

0
1
2
3
4
5
6
7
8
9

tag: table

text: Ordered by code[edit]

text: While there is a general geographic grouping to the zones, some exceptions exist for political and historical reasons. Thus, the geographical indicators below are approximations only.

text: Zone 1: North American Numbering Plan Area[edit]

text: Countries within NANP (North American Numbering Plan) administered areas are assigned area codes as if they were all within one country. The codes below in format +1 XXX represent area code XXX within the +1 NANP zone – not a separate country code.

text: The North American Numbering Plan Area includes:

text:
+1 –  Canada
+1 –  United States, including United States territories:

+1 340 –  United States Virgin Islands
+1 670 –  Northern Mariana Islands
+1 671 –  Guam
+1 684 –  American Samoa
+1 787 / 939 –  Puerto Rico

+1 Many, but not all, Caribbean nations and some Caribbean Dutch and British Overseas Territories:

+1 242 –  Bahamas
+1 246 –  Barbados
+1 264 –  Anguilla
+1 268 –  Antigua and Barbuda
+1 284 –  British Virgin Islands
+1 345 –  Cayman Islands
+1 441 –  Bermuda
+1 473 –  Grenada
+1 649 –  Turks and Caicos Islands
+1 664 –  Montserrat
+1 721 –  Sint Maarten
+1 758 –  Saint Lucia
+1 767 –  Dominica
+1 784 –  Saint Vincent and the Grenadines
+1 809 / 829 / 849 –  Dominican Republic
+1 868 –  Trinidad and Tobago
+1 869 –  Saint Kitts and Nevis
+1 876 –  Jamaica

tag: ul

text: Zone 2: mostly Africa[edit]

text: (but also Aruba, Faroe Islands, Greenland and British Indian Ocean Territory)

text:
+20 –  Egypt
+210 – unassigned
+211 –  South Sudan
+212 –  Morocco
+213 –  Algeria
+214 – unassigned
+215 – unassigned
+216 –  Tunisia
+217 – unassigned
+218 –  Libya
+219 – unassigned
+220 –  Gambia
+221 –  Senegal
+222 –  Mauritania
+223 –  Mali
+224 –  Guinea
+225 –  Ivory Coast
+226 –  Burkina Faso
+227 –  Niger
+228 –  Togo
+229 –  Benin
+230 –  Mauritius
+231 –  Liberia
+232 –  Sierra Leone
+233 –  Ghana
+234 –  Nigeria
+235 –  Chad
+236 –  Central African Republic
+237 –  Cameroon
+238 –  Cape Verde
+239 –  São Tomé and Príncipe
+240 –  Equatorial Guinea
+241 –  Gabon
+242 –  Republic of the Congo
+243 –  Democratic Republic of the Congo
+244 –  Angola
+245 –  Guinea-Bissau
+246 –  British Indian Ocean Territory
+247 –  Ascension Island
+248 –  Seychelles
+249 –  Sudan
+250 –  Rwanda
+251 –  Ethiopia
+252 –  Somalia
+253 –  Djibouti
+254 –  Kenya
+255 –  Tanzania

+255 24 –  Zanzibar, in place of never-implemented +259

+256 –  Uganda
+257 –  Burundi
+258 –  Mozambique
+259 – unassigned, was intended for Zanzibar but never implemented – see +255 Tanzania
+260 –  Zambia
+261 –  Madagascar
+262 –  Réunion

+262 269 / 639 –  Mayotte (land / mobile, formerly with +269 Comoros)

+263 –  Zimbabwe
+264 –  Namibia
+265 –  Malawi
+266 –  Lesotho
+267 –  Botswana
+268 –  Swaziland
+269 –  Comoros (Mayotte was here but moved to +262 Réunion)
+27 –  South Africa
+28x – unassigned (Reserved for country code expansion)[1]
+290 –  Saint Helena

+290 8 –  Tristan da Cunha

+291 –  Eritrea
+292 – unassigned
+293 – unassigned
+294 – unassigned
+295 – discontinued (was assigned to San Marino, see +378)
+296 – unassigned
+297 –  Aruba
+298 –  Faroe Islands
+299 –  Greenland

text: Zones 3–4: Europe[edit]

text: Originally, larger countries such as Spain, the United Kingdom or France, were assigned two-digit codes to compensate for their usually longer domestic numbers. Small countries, such as Iceland, were assigned three-digit codes. Since the 1980s, all new assignments have been three-digit regardless of countries’ populations.

text:
+30 –  Greece
+31 –  Netherlands
+32 –  Belgium
+33 –  France
+34 –  Spain
+350 –  Gibraltar
+351 –  Portugal
+352 –  Luxembourg
+353 –  Ireland
+354 –  Iceland
+355 –  Albania
+356 –  Malta
+357 –  Cyprus
+358 –  Finland

+358 18 –  Åland Islands

+359 –  Bulgaria
+36 –  Hungary

text:
+37 – Discontinued (was assigned to the  German Democratic Republic. See Germany's country code +49)
+370 –  Lithuania
+371 –  Latvia
+372 –  Estonia
+373 –  Moldova

+373 2 / 5 –  Transnistria

+374 –  Armenia

+374 47 / 97 –  Nagorno-Karabakh (landlines / mobile phones)

+375 –  Belarus
+376 –  Andorra (formerly +33 628)
+377 –  Monaco (formerly +33 93)
+378 –  San Marino (formerly +39 549)
+379 –   Vatican City assigned but uses Italian +39 06698.
+38 – Discontinued (was assigned to  Socialist Federal Republic of Yugoslavia until its break-up)
+380 –  Ukraine (formerly used by  SFR Yugoslavia)
+381 –  Serbia (formerly used by  FR Yugoslavia and  Serbia and Montenegro before Montenegro's independence)
+382 –  Montenegro
+383 –  Kosovo[a] (was assigned in 2016, and will be used from 17 December 2017)
+384 – unassigned
+385 –  Croatia
+386 –  Slovenia
+387 –  Bosnia and Herzegovina
+388 – Discontinued (was assigned to the European Telephony Numbering Space)[1][2]
+389 –  Macedonia
+39 –  Italy

+39 06 698 –   Vatican City (assigned +379 but not in use)

+40 –  Romania
+41 –   Switzerland
+42 – Discontinued (was assigned to  Czechoslovakia until its breakup)
+420 –  Czech Republic
+421 –  Slovakia
+422 – unassigned
+423 –  Liechtenstein (formerly +41 75)
+424 – unassigned
+425 – unassigned
+426 – unassigned
+427 – unassigned
+428 – unassigned
+429 – unassigned
+43 –  Austria
+44 –  United Kingdom

+44 1481 –  Guernsey
+44 1534 –  Jersey
+44 1624 –  Isle of Man

+45 –  Denmark
+46 –  Sweden
+47 –  Norway

+47 79 –  Svalbard
+47 79 –  Jan Mayen

+48 –  Poland
+49 –  Germany

text: Zone 5: Lower North America and Central America and South America[edit]

text:
+500 –  Falkland Islands

+500 x –  South Georgia and the South Sandwich Islands

+501 –  Belize
+502 –  Guatemala
+503 –  El Salvador
+504 –  Honduras
+505 –  Nicaragua
+506 –  Costa Rica
+507 –  Panama
+508 –  Saint-Pierre and Miquelon
+509 –  Haiti
+51 –  Peru
+52 –  Mexico
+53 –  Cuba
+54 –  Argentina
+55 –  Brazil
+56 –  Chile
+57 –  Colombia
+58 –  Venezuela
+590 –  Guadeloupe (including Saint Barthélemy, Saint Martin)
+591 –  Bolivia
+592 –  Guyana
+593 –  Ecuador
+594 –  French Guiana
+595 –  Paraguay
+596 –  Martinique
+597 –  Suriname
+598 –  Uruguay
+599 – Former  Netherlands Antilles, now grouped as follows:

+599 3 –  Sint Eustatius
+599 4 –  Saba
+599 5 – formerly  Sint Maarten – Now included in NANP as code +1-721 (see Zone 1, above)
+599 7 –  Bonaire
+599 8 – formerly  Aruba – See country code +297 above
+599 9 –  Curaçao

text: Zone 6: Southeast Asia and Oceania[edit]

text:
+60 –  Malaysia
+61 –  Australia (see also +672 below)

+61 8 9162 –  Cocos Islands
+61 8 9164 –  Christmas Island

+62 –  Indonesia
+63 –  Philippines
+64 –  New Zealand

+64 xx –  Pitcairn Islands

+65 –  Singapore
+66 –  Thailand
+670 –  East Timor - formerly Northern Mariana Islands which is now included in NANP as code +1-670 (See Zone 1, above)
+671 – formerly Guam – Now included in NANP as code +1-671 (See Zone 1, above)
+672 – Australian External Territories (see also +61 Australia above)

