document_title: Scramble for Africa

question_text: what three events allowed for the formal colonization of africa in the late 1800s

question_tokens: what three events allowed for the formal colonization of africa in the late 1800s

text: Scramble for Africa

text: Background[edit]

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text: The Portuguese established the first firm post-Middle Ages European settlements, trade posts, permanent fortifications and ports of call along the coast of the African continent, from the beginning of the Age of Discovery during the 15th century. But Europeans showed comparatively little interest in (and less knowledge of) the interior for some two centuries thereafter.

text: European exploration of the African interior began in earnest at the end of the 18th century. By 1835, Europeans had mapped most of northwestern Africa. In the middle decades of the 19th century, famous European explorers included David Livingstone and H. M. Stanley, each of whom mapped vast areas of Southern Africa and Central Africa. Arduous expeditions in the 1850s and 1860s by Richard Burton, John Speke and James Grant located the great central lakes and the source of the Nile. By the end of the 19th century Europeans had charted the Nile from its source, traced the courses of the Niger, Congo and Zambezi Rivers, and realized the vast resources of Africa.

text: Even as late as the 1870s, European states still controlled only ten percent of the African continent, with all their territories located near the coast. The most important holdings were Angola and Mozambique, held by Portugal; the Cape Colony, held by the United Kingdom; and Algeria, held by France. By 1914, only Ethiopia and Liberia remained independent of European control.[5]

text: Technological advances facilitated European expansion overseas. Industrialisation brought about rapid advancements in transportation and communication, especially in the forms of steam navigation, railways, and telegraphs. Medical advances also played an important role, especially medicines for tropical diseases. The development of quinine, an effective treatment for malaria, made vast expanses of the tropics more accessible for Europeans.

text: Causes[edit]

text: Africa and global markets[edit]

text: Sub-Saharan Africa, one of the last regions of the world largely untouched by "informal imperialism", was also attractive to Europe's ruling elites for economic, political and social reasons. During a time when Britain's balance of trade showed a growing deficit, with shrinking and increasingly protectionist continental markets due to the Long Depression (1873–96), Africa offered Britain, Germany, France, and other countries an open market that would garner them a trade surplus: a market that bought more from the colonial power than it sold overall.[4]

text: In addition, surplus capital was often more profitably invested overseas, where cheap materials, limited competition, and abundant raw materials made a greater premium possible. Another inducement for imperialism arose from the demand for raw materials unavailable in Europe, especially copper, cotton, rubber, palm oil, cocoa, diamonds, tea, and tin, to which European consumers had grown accustomed and upon which European industry had grown dependent. Additionally, Britain wanted the southern and eastern coasts of Africa for stopover ports on the route to Asia and its empire in India.[6]

text: However, in Africa – excluding the area which became the Union of South Africa in 1910 – the amount of capital investment by Europeans was relatively small, compared to other continents. Consequently, the companies involved in tropical African commerce were relatively small, apart from Cecil Rhodes's De Beers Mining Company. Rhodes had carved out Rhodesia for himself; Léopold II of Belgium later, and with considerable brutality, exploited the Congo Free State. These events might detract from the pro-imperialist arguments of colonial lobbies such as the Alldeutscher Verband, Francesco Crispi and Jules Ferry, who argued that sheltered overseas markets in Africa would solve the problems of low prices and over-production caused by shrinking continental markets.

text: John A. Hobson argued in Imperialism that this shrinking of continental markets was a key factor of the global "New Imperialism" period.

text: William Easterly of New York University, however, disagrees with the link made between capitalism and imperialism, arguing that colonialism is used mostly to promote state-led development rather than "corporate" development. He has stated that "imperialism is not so clearly linked to capitalism and free markets... historically there has been a closer link between colonialism/imperialism and state-led approaches to development."[7]

text: Strategic rivalry[edit]

text: The rivalry between Britain, France, Germany, and the other European powers accounts for a large part of the colonisation.

text: While tropical Africa was not a large zone of investment, other overseas regions were. The vast interior between Egypt and the gold and diamond-rich southern Africa had strategic value in securing the flow of overseas trade. Britain was under political pressure to secure lucrative markets against encroaching rivals in China and its eastern colonies, most notably India, Malaya, Australia and New Zealand. Thus, it was crucial to secure the key waterway between East and West—the Suez Canal. However, the prevailing theory [8][9] which suggested that Britain annexed East Africa in the 1880-90s out of geo-strategic concerns connected to the Nile Valley and Britain's position in Egypt—and by extension the sea-route to India via the Suez Canal—was challenged by John Darwin in 1997,[10] a refutation that was further contextualized and consolidated by Jonas F. Gjersø in 2015.[11]

