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Treaty of Tordesillas
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Treaty of Tordesillas
Front page of the Portuguese-owned treaty
Created
7 June 1494 in Tordesillas, Spain
Ratified
2 July 1494 in Spain
5 September 1494 in Portugal
Location
Archivo General de Indias (Spain)
Arquivo Nacional da Torre do Tombo (Portugal)
Author(s)
Pope Alexander VI (later confirmed and ratified by Pope Julius II in 1506)[1]
Signatories
Ferdinand II of Aragon
Isabella I of Castile
John, Prince of Asturias
John II of Portugal[2]
Purpose
To divide trading and colonising rights for all newly discovered lands of the world located between Portugal and Castile (later applied between the Spanish Crown and Portugal) to the exclusion of other European nations.
The Treaty of Tordesillas (Portuguese: Tratado de Tordesilhas [tɾɐˈtaðu ðɨ tuɾðɨˈziʎɐʃ],[3] Spanish: Tratado de Tordesillas [tɾaˈtaðo ðe toɾðeˈsiʎas]), signed at Tordesillas on June 7, 1494, and authenticated at Setúbal, Portugal, divided the newly discovered lands outside Europe between the Portuguese Empire and the Crown of Castile, along a meridian 370 leagues[note 1] west of the Cape Verde islands, off the west coast of Africa. This line of demarcation was about halfway between the Cape Verde islands (already Portuguese) and the islands entered by Christopher Columbus on his first voyage (claimed for Castile and León), named in the treaty as Cipangu and Antilia (Cuba and Hispaniola).
The lands to the east would belong to Portugal and the lands to the west to Castile. The treaty was signed by Spain, 2 July 1494 and by Portugal, 5 September 1494. The other side of the world was divided a few decades later by the Treaty of Zaragoza, signed on 22 April 1529, which specified the antimeridian to the line of demarcation specified in the Treaty of Tordesillas. Originals of both treaties are kept at the Archivo General de Indias in Spain and at the Arquivo Nacional da Torre do Tombo in Portugal.[8]
This treaty would be observed fairly well by Spain and Portugal, despite considerable ignorance as to the geography of the New World; however, it omitted all of the other European powers. Those countries generally ignored the treaty, particularly those that became Protestant after the Protestant Reformation.
The treaty was included by UNESCO in 2007 in its Memory of the World Programme.
Contents
[hide]
1 Signing and enforcement
2 Tordesillas meridian
3 Antimeridian: Moluccas and Treaty of Zaragoza
4 Effect on other European powers
5 The Treaty of Madrid
6 Modern claims
7 See also
8 Notes
9 References
9.1 Citations
9.2 Bibliography
10 External links
Signing and enforcement[edit]
Lines dividing the non-Christian world between Castile and Portugal: the 1494 Tordesillas meridian (purple) and the 1529 Zaragoza antimeridian (green)
The Treaty of Tordesillas was intended to solve the dispute that had been created following the return of Christopher Columbus and his crew, who had sailed for the Crown of Castile. On his way back to Spain he first reached Lisbon, in Portugal. There he asked for another meeting with King John II to show him the newly discovered lands.
After learning of the Castilian-sponsored voyage, the Portuguese King sent a threatening letter to the Catholic Monarchs stating that by the Treaty of Alcáçovas signed in 1479 and confirmed in 1481 with the papal bull Æterni regis, that granted all lands south of the Canary Islands to Portugal, all of the lands discovered by Columbus belonged, in fact, to Portugal. Also, the Portuguese King stated that he was already making arrangements for a fleet (an armada led by Francisco de Almeida) to depart shortly and take possession of the new lands.[citation needed] After reading the letter the Catholic Monarchs knew they did not have any military power in the Atlantic to match the Portuguese, so they pursued a diplomatic way out.[citation needed] On 4 May 1493 Pope Alexander VI (Rodrigo Borgia), an Aragonese from Valencia by birth, decreed in the bull Inter caetera that all lands west of a pole-to-pole line 100 leagues west of any of the islands of the Azores or the Cape Verde Islands should belong to Castile, although territory under Catholic rule as of Christmas 1492 would remain untouched.[9] The bull did not mention Portugal or its lands, so Portugal could not claim newly discovered lands even if they were east of the line. Another bull, Dudum siquidem, entitled Extension of the Apostolic Grant and Donation of the Indies and dated 25 September 1493, gave all mainlands and islands, "at one time or even still belonging to India" to Spain, even if east of the line.[10]
The Portuguese King John II was not pleased with that arrangement, feeling that it gave him far too little land—it prevented him from possessing India, his near term goal.[citation needed] By 1493 Portuguese explorers had reached the southern tip of Africa, the Cape of Good Hope. The Portuguese were unlikely to go to war over the islands encountered by Columbus, but the explicit mention of India was a major issue. As the Pope had not made changes, the Portuguese king opened direct negotiations with the Catholic Monarchs, King Ferdinand and Queen Isabella, to move the line to the west and allow him to claim newly discovered lands east of the line. In the bargain, John accepted Inter caetera as the starting point of discussion with Ferdinand and Isabella, but had the boundary line moved 270 leagues west, protecting the Portuguese route down the coast of Africa and giving the Portuguese rights to lands that now constitute the Eastern quarter of Brazil. As one scholar assessed the results, "both sides must have known that so vague a boundary could not be accurately fixed, and each thought that the other was deceived, [concluding that it was a] diplomatic triumph for Portugal, confirming to the Portuguese not only the true route to India, but most of the South Atlantic".[11]
The treaty effectively countered the bulls of Alexander VI but was subsequently sanctioned by Pope Julius II by means of the bull Ea quae pro bono pacis of 24 January 1506.[12] Even though the treaty was negotiated without consulting the Pope, a few sources call the resulting line the "Papal Line of Demarcation".[13]
Very little of the newly divided area had actually been seen by Europeans, as it was only divided via the treaty. Castile gained lands including most of the Americas, which in 1494 had little proven wealth. The easternmost part of current Brazil was granted to Portugal when in 1500 Pedro Álvares Cabral landed there while he was en route to India. Some historians contend that the Portuguese already knew of the South American bulge that makes up most of Brazil before this time, so his landing in Brazil was not an accident.[14] One scholar points to Cabral's landing on the Brazilian coast 12 degrees farther south than the expected Cape São Roque, such that "the likelihood of making such a landfall as a result of freak weather or navigational error was remote; and it is highly probable that Cabral had been instructed to investigate a coast whose existence was not merely suspected, but already known".[15]
The line was not strictly enforced—the Spanish did not resist the Portuguese expansion of Brazil across the meridian. However, the Catholic Monarchs attempted to stop the Portuguese advance in Asia, by claiming the meridian line ran around the world, dividing the whole world in half rather than just the Atlantic. Portugal pushed back, seeking another papal pronouncement that limited the line of demarcation to the Atlantic. This was given by Pope Leo X, who was friendly toward Portugal and its discoveries, in 1514 in the bull Praecelsae devotionis.[16]
For a period between 1580 and 1640, the treaty was rendered meaningless, as the Spanish King was also King of Portugal. It was superseded by the 1750 Treaty of Madrid which granted Portugal control of the lands it occupied in South America. However, the latter treaty was immediately repudiated by the Catholic Monarch. The First Treaty of San Ildefonso settled the problem, with Spain acquiring territories east of the Uruguay River and Portugal acquiring territories in the Amazon Basin.