+672 1x –  Australian Antarctic Territory
+672 3 –  Norfolk Island

+673 –  Brunei
+674 –  Nauru
+675 –  Papua New Guinea
+676 –  Tonga
+677 –  Solomon Islands
+678 –  Vanuatu
+679 –  Fiji
+680 –  Palau
+681 –  Wallis and Futuna
+682 –  Cook Islands
+683 –  Niue
+684 – formerly American Samoa – Now included in NANP as code +1-684 (See Zone 1, above)
+685 –  Samoa
+686 –  Kiribati
+687 –  New Caledonia
+688 –  Tuvalu
+689 –  French Polynesia
+690 –  Tokelau
+691 –  Federated States of Micronesia
+692 –  Marshall Islands
+693 – unassigned
+694 – unassigned
+695 – unassigned
+696 – unassigned
+697 – unassigned
+698 – unassigned
+699 – unassigned

text: Zone 7: Parts of the former  Soviet Union[edit]

text:
+7 –  Russia
+7 6xx / 7xx –  Kazakhstan
+7 840 / 940 –  Abkhazia - see also +995 44

text: Zone 8: East Asia and special services[edit]

text:
+800 – International Freephone (UIFN)
+801 – unassigned
+802 – unassigned
+803 – unassigned
+804 – unassigned
+805 – unassigned
+806 – unassigned
+807 – unassigned
+808 – reserved for Shared Cost Services
+809 – unassigned
+81 –  Japan
+82 –  South Korea
+83x – unassigned (Reserved for country code expansion)[1]
+84 –  Vietnam
+850 –  North Korea
+851 – unassigned
+852 –  Hong Kong
+853 –  Macau
+854 – unassigned
+855 –  Cambodia
+856 –  Laos
+857 – unassigned, formerly ANAC satellite service
+858 – unassigned, formerly ANAC satellite service
+859 – unassigned
+86 –  China
+870 – Inmarsat "SNAC" service
+871 – unassigned (formerly used by Inmarsat, Atlantic East), discontinued in 2008
+872 – unassigned (formerly used by Inmarsat, Pacific), discontinued in 2008
+873 – unassigned (formerly used by Inmarsat, Indian), discontinued in 2008
+874 – unassigned (formerly used by Inmarsat, Atlantic West), discontinued 2008
+875 – reserved for Maritime Mobile service
+876 – reserved for Maritime Mobile service
+877 – reserved for Maritime Mobile service
+878 – Universal Personal Telecommunications services
+879 – reserved for national non-commercial purposes
+880 –  Bangladesh
+881 – Global Mobile Satellite System
+882 – International Networks
+883 – International Networks
+884 – unassigned
+885 – unassigned
+886 –  Taiwan
+887 – unassigned
+888 – Telecommunications for Disaster Relief by OCHA
+889 – unassigned
+89x – unassigned (Reserved for country code expansion)[1]

text: Zone 9: mostly Asia[edit]

text:
+90 –  Turkey

+90 392 –  Northern Cyprus

+91 –  India
+92 –  Pakistan
+93 –  Afghanistan
+94 –  Sri Lanka
+95 –  Myanmar
+960 –  Maldives
+961 –  Lebanon
+962 –  Jordan
+963 –  Syria
+964 –  Iraq
+965 –  Kuwait
+966 –  Saudi Arabia
+967 –  Yemen
+968 –  Oman
+969 – unassigned – originally South Yemen, now covered under 967 Yemen (formerly North Yemen)
+970 –  Palestine
+971 –  United Arab Emirates
+972 –  Israel
+973 –  Bahrain
+974 –  Qatar
+975 –  Bhutan
+976 –  Mongolia
+977 –    Nepal
+978 – unassigned – originally assigned to Dubai, now covered under 971
+979 – International Premium Rate Service - originally assigned to Abu Dhabi, now covered under 971
+98 –  Iran
+990 – unassigned
+991 – International Telecommunications Public Correspondence Service trial (ITPCS)
+992 –  Tajikistan
+993 –  Turkmenistan
+994 –  Azerbaijan
+995 –  Georgia

+995 34 –  South Ossetia
+995 44 –  Abkhazia[3][4] - see also +7 840, 940

+996 –  Kyrgyzstan
+997 – unassigned
+998 –  Uzbekistan
+999 – reserved for future global service

text: Alphabetical listing by country or region[edit]

text:
Country, Territory or Service
Code
Time Zone
DST

Afghanistan
7004930000000000000♠+93
UTC+04:30

Åland Islands
7004358180000000000♠+358 18
UTC+02:00
UTC+03:00

Albania
7004355000000000000♠+355
UTC+01:00
UTC+02:00

Algeria
7004213000000000000♠+213
UTC+01:00

American Samoa
7004168400000000000♠+1 684[notes 1]
UTC-11:00

Andorra
7004376000000000000♠+376
UTC+01:00
UTC+02:00

Angola
7004244000000000000♠+244
UTC+01:00

Anguilla
7004126400000000000♠+1 264[notes 1]
UTC-04:00

Antigua and Barbuda
7004126800000000000♠+1 268[notes 1]
UTC-04:00

Argentina
7004540000000000000♠+54
UTC-03:00

Armenia
7004374000000000000♠+374
UTC+04:00

Aruba
7004297000000000000♠+297
UTC-04:00

Ascension
7004247000000000000♠+247
UTC+00:00

Australia
7004610000000000000♠+61
UTC+08:00 to +10:30
UTC+08:00 to +11:00

Australian Antarctic Territory
7004672100000000000♠+672 1

Australian External Territories
7004672000000000000♠+672

Austria
7004430000000000000♠+43
UTC+01:00
UTC+02:00

Azerbaijan
7004994000000000000♠+994
UTC+04:00

Bahamas
7004124200000000000♠+1 242[notes 1]
UTC-05:00
UTC-04:00

Bahrain
7004973000000000000♠+973
UTC+03:00

Bangladesh
7004880000000000000♠+880
UTC+06:00

Barbados
7004124600000000000♠+1 246[notes 1]
UTC-04:00

Barbuda
7004126800000000000♠+1 268[notes 1]
UTC-04:00

Belarus
7004375000000000000♠+375
UTC+03:00

Belgium
7004320000000000000♠+32
UTC+01:00
UTC+02:00

Belize
7004501000000000000♠+501
UTC-06:00

Benin
7004229000000000000♠+229
UTC+01:00

Bermuda
7004144100000000000♠+1 441[notes 1]
UTC-04:00
UTC-03:00

Bhutan
7004975000000000000♠+975
UTC+06:00

Bolivia
7004591000000000000♠+591
UTC-04:00

Bonaire
7004599700000000000♠+599 7
UTC-04:00

Bosnia and Herzegovina
7004387000000000000♠+387
UTC+01:00
UTC+02:00

Botswana
7004267000000000000♠+267
UTC+02:00

Brazil
7004550000000000000♠+55
UTC-05:00 to -02:00
UTC-05:00 to -02:00

British Indian Ocean Territory
7004246000000000000♠+246
UTC+06:00

British Virgin Islands
7004128400000000000♠+1 284[notes 1]
UTC-04:00

Brunei Darussalam
7004673000000000000♠+673
UTC+08:00

Bulgaria
7004359000000000000♠+359
UTC+02:00
UTC+03:00

Burkina Faso
7004226000000000000♠+226
UTC+00:00

Burundi
7004257000000000000♠+257
UTC+02:00

Cambodia
7004855000000000000♠+855
UTC+07:00

Cameroon
7004237000000000000♠+237
UTC+01:00

Canada
7004100000000000000♠+1
UTC-08:00 to -03:30
UTC-07:00 to -02:30

Cape Verde
7004238000000000000♠+238
UTC-01:00

Caribbean Netherlands
7004599300000000000♠+599 3, +599 4, +599 7
UTC-04:00

Cayman Islands
7004134500000000000♠+1 345[notes 1]
UTC-05:00

Central African Republic
7004236000000000000♠+236
UTC+01:00

Chad
7004235000000000000♠+235
UTC+01:00

Chatham Island, New Zealand
7004640000000000000♠+64
UTC+12:00
UTC+13:00

Chile
7004560000000000000♠+56
UTC-06:00 to -04:00
UTC-05:00 to -03:00

China
7004860000000000000♠+86
UTC+08:00

Christmas Island
7004618910000000000♠+61 89164
UTC+07:00

Cocos (Keeling) Islands
7004610000000000000♠+61 89162
UTC+06:30

Colombia
7004570000000000000♠+57
UTC-05:00

Comoros
7004269000000000000♠+269
UTC+03:00

Congo
7004242000000000000♠+242
UTC+01:00

Congo, Democratic Republic of the (Zaire)
7004243000000000000♠+243
UTC+01:00 to +02:00