Emerging Protestant maritime powers, particularly England and The Netherlands, and other third parties such as Roman Catholic France, did not recognize the division of the world between only two Roman Catholic nations brokered by the pope.[17]
Tordesillas meridian[edit]
Early Tordesillas lines in South America (1495–1545)
The Treaty of Tordesillas only specified the line of demarcation in leagues from the Cape Verde Islands. It did not specify the line in degrees, nor did it identify the specific island or the specific length of its league. Instead, the treaty stated that these matters were to be settled by a joint voyage which never occurred. The number of degrees can be determined via a ratio of marine leagues to degrees applied to the Earth regardless of its assumed size, or via a specific marine league applied to the true size of the Earth, called "our sphere" by historian Henry Harrisse.[18]
The earliest Aragonese opinion was provided by Jaime Ferrer in 1495 at the request of the Aragonese king and Castilian queen to those monarchs. He stated that the demarcation line was 18° west of the most central island of the Cape Verde Islands, which is Fogo according to Harrisse, having a longitude of 24°25'W of Greenwich, hence Ferrer placed the line at 42°25'W on his sphere, which was 21.1% larger than our sphere. Ferrer also stated that his league contained 32 Olympic stades, or 6.15264 km according to Harrisse, thus Ferrer's line was 2,276.5 km west of Fogo at 47°37'W on our sphere.[19]
Cantino planisphere depicting the meridian, 1502
The earliest surviving Portuguese opinion is on the Cantino planisphere of 1502. Because its demarcation line was midway between Cape Saint Roque (northeast cape of South America) and the mouth of the Amazon River (its estuary is marked Todo este mar he de agua doçe—"All of this sea is fresh water"—and its river is marked Rio grande, "great river"), Harrisse concluded that the line was at 42°30'W on our sphere. Harrisse believed the large estuary just west of the line on the Cantino map was that of the Rio Maranhão (this estuary is now the Baía de São Marcos and the river is now the Mearim), whose flow is so weak that its gulf does not contain fresh water.[20]
In 1518 another Castilian opinion was provided by Martin Fernandez de Enciso. Harrisse concluded that Enciso placed his line at 47°24'W on his sphere (7.7% smaller than ours), but at 45°38'W on our sphere using Enciso's numerical data. Enciso also described the coastal features near which the line passed in a very confused manner. Harrisse concluded from this description that Enciso's line could also be near the mouth of the Amazon between 49° and 50°W.[21]
In 1524 the Castilian pilots (ships' captains) Thomas Duran, Sebastian Cabot (son of John Cabot), and Juan Vespuccius (nephew of Amerigo Vespucci) gave their opinion to the Badajoz Junta, whose failure to resolve the dispute led to the Treaty of Saragossa. They specified that the line was 22° plus nearly 9 miles west of the center of Santo Antão (the westernmost Cape Verde island), which Harrisse concluded was 47°17'W on their sphere (3.1% smaller than ours) and 46°36'W on our sphere.[22]
In 1524 the Portuguese presented a globe to the Badajoz Junta on which the line was marked 21°30' west of Santo Antão (22°6'36" on our sphere).[23]
Antimeridian: Moluccas and Treaty of Zaragoza[edit]
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Main article: Treaty of Zaragoza
Dutch map of the Moluccas (north is at right)
Initially, the line of demarcation did not encircle the Earth. Instead, Spain and Portugal could conquer any new lands they were the first to discover, Spain to the west and Portugal to the east, even if they passed each other on the other side of the globe.[24] But Portugal's discovery of the highly valued Moluccas in 1512 caused Spain to argue in 1518 that the Treaty of Tordesillas divided the Earth into two equal hemispheres. After the surviving ships of Magellan's fleet visited the Moluccas in 1521, Spain claimed that those islands were within its western hemisphere. In 1523, the Treaty of Vitoria called for the Badajoz Junta to meet in 1524, at which the two countries tried to reach an agreement on the anti-meridian but failed.[25] They finally agreed in a treaty signed at Zaragoza that Spain would relinquish its claims to the Moluccas upon the payment of 350,000 ducats (≈100 kg) of gold by Portugal to Spain. To prevent Spain from encroaching upon Portugal's Moluccas, the anti-meridian was to be 297.5 leagues or 17° to the east of the Moluccas, passing through the islands of Las Velas and Santo Thome.[26] This distance is slightly smaller than the 300 leagues determined by Magellan as the westward distance from los Ladrones to the Philippine island of Samar, which is just west of due north of the Moluccas.[27]
The Moluccas are a group of islands west of New Guinea. However, unlike the large modern Indonesian archipelago of the Maluku Islands, to 16th-century Europeans the Moluccas were a small chain of islands, the only place on Earth where cloves grew, just west of the large north Malukan island of Halmahera (called Gilolo at the time). Cloves were so prized by Europeans for their medicinal uses that they were worth their weight in gold.[28][29] 16th- and 17th-century maps and descriptions indicate that the main islands were Ternate, Tidore, Moti, Makian and Bacan, although the last was often ignored even though it was by far the largest island.[30][31][32] The principal island was Ternate at the chain's northern end (0°47'N, only 11 kilometres (7 mi) in diameter) on whose southwest coast the Portuguese built a stone fort (Forte de São João Baptista de Ternate) during 1522–23,[33] which could only be repaired, not modified, according to the Treaty of Saragossa. This north-south chain occupies two degrees of latitude bisected by the equator at about 127°24'E, with Ternate, Tidore, Moti, and Makian north of the equator and Bacan south of it.