Cook Islands
7004682000000000000♠+682
UTC-10:00

Costa Rica
7004506000000000000♠+506
UTC-06:00

Ivory Coast
7004225000000000000♠+225
UTC+00:00

Croatia
7004385000000000000♠+385
UTC+01:00
UTC+02:00

Cuba
7004530000000000000♠+53
UTC-05:00
UTC-04:00

Curaçao
7004599900000000000♠+599 9
UTC-04:00

Cyprus
7004357000000000000♠+357
UTC+02:00
UTC+03:00

Czech Republic
7004420000000000000♠+420
UTC+01:00
UTC+02:00

Denmark
7004450000000000000♠+45
UTC+01:00
UTC+02:00

Diego Garcia
7004246000000000000♠+246
UTC+06:00

Djibouti
7004253000000000000♠+253
UTC+02:00

Dominica
7004176700000000000♠+1 767[notes 1]
UTC-04:00

Dominican Republic
7004180900000000000♠+1 809, 7004182900000000000♠+1 829, 7004184900000000000♠+1 849[notes 1]
UTC-04:00

East Timor
7004670000000000000♠+670
UTC+09:00

Easter Island
7004560000000000000♠+56
UTC-06:00
UTC-05:00

Ecuador
7004593000000000000♠+593
UTC-06:00 to -05:00

Egypt
7004200000000000000♠+20
UTC+02:00

El Salvador
7004503000000000000♠+503
UTC-06:00

Ellipso (Mobile Satellite service)
7004881200000000000♠+881 2, +881 3

EMSAT (Mobile Satellite service)
7004882130000000000♠+882 13

Equatorial Guinea
7004240000000000000♠+240
UTC+01:00

Eritrea
7004291000000000000♠+291
UTC+03:00

Estonia
7004372000000000000♠+372
UTC+02:00
UTC+03:00

Ethiopia
7004251000000000000♠+251
UTC+03:00

Falkland Islands
7004500000000000000♠+500
UTC-03:00

Faroe Islands
7004298000000000000♠+298
UTC+00:00
UTC+01:00

Fiji
7004679000000000000♠+679
UTC+12:00
UTC+13:00

Finland
7004358000000000000♠+358
UTC+02:00
UTC+03:00

France
7004330000000000000♠+33
UTC+01:00
UTC+02:00

French Antilles
7004596000000000000♠+596

French Guiana
7004594000000000000♠+594
UTC-03:00

French Polynesia
7004689000000000000♠+689
UTC-10:00 to -09:00

Gabon
7004241000000000000♠+241
UTC+01:00

Gambia
7004220000000000000♠+220
UTC+00:00

Georgia
7004995000000000000♠+995
UTC+04:00

Germany
7004490000000000000♠+49
UTC+01:00
UTC+02:00

Ghana
7004233000000000000♠+233
UTC+00:00

Gibraltar
7004350000000000000♠+350
UTC+01:00
UTC+02:00

Global Mobile Satellite System (GMSS)
7004881000000000000♠+881

Globalstar (Mobile Satellite Service)
7004881800000000000♠+881 8, +881 9

Greece
7004300000000000000♠+30
UTC+02:00
UTC+03:00

Greenland
7004299000000000000♠+299
UTC-04:00 to +00:00

Grenada
7004147300000000000♠+1 473[notes 1]
UTC-04:00

Guadeloupe
7004590000000000000♠+590
UTC-04:00

Guam
7004167100000000000♠+1 671[notes 1]
UTC+10:00

Guatemala
7004502000000000000♠+502
UTC-06:00

Guernsey
7004441480000000000♠+44 1481, +44 7781, +44 7839, +44 7911
UTC+01:00

Guinea
7004224000000000000♠+224
UTC+00:00

Guinea-Bissau
7004245000000000000♠+245
UTC+00:00

Guyana
7004592000000000000♠+592
UTC-04:00

Haiti
7004509000000000000♠+509
UTC-05:00
UTC-04:00

Honduras
7004504000000000000♠+504
UTC-06:00

Hong Kong
7004852000000000000♠+852
UTC+08:00

Hungary
7004360000000000000♠+36
UTC+01:00
UTC+02:00

Iceland
7004354000000000000♠+354
UTC+00:00

ICO Global (Mobile Satellite Service)
7004881000000000000♠+881 0, +881 1

India
7004910000000000000♠+91
UTC+05:30

Indonesia
7004620000000000000♠+62
UTC+07:00 to +09:00

Inmarsat SNAC
7004870000000000000♠+870

International Freephone Service (UIFN)
7004800000000000000♠+800

International Networks
7004882000000000000♠+882, +883

International Premium Rate Service
7004979000000000000♠+979

International Shared Cost Service (ISCS)
7004808000000000000♠+808

Iran
7004980000000000000♠+98
UTC+03:30
UTC+04:30

Iraq
7004964000000000000♠+964
UTC+03:00

Ireland
7004353000000000000♠+353
UTC+00:00
UTC+01:00

Iridium (Mobile Satellite service)
7004881600000000000♠+881 6, +881 7

Isle of Man
7004441620000000000♠+44 1624, +44 7524, +44 7624, +44 7924
UTC+00:00
UTC+01:00

Israel
7004972000000000000♠+972
UTC+02:00
UTC+03:00

Italy
7004390000000000000♠+39
UTC+01:00
UTC+02:00

Jamaica
7004187600000000000♠+1 876[notes 1]
UTC-05:00

Jan Mayen
7004477900000000000♠+47 79

Japan
7004810000000000000♠+81
UTC+09:00

Jersey
7004441530000000000♠+44 1534
UTC+00:00
UTC+01:00

Jordan
7004962000000000000♠+962
UTC+02:00

Kazakhstan
7004760000000000000♠+7 6, +7 7[notes 2]
UTC+05:00 to +06:00

Kenya
7004254000000000000♠+254
UTC+03:00

Kiribati
7004686000000000000♠+686
UTC+12:00 to +14:00

Korea, North
7004850000000000000♠+850
UTC+08:30

Korea, South
7004820000000000000♠+82
UTC+09:00

Kosovo (Dardania)
7004383000000000000♠(+383 not yet in use), +377 44, +377 45, +386 43, +386 49, +381 28, +381 29, +381 38, +381 39
UTC+01:00
UTC+02:00

Kuwait
7004965000000000000♠+965
UTC+03:00

Kyrgyzstan
7004996000000000000♠+996
UTC+05:00 to +06:00

Laos
7004856000000000000♠+856
UTC+07:00

Latvia
7004371000000000000♠+371
UTC+02:00
UTC+03:00

Lebanon
7004961000000000000♠+961
UTC+02:00
UTC+03:00

Lesotho
7004266000000000000♠+266
UTC+02:00

Liberia
7004231000000000000♠+231
UTC+00:00

Libya
7004218000000000000♠+218
UTC+01:00
UTC+02:00

Liechtenstein
7004423000000000000♠+423
UTC+01:00
UTC+02:00

Lithuania
7004370000000000000♠+370
UTC+02:00
UTC+03:00

Luxembourg
7004352000000000000♠+352
UTC+01:00
UTC+02:00

Macau
7004853000000000000♠+853
UTC+08:00

Macedonia
7004389000000000000♠+389
UTC+01:00
UTC+02:00

Madagascar
7004261000000000000♠+261
UTC+03:00

Malawi
7004265000000000000♠+265
UTC+02:00

Malaysia
7004600000000000000♠+60
UTC+08:00

Maldives
7004960000000000000♠+960
UTC+05:00

Mali
7004223000000000000♠+223
UTC+00:00

Malta
7004356000000000000♠+356
UTC+01:00
UTC+02:00

Marshall Islands
7004692000000000000♠+692
UTC+12:00

Martinique
7004596000000000000♠+596
UTC-04:00

Mauritania
7004222000000000000♠+222
UTC+00:00

Mauritius
7004230000000000000♠+230
UTC+04:00

Mayotte
7004262260000000000♠+262 269, +262 639
UTC+03:00

Mexico
7004520000000000000♠+52
UTC-08:00 to -06:00
UTC-07:00 10 -05:00

Micronesia, Federated States of
7004691000000000000♠+691
UTC+10:00 to +11:00

Midway Island, USA
7004180800000000000♠+1 808[notes 1]
UTC-11:00

Moldova
7004373000000000000♠+373
UTC+02:00
UTC+03:00

Monaco
7004377000000000000♠+377
UTC+01:00
UTC+02:00

Mongolia
7004976000000000000♠+976
UTC+07:00 to +08:00
UTC+08:00 to +09:00

Montenegro
7004382000000000000♠+382
UTC+01:00
UTC+02:00

Montserrat
7004166400000000000♠+1 664[notes 1]
UTC-04:00

Morocco
7004212000000000000♠+212
UTC+00:00
UTC+01:00

Mozambique
7004258000000000000♠+258
UTC+02:00

Myanmar
7004950000000000000♠+95
UTC+06:30

Nagorno-Karabakh
7004374470000000000♠+374 47, +374 97
UTC+04:00

Namibia
7004264000000000000♠+264
UTC+01:00
UTC+02:00

Nauru
7004674000000000000♠+674
UTC+12:00

Nepal
7004977000000000000♠+977
UTC+05:45

Netherlands
7004310000000000000♠+31
CET(UTC+01:00) / AST(UTC-04:00)
CET(UTC+02:00) / AST(UTC-04:00)