Although the treaty's Santo Thome island has not been identified, its "Islas de las Velas" (Islands of the Sails) appear in a 1585 Spanish history of China, on the 1594 world map of Petrus Plancius, on an anonymous map of the Moluccas in the 1598 London edition of Linschoten, and on the 1607 world map of Petro Kærio, identified as a north-south chain of islands in the northwest Pacific, which were also called the "Islas de los Ladrones" (Islands of the Thieves) during that period.[34][35][36] Their name was changed by Spain in 1667 to "Islas de las Marianas" (Mariana Islands), which include Guam at their southern end. Guam's longitude of 144°45'E is east of the Moluccas' longitude of 127°24'E by 17°21', which is remarkably close by 16th-century standards to the treaty's 17° east. This longitude passes through the eastern end of the main north Japanese island of Hokkaidō and through the eastern end of New Guinea, which is where Frédéric Durand placed the demarcation line.[37] Moriarty and Keistman placed the demarcation line at 147°E by measuring 16.4° east from the western end of New Guinea (or 17° east of 130°E).[38] Despite the treaty's clear statement that the demarcation line passes 17° east of the Moluccas, some sources place the line just east of the Moluccas.[39][40][41]
The Treaty of Saragossa did not modify or clarify the line of demarcation in the Treaty of Tordesillas, nor did it validate Spain's claim to equal hemispheres (180° each), so the two lines divided the Earth into unequal hemispheres. Portugal's portion was roughly 191° whereas Spain's portion was roughly 169°. Both portions have a large uncertainty of ±4° because of the wide variation in the opinions regarding the location of the Tordesillas line.
Portugal gained control of all lands and seas west of the Saragossa line, including all of Asia and its neighboring islands so far "discovered," leaving Spain most of the Pacific Ocean. Although the Philippines were not named in the treaty, Spain implicitly relinquished any claim to them because they were well west of the line. Nevertheless, by 1542, King Charles V decided to colonize the Philippines, judging that Portugal would not protest because the archipelago had no spices. Although a number of expeditions sent from New Spain arrived in the Philippines, they were unable to establish a settlement because the return route across the Pacific was unknown. King Philip II succeeded in 1565 when he sent Miguel Lopez de Legazpi and Andres de Urdaneta, establishing the initial Spanish trading post at Cebu and later founding Manila in 1571.
Besides Brazil and the Moluccas, Portugal eventually controlled Angola, Mozambique, Portuguese Guinea, and São Tomé and Príncipe (among other territories and bases) in Africa; several bases or territories as Muscat, Ormus and Bahrein in the Persian Gulf, Goa, Bombay and Daman and Diu (among other coastal cities) in India; Ceylon, and Malacca, bases in present-day Indonesia as Makassar, Solor and Ambon, Portuguese Timor, the entrepôt-base of Macau and the entrepôt-enclave of Dejima (Nagasaki) in the Far East.
Spain, on the other hand, would control vast western regions in the Americas, in areas ranging from the present-day United States to present-day Argentina, an empire that would extend to the Philippines, and bases in Ternate and Formosa (17th century).
Portuguese and Spanish empires (anachronous world maps)
Portuguese Empire
Spanish Empire alongside Iberian Union
Iberian Union (1581–1640)
Effect on other European powers[edit]
The treaty was historically important in dividing Latin America, as well as establishing Spain in the western Pacific until 1898. However, it quickly became obsolete in North America, and later in Asia and Africa, where it affected colonization. It was ignored by other European nations, and with the decline of Spanish and Portuguese power, the home countries were unable to hold many of their claims, much less expand them into poorly explored areas. Thus, with sufficient backing, it became possible for any European state to colonize open territories, or those weakly held by Lisbon or Madrid. With the fall of Malacca to the Dutch, the VOC (Dutch East India Company) took control of Portuguese possessions in Indonesia, claiming Western New Guinea and Western Australia, as New Holland. Eastern Australia remained in the Spanish half of the world until claimed for Britain by James Cook in 1770. The attitude towards the treaty that other governments had was expressed in a statement attributed to France's King Francis I, "Show me Adam's will!"[42]
The Treaty of Madrid[edit]
Main article: Treaty of Madrid (13 January 1750)
In January 13, 1750, King John V of Portugal and Ferdinand VI of Spain signed the Treaty of Madrid, in which both parts sought to establish the borders between Brazil and Spanish America, admitting that the Treaty of Tordesillas, as it had been envisioned in 1494 had been superseded, and was considered void. Spain was acknowledged sovereignty over the Philippines, while Portugal would get the territory of the Amazon River basin. Portugal would relinquish the colony of Sacramento, on the northern bank of the River Plata in modern-day Uruguay, while getting the territory of the Seven Missions.[43]
Modern claims[edit]
The Treaty of Tordesillas was invoked by Chile in the 20th century to defend the principle of an Antarctic sector extending along a meridian to the South Pole, as well as the assertion that the treaty made Spanish (or Portuguese) all undiscovered land south to the Pole.[44]
Indonesia took possession of Netherlands New Guinea in 1962, supporting its claim by stating the Empire of Majapahit had included western New Guinea, and that it was part of the Treaty of Tordesillas.