Nevis
7004186900000000000♠+1 869[notes 1]
UTC-04:00

New Caledonia
7004687000000000000♠+687
UTC+11:00

New Zealand
7004640000000000000♠+64
UTC+12:00
UTC+13:00

Nicaragua
7004505000000000000♠+505
UTC-06:00

Niger
7004227000000000000♠+227
UTC+01:00

Nigeria
7004234000000000000♠+234
UTC+01:00

Niue
7004683000000000000♠+683
UTC-11:00

Norfolk Island
7004672300000000000♠+672 3
UTC+11:00

Northern Cyprus
7004167000000000000♠+90 392
UTC+02:00
UTC+03:00

Northern Ireland
7004442800000000000♠+44 28
UTC+00:00
UTC+01:00

Northern Mariana Islands
7004167000000000000♠+1 670[notes 1]
UTC+10:00

Norway
7004470000000000000♠+47
UTC+01:00
UTC+02:00

Oman
7004968000000000000♠+968
UTC+04:00

Pakistan
7004920000000000000♠+92
UTC+05:00
UTC+06:00

Palau
7004680000000000000♠+680
UTC+09:00

Palestine, State of
7004970000000000000♠+970
UTC+02:00
UTC+03:00

Panama
7004507000000000000♠+507
UTC-05:00

Papua New Guinea
7004675000000000000♠+675
UTC+10:00

Paraguay
7004595000000000000♠+595
UTC-04:00
UTC-03:00

Peru
7004510000000000000♠+51
UTC-05:00

Philippines
7004630000000000000♠+63
UTC+08:00

Pitcairn Islands
7004640000000000000♠+64
UTC-08:00

Poland
7004480000000000000♠+48
UTC+01:00
UTC+02:00

Portugal
7004351000000000000♠+351
UTC+00:00
UTC+01:00

Puerto Rico
7004178700000000000♠+1 787, 7004178700000000000♠+1 939[notes 1]
UTC-04:00

Qatar
7004974000000000000♠+974
UTC+03:00

Réunion
7004262000000000000♠+262
UTC+04:00

Romania
7004400000000000000♠+40
UTC+02:00
UTC+03:00

Russia
7004700000000000000♠+7[notes 2]
UTC+02:00 to +12:00

Rwanda
7004250000000000000♠+250
UTC+02:00

Saba
7004599400000000000♠+599 4
UTC-04:00

Saint Barthélemy
7004590000000000000♠+590
UTC-04:00

Saint Helena
7004290000000000000♠+290
UTC+00:00

Saint Kitts and Nevis
7004186900000000000♠+1 869[notes 1]
UTC-04:00

Saint Lucia
7004175800000000000♠+1 758[notes 1]
UTC-04:00

Saint Martin (France)
7004590000000000000♠+590
UTC-04:00

Saint Pierre and Miquelon
7004508000000000000♠+508
UTC-03:00
UTC-02:00

Saint Vincent and the Grenadines
7004178400000000000♠+1 784[notes 1]
UTC-04:00

Samoa
7004685000000000000♠+685
UTC+13:00
UTC+14:00

San Marino
7004378000000000000♠+378
UTC+01:00
UTC+02:00

São Tomé and Príncipe
7004239000000000000♠+239
UTC+00:00

Saudi Arabia
7004966000000000000♠+966
UTC+03:00

Senegal
7004221000000000000♠+221
UTC+00:00

Serbia
7004381000000000000♠+381
UTC+01:00
UTC+02:00

Seychelles
7004248000000000000♠+248
UTC+04:00

Sierra Leone
7004232000000000000♠+232
UTC+00:00

Singapore
7004650000000000000♠+65
UTC+08:00

Sint Eustatius
7004599300000000000♠+599 3
UTC-04:00

Sint Maarten (Netherlands)
7004172100000000000♠+1 721[notes 1]
UTC-04:00

Slovakia
7004421000000000000♠+421
UTC+01:00
UTC+02:00

Slovenia
7004386000000000000♠+386
UTC+01:00
UTC+02:00

Solomon Islands
7004677000000000000♠+677
UTC+11:00

Somalia
7004252000000000000♠+252
UTC+03:00

South Africa
7004270000000000000♠+27
UTC+02:00

South Georgia and the South Sandwich Islands
7004500000000000000♠+500
UTC-02:00

South Ossetia
7004995340000000000♠+995 34
UTC+03:00

South Sudan
7004211000000000000♠+211
UTC+03:00

Spain
7004340000000000000♠+34
UTC+01:00
UTC+02:00

Sri Lanka
7004940000000000000♠+94
UTC+05:30

Sudan
7004249000000000000♠+249
UTC+03:00

Suriname
7004597000000000000♠+597
UTC-03:00

Svalbard
7004477900000000000♠+47 79
UTC+01:00
UTC+02:00

Swaziland
7004268000000000000♠+268
UTC+02:00

Sweden
7004460000000000000♠+46
UTC+01:00
UTC+02:00

Switzerland
7004410000000000000♠+41
UTC+01:00
UTC+02:00

Syria
7004963000000000000♠+963
UTC+02:00
UTC+03:00

Taiwan
7004886000000000000♠+886
UTC+08:00

Tajikistan
7004992000000000000♠+992
UTC+05:00

Tanzania
7004255000000000000♠+255
UTC+03:00

Telecommunications for Disaster Relief by OCHA
7004255000000000000♠+888

Thailand
7004660000000000000♠+66
UTC+07:00

Thuraya (Mobile Satellite service)
7004882160000000000♠+882 16

Togo
7004228000000000000♠+228
UTC+00:00

Tokelau
7004690000000000000♠+690
UTC+13:00

Tonga
7004676000000000000♠+676
UTC+13:00

Transnistria
7004373200000000000♠+373 2, +373 5
UTC+02:00
UTC+03:00

Trinidad and Tobago
7004186800000000000♠+1 868[notes 1]
UTC-04:00

Tristan da Cunha
7004290800000000000♠+290 8
UTC+00:00

Tunisia
7004216000000000000♠+216
UTC+01:00

Turkey
7004900000000000000♠+90
UTC+03:00

Turkmenistan
7004993000000000000♠+993
UTC+05:00

Turks and Caicos Islands
7004164900000000000♠+1 649[notes 1]
UTC-05:00
UTC-04:00

Tuvalu
7004688000000000000♠+688
UTC+12:00

Uganda
7004256000000000000♠+256
UTC+03:00

Ukraine
7004380000000000000♠+380
UTC+02:00
UTC+03:00

United Arab Emirates
7004971000000000000♠+971
UTC+04:00

United Kingdom
7004440000000000000♠+44
UTC+00:00
UTC+01:00

United States
7004100000000000000♠+1
UTC-10:00 to -05:00
UTC-10:00 to -04:00

Universal Personal Telecommunications (UPT)
7004878000000000000♠+878

Uruguay
7004598000000000000♠+598
UTC-03:00
No DST since June 2015

US Virgin Islands
7004134000000000000♠+1 340[notes 1]
UTC-04:00

Uzbekistan
7004998000000000000♠+998
UTC+05:00

Vanuatu
7004678000000000000♠+678
UTC+11:00

Venezuela
7004580000000000000♠+58
UTC-04:00

Vatican City State (Holy See)
7004390660000000000♠+39 06 698, assigned +379
UTC+01:00
UTC+02:00

Vietnam
7004840000000000000♠+84
UTC+07:00

Wake Island, USA
7004180800000000000♠+1 808[notes 1]
UTC+12:00

Wallis and Futuna
7004681000000000000♠+681
UTC+12:00

Yemen
7004967000000000000♠+967
UTC+03:00

Zambia
7004260000000000000♠+260
UTC+02:00

Zanzibar
7004255240000000000♠+255 24
UTC+03:00

Zimbabwe
7004263000000000000♠+263
UTC+02:00anil

text: Locations with no country code[edit]

text: In Antarctica, dialing is dependent on the parent country of each base:

text:
Base
Calling Code
Country
Note

Aboa
+358
 Finland

Almirante Brown Antarctic Base
+54
 Argentina

Amundsen–Scott South Pole Station
+1
 United States

Artigas Base
+598
 Uruguay

Asuka Station
+81
 Japan

Base Presidente Eduardo Frei Montalva and Villa Las Estrellas
+56
 Chile

Belgrano II
+54
 Argentina

Bellingshausen Station
+7
 Russia

Bernardo O'Higgins Station
+56
 Chile

Byrd Station
+1
 United States

Captain Arturo Prat Base
+56
 Chile

Casey Station
+672
 Australia
can be direct dialed

Comandante Ferraz Brazilian Antarctic Base
+55
 Brazil

Concordia Station
+39
+33
 Italy
 France

Davis Station
+672
 Australia
can be direct dialed

Dome Fuji Station
+81
 Japan

Dumont d'Urville Station
+33
 France

Esperanza Base
+54
 Argentina

Gabriel de Castilla Spanish Antarctic Station
+34
 Spain

Georg-von-Neumayer-Station (Replaced by Neumayer Station)
+49
 Germany

Gonzalez Videla Station
+56
 Chile

Great Wall Station
+86
 China

Halley Research Station
+44
 United Kingdom

Henryk Arctowski Polish Antarctic Station
+48
 Poland

Jang Bogo Station
+82
 South Korea

Jinnah Antarctic Station
+92
 Pakistan

Juan Carlos I Base
+34
 Spain

Jubany
+54
 Argentina

King Sejong Station
+82
 South Korea

Kohnen-Station
+49
 Germany

Kunlun Station
+86
 China

Law-Racoviţă Station
+40
 Romania

Leningradskaya Station
+7
 Russia

Machu Picchu Research Station
+51
 Peru

Macquarie Island Station
+672
 Australia
can be direct dialed

Maitri Station
+91
 India

Marambio Base
+54
 Argentina

Mario Zucchelli Station
+39
 Italy

Mawson Station
+672
 Australia
can be direct dialed

McMurdo Station
+1
 United States
can be reached by +64 code to Scott Base (NZ)

Mendel Polar Station
+420
 Czech Republic

Mirny Station
+7
 Russia

Mizuho Station
+81
 Japan

Molodyozhnaya Station
+7
+375
 Russia
 Belarus

Neumayer Station
+49
 Germany

Novolazarevskaya Station
+7
 Russia

Orcadas Base
+54
 Argentina

Palmer Station
+1
 United States

Princess Elisabeth Base
+32
 Belgium

Professor Julio Escudero Base
+56
 Chile

Progress Station
+7
 Russia

Rothera Research Station
+44
 United Kingdom

Russkaya Station
+7
 Russia

San Martín Base
+54
 Argentina

SANAE IV (South African National Antarctic Expeditions)
+27
 South Africa

Signy Research Station
+44
 United Kingdom

St. Kliment Ohridski Base
+359
 Bulgaria

Scott Base
+64
 New Zealand
can be reached via +64 2409 and four digits on McMurdo exchange

Showa Station
+81
 Japan

Svea
+46
 Sweden

Tor Station
+47
 Norway

Troll Station
+47
 Norway

Wasa Research Station
+46
 Sweden

Vostok Station
+7
 Russia

Vernadsky Research Base
+380
 Ukraine

Zhongshan (Sun Yat-Sen) Station
+86
 China

text: Other places with no calling codes:

text:
Location
Calling Code
Country
Reasons for no Code

Kerguelen Archipelago
+262
 France
No permanent local switches

Pitcairn Islands
+64
 Pitcairn Islands
No connections to local switching; there are only satellite phones on the island, replacing a single wired party line; a single pay phone also exists


document_title: Cell nucleus

question_text: chromosomes are found in what part of the cell

text: Cell nucleus

text: History

text: The nucleus was the first organelle to be discovered. What is most likely the oldest preserved drawing dates back to the early microscopist Antonie van Leeuwenhoek (1632–1723). He observed a "lumen", the nucleus, in the red blood cells of salmon.[2] Unlike mammalian red blood cells, those of other vertebrates still contain nuclei.

text: The nucleus was also described by Franz Bauer in 1804[3] and in more detail in 1831 by Scottish botanist Robert Brown in a talk at the Linnean Society of London. Brown was studying orchids under microscope when he observed an opaque area, which he called the "areola" or "nucleus", in the cells of the flower's outer layer.[4]

text: He did not suggest a potential function. In 1838, Matthias Schleiden proposed that the nucleus plays a role in generating cells, thus he introduced the name "cytoblast" (cell builder). He believed that he had observed new cells assembling around "cytoblasts". Franz Meyen was a strong opponent of this view, having already described cells multiplying by division and believing that many cells would have no nuclei. The idea that cells can be generated de novo, by the "cytoblast" or otherwise, contradicted work by Robert Remak (1852) and Rudolf Virchow (1855) who decisively propagated the new paradigm that cells are generated solely by cells ("Omnis cellula e cellula"). The function of the nucleus remained unclear.[5]

text: Between 1877 and 1878, Oscar Hertwig published several studies on the fertilization of sea urchin eggs, showing that the nucleus of the sperm enters the oocyte and fuses with its nucleus. This was the first time it was suggested that an individual develops from a (single) nucleated cell. This was in contradiction to Ernst Haeckel's theory that the complete phylogeny of a species would be repeated during embryonic development, including generation of the first nucleated cell from a "monerula", a structureless mass of primordial mucus ("Urschleim"). Therefore, the necessity of the sperm nucleus for fertilization was discussed for quite some time. However, Hertwig confirmed his observation in other animal groups, including amphibians and molluscs. Eduard Strasburger produced the same results for plants in 1884. This paved the way to assign the nucleus an important role in heredity. In 1873, August Weismann postulated the equivalence of the maternal and paternal germ cells for heredity. The function of the nucleus as carrier of genetic information became clear only later, after mitosis was discovered and the Mendelian rules were rediscovered at the beginning of the 20th century; the chromosome theory of heredity was therefore developed.[5]

text: Structures

text: The nucleus is the largest cellular organelle in animal cells.[6] In mammalian cells, the average diameter of the nucleus is approximately 6 micrometres (µm), which occupies about 10% of the total cell volume.[7] The viscous liquid within it is called nucleoplasm (or karyolymph), and is similar in composition to the cytosol found outside the nucleus.[8] It appears as a dense, roughly spherical or irregular organelle. The composition by dry weight of the nucleus is approximately: DNA 9%, RNA 1%, Histone Protein 11%, Residual Protein 14%, Acidic Proteins 65%.

text: Nuclear envelope and pores

text:
The eukaryotic cell nucleus. Visible in this diagram are the ribosome-studded double membranes of the nuclear envelope, the DNA (complexed as chromatin), and the nucleolus. Within the cell nucleus is a viscous liquid called nucleoplasm, similar to the cytoplasm found outside the nucleus.

A cross section of a nuclear pore on the surface of the nuclear envelope (1). Other diagram labels show (2) the outer ring, (3) spokes, (4) basket, and (5) filaments.

text: The nuclear envelope, otherwise known as nuclear membrane, consists of two cellular membranes, an inner and an outer membrane, arranged parallel to one another and separated by 10 to 50 nanometres (nm). The nuclear envelope completely encloses the nucleus and separates the cell's genetic material from the surrounding cytoplasm, serving as a barrier to prevent macromolecules from diffusing freely between the nucleoplasm and the cytoplasm.[9] The outer nuclear membrane is continuous with the membrane of the rough endoplasmic reticulum (RER), and is similarly studded with ribosomes.[9] The space between the membranes is called the perinuclear space and is continuous with the RER lumen.