The Treaty of Tordesillas was also invoked by Argentina in the 20th century as part of its claim to the Falkland/Malvinas Islands.[45]
See also[edit]
Catholic Church and the Age of Discovery
History of Portugal (1415–1578)
List of treaties
Notes[edit]
Jump up ^ 370 leagues equals 2,193 km, 1,362 statute miles, or 1,184 nautical miles.
These figures use the legua náutica (nautical league) of four Roman miles totaling 5.926 km, which was used by Spain during the 15th, 16th, and 17th centuries for navigation.[4] In 1897 Henry Harrise noted that Jaime Ferrer, the expert consulted by King Ferdinand and Queen Isabella, stated that a league was four miles of six stades each.[5] Modern scholars agree that the geographic stade was the Roman or Italian stade, not any of several other Greek stades, supporting these figures.[6][7] Harrise is in the minority when he uses the stade of 192.27 m marked within the stadium at Olympia, Greece, resulting in a league (32 stades) of 6.153 km, 3.8% larger.
References[edit]
Citations[edit]
Jump up ^ [1]
Jump up ^ Emma Helen Blair, ed., The Philippine Islands, 1493–1803 (Cleveland, Ohio: 1903). Frances Gardiner Davenport, ed., European Treaties Bearing on the History of the United States and Its Dependencies to 1648 (Washington, DC: Carnegie Institute of Washington, 1917), 100.
Jump up ^ In the European Portuguese pronunciation. Brazilians might variously pronounce it as [tɾɐˈtadʊ dʑɪ toɾdeˈziʎəs] in São Paulo, [tɾəˈtadu dʑi to̞ʀde̞ˈziʎəɕ] in Rio de Janeiro and [tɾaˈtadu dʑi tɔʁdɛˈziʎəs] in Salvador, Bahia and [tɾɐˈtadu di tɔɦde̞ˈziʎəs] in Recife.
Jump up ^ Chardon, Roland (1980). "The linear league in North America". Annals of the Association of American Geographers. 70: 129–153 [pp. 142, 144, 151]. doi:10.1111/j.1467-8306.1980.tb01304.x. JSTOR 2562946.
Jump up ^ Harrisse, pp. 85–97, 176–190.
Jump up ^ Newlyn Walkup, Eratosthenes and the mystery of the stades
Jump up ^ Engels, Donald (1985). "The length of Eratosthenes' stade". American Journal of Philology. 106: 298–311. doi:10.2307/295030. JSTOR 295030.
Jump up ^ Davenport, 85, 171.
Jump up ^ Pope Alexander VI (4 May 1493). Inter Caetera. Wikisource.
Jump up ^ Pope Alexander VI (25 September 1493) (in Latin). Dudum siquidem. Wikisource.
Jump up ^ Parry, J. H. (1973). The Age of Reconnaissance: Discovery, Exploration, and Settlement, 1450–1650. London: Cardinal. p. 194. ISBN 0-297-16603-4.
Jump up ^ Davenport, ed., 107–111.
Jump up ^ Leslie Ronald Marchant, The Papal Line of Demarcation and Its Impact in the Eastern Hemisphere on the Political Division of Australia, 1479–1829 (Greenwood, Western Australia: Woodside Valley Foundation, 2008) ISBN 978-1-74126-423-4.
Jump up ^ Crow, John A. (1992). The Epic of Latin America (Fourth ed.). University of California Press. p. 136. ISBN 0-520-07723-7.
Jump up ^ Parry, Age of Reconnaissance p. 198.
Jump up ^ Parry, Age of Reconnaissance p. 202.
Jump up ^ Parry, Age of Reconnaissance p. 205.
Jump up ^ Henry Harrisse, The Diplomatic History of America: Its first chapter 1452—1493—1494 (London: Stevens, 1897). pp. 194
Jump up ^ Harrisse, pp. 91–97, 178–190.
Jump up ^ Harrisse, pp. 100–102, 190–192.
Jump up ^ Harrisse, pp. 103–108, 122, 192–200.
Jump up ^ Harrisse, pp. 138–139, 207–208.
Jump up ^ Harrisse, pp. 207–208.
Jump up ^ Edward Gaylord Bourne, "Historical Introduction", in Blair.
Jump up ^ Emma Helen Blair, The Philippine Islands, 1493–1803, part 2
Jump up ^ Emma Helen Blair, The Philippine Islands, 1493–1803, part 3
Jump up ^ Lord Stanley of Alderley, The first voyage round the world, by Magellan, London: Hakluyt, 1874, p. 71
Jump up ^ Andaya, pp. 1–3
Jump up ^ Corn, p. xxiv. "I split the nut, once more valuable than gold."
Jump up ^ Gavan Daws and Marty Fujita, Archipelago: The Islands of Indonesia, (Berkeley: University of California Press, 1999), p. 98, ISBN 0-520-21576-1 (early 1500s).
Jump up ^ "The Portuguese in the Moluccas and in the Lesser Sunda Islands by Marco Ramerini, 1600s". Colonialvoyage.com. Archived from the original on 2011-05-14. Retrieved 2010-06-07.
Jump up ^ Lord Stanley of Alderley, The first voyage round the world, by Magellan, London: Hakluyt Society, 1874, pp. 126–27.
Jump up ^ Andaya, p. 117. After the Iberian Union (1580–1640) and the effective Dutch conquest of the Moluccas (1605–1611, pp. 152–3), the fort was destroyed by the Spanish in 1666 during their retreat to the Philippines. (p. 156)
Jump up ^ Knowlton, p.341. The islands were named both las Velas and los Ladrones in a quote from Father Juan González de Mendoza in Historia de las cosas más notables, ritos y costumbres del gran Reino de la China (History of the most remarkable things, rites and customs of the great Kingdom of China, 1585).
Jump up ^ Cortesao, p.224, with detailed maps naming each island on several maps.