text: Nuclear pores, which provide aqueous channels through the envelope, are composed of multiple proteins, collectively referred to as nucleoporins. The pores are about 125 million daltons in molecular weight and consist of around 50 (in yeast) to several hundred proteins (in vertebrates).[6] The pores are 100 nm in total diameter; however, the gap through which molecules freely diffuse is only about 9 nm wide, due to the presence of regulatory systems within the center of the pore. This size selectively allows the passage of small water-soluble molecules while preventing larger molecules, such as nucleic acids and larger proteins, from inappropriately entering or exiting the nucleus. These large molecules must be actively transported into the nucleus instead. The nucleus of a typical mammalian cell will have about 3000 to 4000 pores throughout its envelope,[10] each of which contains an eightfold-symmetric ring-shaped structure at a position where the inner and outer membranes fuse.[11] Attached to the ring is a structure called the nuclear basket that extends into the nucleoplasm, and a series of filamentous extensions that reach into the cytoplasm. Both structures serve to mediate binding to nuclear transport proteins.[6]

text: Most proteins, ribosomal subunits, and some DNAs are transported through the pore complexes in a process mediated by a family of transport factors known as karyopherins. Those karyopherins that mediate movement into the nucleus are also called importins, whereas those that mediate movement out of the nucleus are called exportins. Most karyopherins interact directly with their cargo, although some use adaptor proteins.[12] Steroid hormones such as cortisol and aldosterone, as well as other small lipid-soluble molecules involved in intercellular signaling, can diffuse through the cell membrane and into the cytoplasm, where they bind nuclear receptor proteins that are trafficked into the nucleus. There they serve as transcription factors when bound to their ligand; in the absence of a ligand, many such receptors function as histone deacetylases that repress gene expression.[6]

text: Nuclear lamina

text: In animal cells, two networks of intermediate filaments provide the nucleus with mechanical support: The nuclear lamina forms an organized meshwork on the internal face of the envelope, while less organized support is provided on the cytosolic face of the envelope. Both systems provide structural support for the nuclear envelope and anchoring sites for chromosomes and nuclear pores.[7]

text: The nuclear lamina is composed mostly of lamin proteins. Like all proteins, lamins are synthesized in the cytoplasm and later transported to the nucleus interior, where they are assembled before being incorporated into the existing network of nuclear lamina.[13][14] Lamins found on the cytosolic face of the membrane, such as emerin and nesprin, bind to the cytoskeleton to provide structural support. Lamins are also found inside the nucleoplasm where they form another regular structure, known as the nucleoplasmic veil,[15] that is visible using fluorescence microscopy. The actual function of the veil is not clear, although it is excluded from the nucleolus and is present during interphase.[16] Lamin structures that make up the veil, such as LEM3, bind chromatin and disrupting their structure inhibits transcription of protein-coding genes.[17]

text: Like the components of other intermediate filaments, the lamin monomer contains an alpha-helical domain used by two monomers to coil around each other, forming a dimer structure called a coiled coil. Two of these dimer structures then join side by side, in an antiparallel arrangement, to form a tetramer called a protofilament. Eight of these protofilaments form a lateral arrangement that is twisted to form a ropelike filament. These filaments can be assembled or disassembled in a dynamic manner, meaning that changes in the length of the filament depend on the competing rates of filament addition and removal.[7]

text: Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as laminopathies. The most notable laminopathy is the family of diseases known as progeria, which causes the appearance of premature aging in its sufferers. The exact mechanism by which the associated biochemical changes give rise to the aged phenotype is not well understood.[18]

text: Chromosomes

text: The cell nucleus contains the majority of the cell's genetic material in the form of multiple linear DNA molecules organized into structures called chromosomes. Each human cell contains roughly two meters of DNA. During most of the cell cycle these are organized in a DNA-protein complex known as chromatin, and during cell division the chromatin can be seen to form the well-defined chromosomes familiar from a karyotype. A small fraction of the cell's genes are located instead in the mitochondria.

text: There are two types of chromatin. Euchromatin is the less compact DNA form, and contains genes that are frequently expressed by the cell.[19] The other type, heterochromatin, is the more compact form, and contains DNA that is infrequently transcribed. This structure is further categorized into facultative heterochromatin, consisting of genes that are organized as heterochromatin only in certain cell types or at certain stages of development, and constitutive heterochromatin that consists of chromosome structural components such as telomeres and centromeres.[20] During interphase the chromatin organizes itself into discrete individual patches,[21] called chromosome territories.[22] Active genes, which are generally found in the euchromatic region of the chromosome, tend to be located towards the chromosome's territory boundary.[23]

text: Antibodies to certain types of chromatin organization, in particular, nucleosomes, have been associated with a number of autoimmune diseases, such as systemic lupus erythematosus.[24] These are known as anti-nuclear antibodies (ANA) and have also been observed in concert with multiple sclerosis as part of general immune system dysfunction.[25] As in the case of progeria, the role played by the antibodies in inducing the symptoms of autoimmune diseases is not obvious.

text: Nucleolus

text: The nucleolus is a discrete densely stained structure found in the nucleus. It is not surrounded by a membrane, and is sometimes called a suborganelle. It forms around tandem repeats of rDNA, DNA coding for ribosomal RNA (rRNA). These regions are called nucleolar organizer regions (NOR). The main roles of the nucleolus are to synthesize rRNA and assemble ribosomes. The structural cohesion of the nucleolus depends on its activity, as ribosomal assembly in the nucleolus results in the transient association of nucleolar components, facilitating further ribosomal assembly, and hence further association. This model is supported by observations that inactivation of rDNA results in intermingling of nucleolar structures.[26]

text: In the first step of ribosome assembly, a protein called RNA polymerase I transcribes rDNA, which forms a large pre-rRNA precursor. This is cleaved into the subunits 5.8S, 18S, and 28S rRNA.[27] The transcription, post-transcriptional processing, and assembly of rRNA occurs in the nucleolus, aided by small nucleolar RNA (snoRNA) molecules, some of which are derived from spliced introns from messenger RNAs encoding genes related to ribosomal function. The assembled ribosomal subunits are the largest structures passed through the nuclear pores.[6]

text: When observed under the electron microscope, the nucleolus can be seen to consist of three distinguishable regions: the innermost fibrillar centers (FCs), surrounded by the dense fibrillar component (DFC), which in turn is bordered by the granular component (GC). Transcription of the rDNA occurs either in the FC or at the FC-DFC boundary, and, therefore, when rDNA transcription in the cell is increased, more FCs are detected. Most of the cleavage and modification of rRNAs occurs in the DFC, while the latter steps involving protein assembly onto the ribosomal subunits occur in the GC.[27]

text: Other subnuclear bodies

text:
Subnuclear structure sizes

Structure name
Structure diameter

Cajal bodies
0.2–2.0 µm
[28]

Clastosomes
0.2-0.5 µm
[29]

PIKA
5 µm
[30]

PML bodies
0.2–1.0 µm
[31]

Paraspeckles
0.2–1.0 µm
[32]

Speckles
20–25 nm
[30]

text: Besides the nucleolus, the nucleus contains a number of other non-membrane-delineated bodies. These include Cajal bodies, Gemini of coiled bodies, polymorphic interphase karyosomal association (PIKA), promyelocytic leukaemia (PML) bodies, paraspeckles, and splicing speckles. Although little is known about a number of these domains, they are significant in that they show that the nucleoplasm is not a uniform mixture, but rather contains organized functional subdomains.[31]

text: Other subnuclear structures appear as part of abnormal disease processes. For example, the presence of small intranuclear rods has been reported in some cases of nemaline myopathy. This condition typically results from mutations in actin, and the rods themselves consist of mutant actin as well as other cytoskeletal proteins.[33]

text: Cajal bodies and gems

text: A nucleus typically contains between 1 and 10 compact structures called Cajal bodies or coiled bodies (CB), whose diameter measures between 0.2 µm and 2.0 µm depending on the cell type and species.[28] When seen under an electron microscope, they resemble balls of tangled thread[30] and are dense foci of distribution for the protein coilin.[34] CBs are involved in a number of different roles relating to RNA processing, specifically small nucleolar RNA (snoRNA) and small nuclear RNA (snRNA) maturation, and histone mRNA modification.[28]

text: Similar to Cajal bodies are Gemini of Cajal bodies, or gems, whose name is derived from the Gemini constellation in reference to their close "twin" relationship with CBs. Gems are similar in size and shape to CBs, and in fact are virtually indistinguishable under the microscope.[34] Unlike CBs, gems do not contain small nuclear ribonucleoproteins (snRNPs), but do contain a protein called survival of motor neuron (SMN) whose function relates to snRNP biogenesis. Gems are believed to assist CBs in snRNP biogenesis,[35] though it has also been suggested from microscopy evidence that CBs and gems are different manifestations of the same structure.[34] Later ultrastructural studies have shown gems to be twins of Cajal bodies with the difference being in the coilin component; Cajal bodies are SMN positive and coilin positive, and gems are SMN positive and coilin negative.[36]

text: RAFA and PTF domains

text: RAFA domains, or polymorphic interphase karyosomal associations, were first described in microscopy studies in 1991. Their function remains unclear, though they were not thought to be associated with active DNA replication, transcription, or RNA processing.[37] They have been found to often associate with discrete domains defined by dense localization of the transcription factor PTF, which promotes transcription of small nuclear RNA (snRNA).[38]

text: PML bodies

text: Promyelocytic leukaemia bodies (PML bodies) are spherical bodies found scattered throughout the nucleoplasm, measuring around 0.1–1.0 µm. They are known by a number of other names, including nuclear domain 10 (ND10), Kremer bodies, and PML oncogenic domains. PML bodies are named after one of their major components, the promyelocytic leukemia protein (PML). They are often seen in the nucleus in association with Cajal bodies and cleavage bodies.[31] PML bodies belong to the nuclear matrix, an ill-defined super-structure of the nucleus proposed to anchor and regulate many nuclear functions, including DNA replication, transcription, or epigenetic silencing.[39] The PML protein is the key organizer of these domains that recruits an ever-growing number of proteins, whose only common known feature to date is their ability to be SUMOylated. Yet, pml-/- mice (which have their PML gene deleted) cannot assemble nuclear bodies, develop normally and live well, demonstrating that PML bodies are dispensable for most basic biological functions.[39]

text: Splicing speckles

text: Speckles are subnuclear structures that are enriched in pre-messenger RNA splicing factors and are located in the interchromatin regions of the nucleoplasm of mammalian cells. At the fluorescence-microscope level they appear as irregular, punctate structures, which vary in size and shape, and when examined by electron microscopy they are seen as clusters of interchromatin granules. Speckles are dynamic structures, and both their protein and RNA-protein components can cycle continuously between speckles and other nuclear locations, including active transcription sites. Studies on the composition, structure and behaviour of speckles have provided a model for understanding the functional compartmentalization of the nucleus and the organization of the gene-expression machinery[40] splicing snRNPs[41][42] and other splicing proteins necessary for pre-mRNA processing.[43] Because of a cell's changing requirements, the composition and location of these bodies changes according to mRNA transcription and regulation via phosphorylation of specific proteins.[44] The splicing speckles are also known as nuclear speckles (nuclear specks), splicing factor compartments (SF compartments), interchromatin granule clusters (IGCs), B snurposomes.[45] B snurposomes are found in the amphibian oocyte nuclei and in Drosophila melanogaster embryos. B snurposomes appear alone or attached to the Cajal bodies in the electron micrographs of the amphibian nuclei.[46] IGCs function as storage sites for the splicing factors.[47]

text: Paraspeckles

text: Discovered by Fox et al. in 2002, paraspeckles are irregularly shaped compartments in the nucleus' interchromatin space.[48] First documented in HeLa cells, where there are generally 10–30 per nucleus,[49] paraspeckles are now known to also exist in all human primary cells, transformed cell lines, and tissue sections.[50] Their name is derived from their distribution in the nucleus; the "para" is short for parallel and the "speckles" refers to the splicing speckles to which they are always in close proximity.[49]

text: Paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity. They are transcription dependent[48] and in the absence of RNA Pol II transcription, the paraspeckle disappears and all of its associated protein components (PSP1, p54nrb, PSP2, CFI(m)68, and PSF) form a crescent shaped perinucleolar cap in the nucleolus. This phenomenon is demonstrated during the cell cycle. In the cell cycle, paraspeckles are present during interphase and during all of mitosis except for telophase. During telophase, when the two daughter nuclei are formed, there is no RNA Pol II transcription so the protein components instead form a perinucleolar cap.[50]

text: Perichromatin fibrils

text: Perichromatin fibrils are visible only under electron microscope. They are located next to the transcriptionally active chromatin and are hypothesized to be the sites of active pre-mRNA processing.[47]

text: Clastosome

text: Clastosomes are small nuclear bodies (0.2-0.5 µm) described as donut-shaped due to the peripheral capsule around these bodies.[29] This name is derived from the Greek klastos, broken and soma, body.[29] Clastosomes are not typically present in normal cells, making them hard to detect. They form under high proteolysis conditions within the nucleus and degrade once there is a decrease in activity or if cells are treated with proteasome inhibitors.[29][51] The scarcity of clastosomes in cells indicates that they are not required for proteasome function.[52] Osmotic stress has also been shown to cause the formation of clastosomes. [53] These nuclear bodies contain catalytic and regulatory sub-units of the proteasome and its substrates, indicating that clastosomes are sites for degrading proteins.[52]

text: The Clastosome would be an important addition to the ‘Cell Nucleus’ Wiki page because it plays a role in cell function. With further research, we may discover that this nuclear body has a more influential job in cell function than what is already known. [29]

text: Function

text: The nucleus provides a site for genetic transcription that is segregated from the location of translation in the cytoplasm, allowing levels of gene regulation that are not available to prokaryotes. The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell cycle.

text: The nucleus is an organelle found in eukaryotic cells. Inside its fully enclosed nuclear membrane, it contains the majority of the cell's genetic material. This material is organized as DNA molecules, along with a variety of proteins, to form chromosomes.

text: Cell compartmentalization

text: The nuclear envelope allows the nucleus to control its contents, and separate them from the rest of the cytoplasm where necessary. This is important for controlling processes on either side of the nuclear membrane. In most cases where a cytoplasmic process needs to be restricted, a key participant is removed to the nucleus, where it interacts with transcription factors to downregulate the production of certain enzymes in the pathway. This regulatory mechanism occurs in the case of glycolysis, a cellular pathway for breaking down glucose to produce energy. Hexokinase is an enzyme responsible for the first the step of glycolysis, forming glucose-6-phosphate from glucose. At high concentrations of fructose-6-phosphate, a molecule made later from glucose-6-phosphate, a regulator protein removes hexokinase to the nucleus,[54] where it forms a transcriptional repressor complex with nuclear proteins to reduce the expression of genes involved in glycolysis.[55]

text: In order to control which genes are being transcribed, the cell separates some transcription factor proteins responsible for regulating gene expression from physical access to the DNA until they are activated by other signaling pathways. This prevents even low levels of inappropriate gene expression. For example, in the case of NF-κB-controlled genes, which are involved in most inflammatory responses, transcription is induced in response to a signal pathway such as that initiated by the signaling molecule TNF-α, binds to a cell membrane receptor, resulting in the recruitment of signalling proteins, and eventually activating the transcription factor NF-κB. A nuclear localisation signal on the NF-κB protein allows it to be transported through the nuclear pore and into the nucleus, where it stimulates the transcription of the target genes.[7]

text: The compartmentalization allows the cell to prevent translation of unspliced mRNA.[56] Eukaryotic mRNA contains introns that must be removed before being translated to produce functional proteins. The splicing is done inside the nucleus before the mRNA can be accessed by ribosomes for translation. Without the nucleus, ribosomes would translate newly transcribed (unprocessed) mRNA, resulting in malformed and nonfunctional proteins.

text: Gene expression

text: Gene expression first involves transcription, in which DNA is used as a template to produce RNA. In the case of genes encoding proteins, that RNA produced from this process is messenger RNA (mRNA), which then needs to be translated by ribosomes to form a protein. As ribosomes are located outside the nucleus, mRNA produced needs to be exported.[57]

text: Since the nucleus is the site of transcription, it also contains a variety of proteins that either directly mediate transcription or are involved in regulating the process. These proteins include helicases, which unwind the double-stranded DNA molecule to facilitate access to it, RNA polymerases, which bind to the DNA promoter to synthesize the growing RNA molecule, topoisomerases, which change the amount of supercoiling in DNA, helping it wind and unwind, as well as a large variety of transcription factors that regulate expression.[58]

text: Processing of pre-mRNA

text: Newly synthesized mRNA molecules are known as primary transcripts or pre-mRNA. They must undergo post-transcriptional modification in the nucleus before being exported to the cytoplasm; mRNA that appears in the cytoplasm without these modifications is degraded rather than used for protein translation. The three main modifications are 5' capping, 3' polyadenylation, and RNA splicing. While in the nucleus, pre-mRNA is associated with a variety of proteins in complexes known as heterogeneous ribonucleoprotein particles (hnRNPs). Addition of the 5' cap occurs co-transcriptionally and is the first step in post-transcriptional modification. The 3' poly-adenine tail is only added after transcription is complete.

text: RNA splicing, carried out by a complex called the spliceosome, is the process by which introns, or regions of DNA that do not code for protein, are removed from the pre-mRNA and the remaining exons connected to re-form a single continuous molecule. This process normally occurs after 5' capping and 3' polyadenylation but can begin before synthesis is complete in transcripts with many exons.[6] Many pre-mRNAs, including those encoding antibodies, can be spliced in multiple ways to produce different mature mRNAs that encode different protein sequences. This process is known as alternative splicing, and allows production of a large variety of proteins from a limited amount of DNA.

text: Dynamics and regulation

text: Nuclear transport

text: The entry and exit of large molecules from the nucleus is tightly controlled by the nuclear pore complexes. Although small molecules can enter the nucleus without regulation,[59] macromolecules such as RNA and proteins require association karyopherins called importins to enter the nucleus and exportins to exit. "Cargo" proteins that must be translocated from the cytoplasm to the nucleus contain short amino acid sequences known as nuclear localization signals, which are bound by importins, while those transported from the nucleus to the cytoplasm carry nuclear export signals bound by exportins. The ability of importins and exportins to transport their cargo is regulated by GTPases, enzymes that hydrolyze the molecule guanosine triphosphate to release energy. The key GTPase in nuclear transport is Ran, which can bind either GTP or GDP (guanosine diphosphate), depending on whether it is located in the nucleus or the cytoplasm. Whereas importins depend on RanGTP to dissociate from their cargo, exportins require RanGTP in order to bind to their cargo.[12]

text: Nuclear import depends on the importin binding its cargo in the cytoplasm and carrying it through the nuclear pore into the nucleus. Inside the nucleus, RanGTP acts to separate the cargo from the importin, allowing the importin to exit the nucleus and be reused. Nuclear export is similar, as the exportin binds the cargo inside the nucleus in a process facilitated by RanGTP, exits through the nuclear pore, and separates from its cargo in the cytoplasm.