Jump up ^ ed. John O. E. Clark, 100 Maps (New York: Sterling, 2005) p. 115, ISBN 1-4027-2885-9.
Jump up ^ Le Réseau Asie (2006-09-15). "The cartography of the Orientals and Southern Europeans in the beginning of the western exploration of South-East Asia from the middle of the XVth century to the beginning of the XVIIth century by Frédéric Durand". Reseau-asie.com. Retrieved 2010-06-07.
Jump up ^ "Philip II Orders the Journey of the First Manila Galleon". The Journal of San Diego History (Volume 12, Number 2 ed.). April 1966. Retrieved 2010-06-07.
Jump up ^ Lines in the sea by Giampiero Francalanci and others, p.3 about 129°E or only 1.5° east of the Moluccas.
Jump up ^ Lines of Demarcation 1529 about 134°E or 6.5° east of the Moluccas.
Jump up ^ Infoblatt Das Zeitalter der großen Entdeckungsfahrten about 135°E or 7.5° east of the Moluccas.
Jump up ^ Miller, James Rodger (2000-06-01). Skyscrapers hide the heavens: a history of Indian-white relations in Canada. p. 20. ISBN 9780802081537. Retrieved 2010-06-07.
Jump up ^ José Damião Rodrigues, Pedro Aires Oliveira (2014) História da Expansão e do Império Português ed. Esfera dos Livros, p.266-267
Jump up ^ "National Interests And Claims In The Antarctic" (PDF). Retrieved 2010-06-07.
Jump up ^ Laver, Roberto (2001). The Falklands/Malvinas case. Springer. pp. 67–69. ISBN 978-90-411-1534-8.
Bibliography[edit]
Edward G. Bourne, 'The History and Determination of the Line of Demarcation by Pope Alexander VI, between the Spanish and Portuguese Fields of Discovery and Colonization', American Historical Association, Annual Report for 1891, Washington, 1892; Senate Miscellaneous Documents, Washington, Vol.5, 1891–92, pp. 103–30.
James R Akerman, The Imperial Map (Chicago: University of Chicago Press, 2009) 138.
Leonard Y. Andaya, The world of Maluku: Eastern Indonesia in the early modern period (Honolulu: University of Hawaii Press, 1993, ISBN 0-8248-1490-8.
Emma Helen Blair, ed., The Philippine Islands, 1493-1803 (vol 1 of 55) (Cleveland, Ohio: 1903-1909), containing complete English translations of both treaties and related documents.
Stephen R. Bown, 1494: How a family feud in medieval Spain divided the world in half (New York: Thomas Dunne Books, 2012) ISBN 978-0-312-61612-0.
Charles Corn, The Scents of Eden: A Narrative of the Spice Trade, (New York: Kodansha, 1998), ISBN 1-56836-202-1.
Cortesao, Armando (1939). "Antonio Pereira and his map of circa 1545". Geographical Review. 29: 205–225. doi:10.2307/209943.
Frances Gardiner Davenport, ed., European Treaties bearing on the History of the United States and its Dependencies to 1648 (Washington, DC: Carnegie Institution of Washington, 1917/1967).
Translation of the Treaty of Tordesillas by Davenport.
Henry Harrisse, The Diplomatic History of America: Its first chapter 1452—1493—1494 (London: Stevens, 1897).
Knowlton, Edgar C. (1963). "China and the Philippines in El Periquillo Sarniento". Hispanic Review. 31: 336–347. doi:10.2307/472212.
External links[edit]
Wikimedia Commons has media related to Treaty of Tordesillas (Tratado de Tordesillas).
Treaty of Tordesillas (about.com)
Treaty of Tordesillas (Portuguese) from Archivo General de Indias
Treaty of Tordesillas[permanent dead link] English translation from Blair—BROKEN LINK
Compact Between the Catholic Sovereigns and the King of Portugal Regarding the Demarcation and the Division of the Ocean Sea[permanent dead link] English translation from Blair—BROKEN LINK
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LCCN: n82219907
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V(D)J recombination
From Wikipedia, the free encyclopedia
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for the European car classification category please see D-segment
V(D)J recombination is the unique mechanism of genetic recombination that occurs only in developing lymphocytes during the early stages of T and B cell maturation. It involves somatic recombination, and results in the highly diverse repertoire of antibodies/immunoglobulins (Igs) and T cell receptors (TCRs) found on B cells and T cells, respectively. The process is a defining feature of the adaptive immune system and its development was a key event in the evolution of jawed vertebrates.
V(D)J recombination occurs in the primary lymphoid organs (bone marrow for B cells and thymus for T cells) and in a nearly random fashion rearranges variable (V), joining (J), and in some cases, diversity (D) gene segments. The process ultimately results in novel amino acid sequences in the antigen-binding regions of Igs and TCRs that allow for the recognition of antigens from nearly all pathogens including bacteria, viruses, parasites, and worms as well as "altered self cells" as seen in cancer. The recognition can also be allergic in nature (e.g., to pollen or other allergens) or may be "autoreactive" and lead to autoimmunity.
In 1987, Susumu Tonegawa was awarded the Nobel Prize in Physiology or Medicine[1] "for his discovery of the genetic principle for generation of antibody diversity".
Contents
[hide]
1 Background
2 Immunoglobulins
2.1 Heavy chain
2.2 Light chain
3 T cell receptors
4 Mechanism
4.1 Key enzymes and components
4.2 Process
5 See also
6 References
7 Further reading
Background[edit]
Human antibody molecules (and B cell receptors) are composed of heavy and light chains (each of which contains both constant (C) and variable (V) regions), which are encoded by genes on three loci:
The immunoglobulin heavy locus (IGH@) on chromosome 14, containing the gene segments for the immunoglobulin heavy chain.
The immunoglobulin kappa (κ) locus (IGK@) on chromosome 2, containing the gene segments for part of the immunoglobulin light chain.
The immunoglobulin lambda (λ) locus (IGL@) on chromosome 22, containing the gene segments for the remainder of the immunoglobulin light chain.
Each heavy chain and light chain gene contains multiple copies of three different types of gene segments for the variable regions of the antibody proteins. For example, the human immunoglobulin heavy chain region contains 2 Constant (Cμ and Cδ) gene segments and 44 Variable (V) gene segments,[2] plus 27 Diversity (D) gene segments and 6 Joining (J) gene segments.[3] The light chains also possess 2 Constant (Cμ and Cδ) gene segments and numerous V and J gene segments, but do not have D gene segments. DNA rearrangement causes one copy of each type of gene segment to go in any given lymphocyte, generating an enormous antibody repertoire; roughly 3×1011 combinations are possible, although some are removed due to self reactivity.
Most T cell receptors are composed of an alpha chain and a beta chain. The T cell receptor genes are similar to immunoglobulin genes in that they too contain multiple V, D and J gene segments in their beta chains (and V and J gene segments in their alpha chains) that are rearranged during the development of the lymphocyte to provide that cell with a unique antigen receptor. The T cell receptor in this sense is the topological equivalent to an antigen-binding fragment of the antibody, both being part of the immunoglobulin superfamily.
Failure of the cell to create a successful product that does not self-react leads to apoptosis. Autoimmunity is prevented by eliminating lymphocytes that self-react in the thymus by testing them against an array of self antigens expressed through the function of Aire. Interestingly, the immunoglobulin lambda light chain locus contains protein-coding genes that can be lost with its rearrangement.[4] This is based on a physiological mechanism and is not pathogenetic for leukemias or lymphomas.[4]
Immunoglobulins[edit]
Simplistic overview of V(D)J recombination of immunoglobulin heavy chains
Heavy chain[edit]
In the developing B cell, the first recombination event to occur is between one D and one J gene segment of the heavy chain locus. Any DNA between these two gene segments is deleted. This D-J recombination is followed by the joining of one V gene segment, from a region upstream of the newly formed DJ complex, forming a rearranged VDJ gene segment. All other gene segments between V and D segments are now deleted from the cell’s genome. Primary transcript (unspliced RNA) is generated containing the VDJ region of the heavy chain and both the constant mu and delta chains (Cμ and Cδ). (i.e. the primary transcript contains the segments: V-D-J-Cμ-Cδ). The primary RNA is processed to add a polyadenylated (poly-A) tail after the Cμ chain and to remove sequence between the VDJ segment and this constant gene segment. Translation of this mRNA leads to the production of the Ig M heavy chain protein.
Light chain[edit]
The kappa (κ) and lambda (λ) chains of the immunoglobulin light chain loci rearrange in a very similar way, except that the light chains lack a D segment. In other words, the first step of recombination for the light chains involves the joining of the V and J chains to give a VJ complex before the addition of the constant chain gene during primary transcription. Translation of the spliced mRNA for either the kappa or lambda chains results in formation of the Ig κ or Ig λ light chain protein. Interestingly, the immunoglobulin lambda light chain locus contains protein-coding genes that can be lost with its rearrangement.[4] This is based on a physiological mechanism and is not pathogenetic for leukemias or lymphomas.[4] However, the rearrangement of several lambda variable subgenes can activate expression of an overlapping miRNA gene, which has consequences for gene expression regulation.[5]
Assembly of the Ig μ heavy chain and one of the light chains results in the formation of membrane bound form of the immunoglobulin IgM that is expressed on the surface of the immature B cell.
T cell receptors[edit]
During thymocyte development, the T cell receptor (TCR) chains undergo essentially the same sequence of ordered recombination events as that described for immunoglobulins. D-to-J recombination occurs first in the β chain of the TCR. This process can involve either the joining of the Dβ1 gene segment to one of six Jβ1 segments or the joining of the Dβ2 gene segment to one of six Jβ2 segments.[6] DJ recombination is followed (as above) with Vβ-to-DβJβ rearrangements. All gene segments between the Vβ-Dβ-Jβ gene segments in the newly formed complex are deleted and the primary transcript is synthesized that incorporates the constant domain gene (Vβ-Dβ-Jβ-Cβ). mRNA transcription splices out any intervening sequence and allows translation of the full length protein for the TCR Cβ chain.
The rearrangement of the alpha (α) chain of the TCR follows β chain rearrangement, and resembles V-to-J rearrangement described for Ig light chains (see above). The assembly of the β- and α- chains results in formation of the αβ-TCR that is expressed on a majority of T cells.
Mechanism[edit]
Key enzymes and components[edit]
The process of V(D)J recombination is mediated by VDJ recombinase, which is a diverse collection of enzymes. The key enzymes involved are recombination activating genes 1 and 2 (RAG), terminal deoxynucleotidyl transferase (TdT), and Artemis nuclease, a member of the ubiquitous non-homologous end joining (NHEJ) pathway for DNA repair.[7] Several other enzymes are known to be involved in the process and include DNA-dependent protein kinase (DNA-PK), X-ray repair cross-complementing protein 4 (XRCC4), DNA ligase IV, non-homologous end-joining factor 1 (NHEJ1; also known as Cernunnos or XRCC4-like factor [XLF]), the recently discovered Paralog of XRCC4 and XLF (PAXX), and DNA polymerases λ and μ.[8] Some enzymes involved are specific to lymphocytes (e.g., RAG, TdT), while others are found in other cell types and even ubiquitously (e.g., NHEJ components).