text: Specialized export proteins exist for translocation of mature mRNA and tRNA to the cytoplasm after post-transcriptional modification is complete. This quality-control mechanism is important due to these molecules' central role in protein translation. Mis-expression of a protein due to incomplete excision of exons or mis-incorporation of amino acids could have negative consequences for the cell; thus, incompletely modified RNA that reaches the cytoplasm is degraded rather than used in translation.[6]

text: Assembly and disassembly

text: During its lifetime, a nucleus may be broken down or destroyed, either in the process of cell division or as a consequence of apoptosis (the process of programmed cell death). During these events, the structural components of the nucleus — the envelope and lamina — can be systematically degraded. In most cells, the disassembly of the nuclear envelope marks the end of the prophase of mitosis. However, this disassembly of the nucleus is not a universal feature of mitosis and does not occur in all cells. Some unicellular eukaryotes (e.g., yeasts) undergo so-called closed mitosis, in which the nuclear envelope remains intact. In closed mitosis, the daughter chromosomes migrate to opposite poles of the nucleus, which then divides in two. The cells of higher eukaryotes, however, usually undergo open mitosis, which is characterized by breakdown of the nuclear envelope. The daughter chromosomes then migrate to opposite poles of the mitotic spindle, and new nuclei reassemble around them.

text: At a certain point during the cell cycle in open mitosis, the cell divides to form two cells. In order for this process to be possible, each of the new daughter cells must have a full set of genes, a process requiring replication of the chromosomes as well as segregation of the separate sets. This occurs by the replicated chromosomes, the sister chromatids, attaching to microtubules, which in turn are attached to different centrosomes. The sister chromatids can then be pulled to separate locations in the cell. In many cells, the centrosome is located in the cytoplasm, outside the nucleus; the microtubules would be unable to attach to the chromatids in the presence of the nuclear envelope.[60] Therefore, the early stages in the cell cycle, beginning in prophase and until around prometaphase, the nuclear membrane is dismantled.[15] Likewise, during the same period, the nuclear lamina is also disassembled, a process regulated by phosphorylation of the lamins by protein kinases such as the CDC2 protein kinase.[61] Towards the end of the cell cycle, the nuclear membrane is reformed, and around the same time, the nuclear lamina are reassembled by dephosphorylating the lamins.[61]

text: However, in dinoflagellates, the nuclear envelope remains intact, the centrosomes are located in the cytoplasm, and the microtubules come in contact with chromosomes, whose centromeric regions are incorporated into the nuclear envelope (the so-called closed mitosis with extranuclear spindle). In many other protists (e.g., ciliates, sporozoans) and fungi, the centrosomes are intranuclear, and their nuclear envelope also does not disassemble during cell division.

text: Apoptosis is a controlled process in which the cell's structural components are destroyed, resulting in death of the cell. Changes associated with apoptosis directly affect the nucleus and its contents, for example, in the condensation of chromatin and the disintegration of the nuclear envelope and lamina. The destruction of the lamin networks is controlled by specialized apoptotic proteases called caspases, which cleave the lamin proteins and, thus, degrade the nucleus' structural integrity. Lamin cleavage is sometimes used as a laboratory indicator of caspase activity in assays for early apoptotic activity.[15] Cells that express mutant caspase-resistant lamins are deficient in nuclear changes related to apoptosis, suggesting that lamins play a role in initiating the events that lead to apoptotic degradation of the nucleus.[15] Inhibition of lamin assembly itself is an inducer of apoptosis.[62]

text: The nuclear envelope acts as a barrier that prevents both DNA and RNA viruses from entering the nucleus. Some viruses require access to proteins inside the nucleus in order to replicate and/or assemble. DNA viruses, such as herpesvirus replicate and assemble in the cell nucleus, and exit by budding through the inner nuclear membrane. This process is accompanied by disassembly of the lamina on the nuclear face of the inner membrane.[15]

text: Disease-related dynamics

text: Initially, it has been suspected that immunoglobulins in general and autoantibodies in particular do not enter the nucleus. Now there is a body of evidence that under pathological conditions (e.g. lupus erythematosus) IgG can enter the nucleus.[63]

text: Nuclei per cell

text: Most eukaryotic cell types usually have a single nucleus, but some have no nuclei, while others have several. This can result from normal development, as in the maturation of mammalian red blood cells, or from faulty cell division.

text: Anucleated cells

text: An anucleated cell contains no nucleus and is, therefore, incapable of dividing to produce daughter cells. The best-known anucleated cell is the mammalian red blood cell, or erythrocyte, which also lacks other organelles such as mitochondria, and serves primarily as a transport vessel to ferry oxygen from the lungs to the body's tissues. Erythrocytes mature through erythropoiesis in the bone marrow, where they lose their nuclei, organelles, and ribosomes. The nucleus is expelled during the process of differentiation from an erythroblast to a reticulocyte, which is the immediate precursor of the mature erythrocyte.[64] The presence of mutagens may induce the release of some immature "micronucleated" erythrocytes into the bloodstream.[65][66] Anucleated cells can also arise from flawed cell division in which one daughter lacks a nucleus and the other has two nuclei.

text: In flowering plants, this condition occurs in sieve tube elements.

text: Multinucleated cells

text: Multinucleated cells contain multiple nuclei. Most acantharean species of protozoa[67] and some fungi in mycorrhizae[68] have naturally multinucleated cells. Other examples include the intestinal parasites in the genus Giardia, which have two nuclei per cell.[69] In humans, skeletal muscle cells, called myocytes and syncytium, become multinucleated during development; the resulting arrangement of nuclei near the periphery of the cells allows maximal intracellular space for myofibrils.[6] Multinucleated and binucleated cells can also be abnormal in humans; for example, cells arising from the fusion of monocytes and macrophages, known as giant multinucleated cells, sometimes accompany inflammation[70] and are also implicated in tumor formation.[71]

text: A number of dinoflagellates are known to have two nuclei.[72] Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic diatom. The mitochondria and the plastids of the diatom somehow remain functional.

text: Evolution

text: As the major defining characteristic of the eukaryotic cell, the nucleus' evolutionary origin has been the subject of much speculation. Four major hypotheses have been proposed to explain the existence of the nucleus, although none have yet earned widespread support.[73]

text: The first model known as the "syntrophic model" proposes that a symbiotic relationship between the archaea and bacteria created the nucleus-containing eukaryotic cell. (Organisms of the Archaea and Bacteria domain have no cell nucleus.[74]) It is hypothesized that the symbiosis originated when ancient archaea, similar to modern methanogenic archaea, invaded and lived within bacteria similar to modern myxobacteria, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and chloroplasts, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria.[75] The archaeal origin of the nucleus is supported by observations that archaea and eukarya have similar genes for certain proteins, including histones. Observations that myxobacteria are motile, can form multicellular complexes, and possess kinases and G proteins similar to eukarya, support a bacterial origin for the eukaryotic cell.[76]

text: A second model proposes that proto-eukaryotic cells evolved from bacteria without an endosymbiotic stage. This model is based on the existence of modern planctomycetes bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures.[77] A similar proposal states that a eukaryote-like cell, the chronocyte, evolved first and phagocytosed archaea and bacteria to generate the nucleus and the eukaryotic cell.[78]

text: The most controversial model, known as viral eukaryogenesis, posits that the membrane-bound nucleus, along with other eukaryotic features, originated from the infection of a prokaryote by a virus. The suggestion is based on similarities between eukaryotes and viruses such as linear DNA strands, mRNA capping, and tight binding to proteins (analogizing histones to viral envelopes). One version of the proposal suggests that the nucleus evolved in concert with phagocytosis to form an early cellular "predator".[79] Another variant proposes that eukaryotes originated from early archaea infected by poxviruses, on the basis of observed similarity between the DNA polymerases in modern poxviruses and eukaryotes.[80][81] It has been suggested that the unresolved question of the evolution of sex could be related to the viral eukaryogenesis hypothesis.[82]

text: A more recent proposal, the exomembrane hypothesis, suggests that the nucleus instead originated from a single ancestral cell that evolved a second exterior cell membrane; the interior membrane enclosing the original cell then became the nuclear membrane and evolved increasingly elaborate pore structures for passage of internally synthesized cellular components such as ribosomal subunits.[83]