To maintain the specificity of recombination, V(D)J recombinase recognizes and binds to Recombination Signal Sequences (RSSs) flanking the variable (V), diversity (D), and joining (J) genes segments. RSSs are composed of three elements: a heptamer of seven conserved nucleotides, a spacer region of 12 or 23 basepairs in length, and a nonamer of nine conserved nucleotides. While the majority of RSSs vary in sequence, the consensus heptamer and nonamer sequences are CACAGTG and ACAAAAACC, respectively; and although the sequence of the spacer region is poorly conserved, the length is highly conserved.[9][10] The length of the spacer region corresponds to approximately one (12 basepairs) or two turns (23 basepairs) of the DNA helix. Following what is known as the 12/23 Rule, gene segments to be recombined are usually adjacent to RSSs of different spacer lengths (i.e., one has a "12RSS" and one has a "23RSS").[11] This is an important feature in the regulation of V(D)J recombination.[12]
Process[edit]
V(D)J recombination begins when V(D)J recombinase (through the activity of RAG1) binds an RSS flanking a coding gene segment (V, D, or J) and creates a single-strand nick in the DNA between the first base of the RSS (just before the heptamer) and the coding segment. This is essentially energetically neutral (no need for ATP hydrolysis) and results in the formation of a free 3' hydroxyl group and a 5' phosphate group on the same strand. The reactive hydroxyl group is positioned by the recombinase to attack the phosphodiester bond of opposite strand, forming two DNA ends: a hairpin (stem-loop) on the coding segment and a blunt end on the signal segment.[13] The current model is that DNA nicking and hairpin formation occurs on both strands simultaneously (or nearly so) in a complex known as a recombination center.[14][15][16][17]
The blunt signal ends are flushly ligated together to form a circular piece of DNA containing all of the intervening sequences between the coding segments known as a signal joint (although circular in nature, this is not to be confused with a plasmid). While originally thought to be lost during successive cell divisions, there is evidence that signal joints may re-enter the genome and lead to pathologies by activating oncogenes or interrupting tumor suppressor gene function(s)[Ref].
The coding ends are processed further prior to their ligation by several events that ultimately lead to junctional diversity.[18] Processing begins when DNA-PK binds to each broken DNA end and recruits several other proteins including Artemis, XRCC4, DNA ligase IV, Cernunnos, and several DNA polymerases.[19] DNA-PK forms a complex that leads to its autophosphorylation, resulting in activation of Artemis. The coding end hairpins are opened by the activity of Artemis.[20] If they are opened at the center, a blunt DNA end will result; however in many cases, the opening is "off-center" and results in extra bases remaining on one strand (an overhang). These are known as palindromic (P) nucleotides due to the palindromic nature of the sequence produced when DNA repair enzymes resolve the overhang.[21] The process of hairpin opening by Artemis is a crucial step of V(D)J recombination and is defective in the severe combined immunodeficiency (scid) mouse model.
Next, XRCC4, Cernunnos, and DNA-PK align the DNA ends and recruit terminal deoxynucleotidyl transferase (TdT), a template-independent DNA polymerase that adds non-templated (N) nucleotides to the coding end. The addition is mostly random, but TdT does exhibit a preference for G/C nucleotides.[22] As with all known DNA polymerases, the TdT adds nucleotides to one strand in a 5' to 3' direction.[23]
Lastly, exonucleases can remove bases from the coding ends (including any P or N nucleotides that may have formed). DNA polymerases λ and μ then insert additional nucleotides as needed to make the two ends compatible for joining. This is a stochastic process, therefore any combination of the addition of P and N nucleotides and exonucleolytic removal can occur (or none at all). Finally, the processed coding ends are ligated together by DNA ligase IV.[24]
All of these processing events result in an antigen-binding region that is highly variable, even when the same gene segments are recombined. V(D)J recombination allows for the generation of immunoglobulins and T cell receptors to antigens that neither the organism nor its ancestor(s) need to have previously encountered, allowing for an adaptive immune response to novel pathogens that develop or to those that frequently change (e.g., seasonal influenza). However, a major caveat to this process is that the DNA sequence must remain in-frame in order to maintain the correct amino acid sequence in the final protein product. If the resulting sequence is out-of-frame, the development of the cell will be arrested, and the cell will not survive to maturity. V(D)J recombination is therefore a very costly process that must be (and is) strictly regulated and controlled.
See also[edit]
Antibody
B cell
B cell receptor
T cell
T cell receptor
Recombination-activating gene
NKT cell
References[edit]
Jump up ^ "The Nobel Prize in Physiology or Medicine 1987". nobelprize.org. Retrieved 26 December 2014.
Jump up ^ Matsuda, F; Ishii, K; Bourvagnet, P; Kuma, K; Hayashida, H; Miyata, T; Honjo, T (1998). "The complete nucleotide sequence of the human immunoglobulin heavy chain variable region locus". The Journal of Experimental Medicine. 188 (11): 2151–62. doi:10.1084/jem.188.11.2151. PMC 2212390  . PMID 9841928.
Jump up ^ Li A, Rue M, Zhou J, et al. (June 2004). "Utilization of Ig heavy chain variable, diversity, and joining gene segments in children with B-lineage acute lymphoblastic leukemia: implications for the mechanisms of VDJ recombination and for pathogenesis". Blood. 103 (12): 4602–9. doi:10.1182/blood-2003-11-3857. PMID 15010366.
^ Jump up to: a b c d Mraz, M.; Stano Kozubik, K.; Plevova, K.; Musilova, K.; Tichy, B.; Borsky, M.; Kuglik, P.; Doubek, M.; Brychtova, Y.; Mayer, J.; Pospisilova, S. (2013). "The origin of deletion 22q11 in chronic lymphocytic leukemia is related to the rearrangement of immunoglobulin lambda light chain locus". Leukemia Research. 37 (7): 802–808. doi:10.1016/j.leukres.2013.03.018. PMID 23608880.
Jump up ^ Mraz, M.; Dolezalova, D.; Plevova, K.; Stano Kozubik, K.; Mayerova, V.; Cerna, K.; Musilova, K.; Tichy, B.; Pavlova, S.; Borsky, M.; Verner, J.; Doubek, M.; Brychtova, Y.; Trbusek, M.; Hampl, A.; Mayer, J.; Pospisilova, S. (2012). "MicroRNA-650 expression is influenced by immunoglobulin gene rearrangement and affects the biology of chronic lymphocytic leukemia". Blood. 119 (9): 2110–2113. doi:10.1182/blood-2011-11-394874. PMID 22234685.
Jump up ^ K. Abbas, Abul (2015). Cellular and Molecular Immunology, 8e. ELSEVIER. p. 192. ISBN 978-0323222754.
Jump up ^ Ma, Yunmei; Lu, Haihui; Schwarz, Klaus; Lieber, Michael (September 2005). "Repair of Double-Strand DNA Breaks by the Human Nonhomologous DNA End Joining Pathway: the Iterative Processing Model". Cell Cycle. 4 (9): 1193–1200. doi:10.4161/cc.4.9.1977. PMID 16082219.
Jump up ^ Malu, Shruti; Malshetty, Vidyasagar; Francis, Dailia; Cortes, Patricia (2012). "Role of non-homologous end joining in V(D)J recombination". Immunologic Research. 54 (1–3): 233–246. doi:10.1007/s12026-012-8329-z. PMID 22569912.
Jump up ^ Ramsden, Dale; Baetz, Kristin; Wu, Gillian (1994). "Conservation of Sequence in Recombination Signal Sequence Spacers". Nucleic Acids Research. 22 (10): 1785–1796. doi:10.1093/nar/22.10.1785. PMC 308075  . PMID 8208601.
Jump up ^ Cowell, Lindsay; Davila, Marco; Ramsden, Dale; Kelsoe, Garnett (2004). "Computational tools for understanding sequence variability in recombination signals". Immunological Reviews. 200: 57–69. doi:10.1111/j.0105-2896.2004.00171.x. PMID 15242396.
Jump up ^ van Gent, Dik; Ramsden, Dale; Gellert, Martin (1996). "The RAG1 and RAG2 Proteins Establish the 12/23 Rule in V(D)J Recombination". Cell. 85 (1): 107–13. doi:10.1016/s0092-8674(00)81086-7. PMID 8620529.
Jump up ^ Hiom, Kevin; Gellert, Martin (1998). "Assembly of a 12/23 Paired Signal Complex: a Critical Control Point in V(D)J Recombination". Molecular Cell. 1 (7): 1011–1019. doi:10.1016/s1097-2765(00)80101-x. PMID 9651584.
Jump up ^ Schatz, David; Swanson, Patrick (2011). "V(D)J Recombination: Mechanisms of Initiation". Annual Review of Genetics. 45: 167–202. doi:10.1146/annurev-genet-110410-132552. PMID 21854230.
Jump up ^ Schatz, David; Ji, Yanhong (2011). "Recombination Centres and the Orchestration of V(D)J Recombination". Nature Reviews Immunology. 11 (4): 251–263. doi:10.1038/nri2941. PMID 21394103.
Jump up ^ Curry, John; Geier, Jamie; Schlissel, Mark (2005). "Single-Strand Recombination Signal Sequence Nicks in vivo: Evidence for a Capture Model of Synapsis". Nature Immunology. 6 (12): 1272–1279. doi:10.1038/ni1270. PMID 16286921.
Jump up ^ Agrawal, Alka; Schatz, David (1997). "RAG1 and RAG2 Form a Stable Postcleavage Synaptic Complex with DNA Containing Signal Ends in V(D)J Recombination". Cell. 89 (1): 43–53. doi:10.1016/s0092-8674(00)80181-6. PMID 9094713.
Jump up ^ Fugmann, Sebastian; Lee, AIfred; Shockett, Penny; Villey, Isabelle; Schatz, David (2000). "The RAG Proteins and V(D)J Recombination: Complexes, Ends, and Transposition". Annual Review of Immunology. 18: 495–527. doi:10.1146/annurev.immunol.18.1.495. PMID 10837067.
Jump up ^ Lewis, Susanna (1994). "The Mechanism of V(D)J Joining: Lessons from Molecular, Immunological, and Comparative Analyses". Advances in Immunology. 56: 27–150. doi:10.1016/s0065-2776(08)60450-2. PMID 8073949.
Jump up ^ Helmink, Beth; Sleckman, Barry (2012). "The response to and repair of RAG-mediated DNA double-strand breaks". Annual Review of Immunology. 30: 175–202. doi:10.1146/annurev-immunol-030409-101320. PMC 4038028  . PMID 22224778.
Jump up ^ Ma, Yunmei; Schwarz, Klaus; Lieber, Michael (2005). "The Artemis:DNA-PKcs Endonuclease Cleaves DNA Loops, Flaps, and Gaps". DNA Repair. 4 (7): 845–851. doi:10.1016/j.dnarep.2005.04.013. PMID 15936993.
Jump up ^ Lu, Haihui; Schwarz, Klaus; Lieber, Michael (2007). "Extent to Which Hairpin Opening by the Artemis:DNA-PKcs Complex can Contribute to Junctional Diversity in V(D)J Recombination". Nucleic Acids Research. 35 (20): 6917–6923. doi:10.1093/nar/gkm823. PMC 2175297  . PMID 17932067.
Jump up ^ Gauss, George; Lieber, Michael (1996). "Mechanistic Constraints on Diversity in Human V(D)J Recombination". Molecular and Cellular Biology. 16 (1): 258–269. PMC 230999  . PMID 8524303.
Jump up ^ Benedict, Cindy; Gilfillan, Susan; Thai, To-Ha; Kearney, John (2000). "Terminal Deoxynucleotidyl Transferase and Repertoire Development". Immunological Reviews. 175: 150–157. doi:10.1111/j.1600-065x.2000.imr017518.x. PMID 10933600.
Jump up ^ D.C. van Gent & M. van der Burg (2007). "Non-Homologous End-Joining, a Sticky Affair". Oncogene. 26 (56): 7731–7740. doi:10.1038/sj.onc.1210871. PMID 18066085.
Further reading[edit]
Hartwell LH, Hood L, Goldberg ML, Reynolds AE, Silver LM, Veres RC (2000). Chapter 24, Evolution at the molecular level. In: Genetics. New York: McGraw-Hill. pp. 805–807. ISBN 0-07-299587-4.
V(D)J Recombination. Series: Advances in Experimental Medicine and Biology, Vol. 650 Ferrier, Pierre (Ed.) Landes Bioscience 2009, XII, 199 p. ISBN 978-1-4419-0295-5
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