text: The scramble for African territory also reflected concern for the acquisition of military and naval bases, for strategic purposes and the exercise of power. The growing navies, and new ships driven by steam power, required coaling stations and ports for maintenance. Defense bases were also needed for the protection of sea routes and communication lines, particularly of expensive and vital international waterways such as the Suez Canal.[12]

text: Colonies were also seen as assets in "balance of power" negotiations, useful as items of exchange at times of international bargaining. Colonies with large native populations were also a source of military power; Britain and France used large numbers of British Indian and North African soldiers, respectively, in many of their colonial wars (and would again be utilized in the coming World Wars). In the age of nationalism there was pressure for a nation to acquire an empire as a status symbol; the idea of "greatness" became linked with the sense of duty underlying many nations' strategies.[12]

text: In the early 1880s, Pierre Savorgnan de Brazza was exploring the Kingdom of Kongo for France, at the same time Henry Morton Stanley explored it on behalf of Léopold II of Belgium, who would have it as his personal Congo Free State (see section below). France occupied Tunisia in May 1881, which may have convinced Italy to join the German-Austrian Dual Alliance in 1882, thus forming the Triple Alliance. The same year, Britain occupied Egypt (hitherto an autonomous state owing nominal fealty to the Ottoman Empire), which ruled over Sudan and parts of Chad, Eritrea, and Somalia. In 1884, Germany declared Togoland, the Cameroons and South West Africa to be under its protection; and France occupied Guinea. French West Africa (AOF) was founded in 1895, and French Equatorial Africa in 1910.[13][14]

text: Germany's Weltpolitik[edit]

text: Germany was hardly a colonial power before the New Imperialism period, but would eagerly participate in this race. Fragmented in various states, Germany was only unified under Prussia's rule after the 1866 Battle of Königgrätz and the 1870 Franco-Prussian War. A rising industrial power close on the heels of Britain, Germany began its world expansion in the 1880s. After isolating France by the Dual Alliance with Austria-Hungary and then the 1882 Triple Alliance with Italy, Chancellor Otto von Bismarck proposed the 1884–85 Berlin Conference, which set the rules of effective control of a foreign territory. Weltpolitik (world policy) was the foreign policy adopted by Kaiser Wilhelm II in 1890, with the aim of transforming Germany into a global power through aggressive diplomacy, the acquisition of overseas colonies, and the development of a large navy.

text: Some Germans, claiming themselves of Friedrich List's thought, advocated expansion in the Philippines and Timor; others proposed to set themselves up in Formosa (modern Taiwan), etc. At the end of the 1870s, these isolated voices began to be relayed by a real imperialist policy[citation needed], backed by mercantilist thesis. In 1881, Hübbe-Schleiden, a lawyer, published Deutsche Kolonisation, according to which the "development of national consciousness demanded an independent overseas policy".[15] Pan-Germanism was thus linked to the young nation's imperialist drives[citation needed]. In the beginning of the 1880s, the Deutscher Kolonialverein was created, and got its own magazine in 1884, the Kolonialzeitung. This colonial lobby was also relayed by the nationalist Alldeutscher Verband. Generally, Bismarck was opposed to widespread German colonialism[citation needed], but he had to resign at the insistence of the new German Emperor Wilhelm II on 18 March 1890. Wilhelm II instead adopted a very aggressive policy of colonisation and colonial expansion.

text: Germany's expansionism would lead to the Tirpitz Plan, implemented by Admiral von Tirpitz, who would also champion the various Fleet Acts starting in 1898, thus engaging in an arms race with Britain. By 1914, they had given Germany the second-largest naval force in the world (roughly three-fifths the size of the Royal Navy). According to von Tirpitz, this aggressive naval policy was supported by the National Liberal Party rather than by the conservatives, implying that imperialism was supported by the rising middle classes.[16]

text: Germany became the third-largest colonial power in Africa. Nearly all of its overall empire of 2.6 million square kilometres and 14 million colonial subjects in 1914 was found in its African possessions of Southwest Africa, Togoland, the Cameroons, and Tanganyika. Following the 1904 Entente cordiale between France and the British Empire, Germany tried to isolate France in 1905 with the First Moroccan Crisis. This led to the 1905 Algeciras Conference, in which France's influence on Morocco was compensated by the exchange of other territories, and then to the Agadir Crisis in 1911. Along with the 1898 Fashoda Incident between France and Britain, this succession of international crises reveals the bitterness of the struggle between the various imperialist nations, which ultimately led to World War I.

text: Italy's expansion[edit]

text: Italy took possession of parts of Eritrea in 1870[17][18] and 1882. Following its defeat in the First Italo–Ethiopian War (1895–1896), it acquired Italian Somaliland in 1889–90 and the whole of Eritrea (1899). In 1911, it engaged in a war with the Ottoman Empire, in which it acquired Tripolitania and Cyrenaica (modern Libya). In 1919 Enrico Corradini—who fully supported the war, and later merged his group in the early fascist party (PNF)—developed the concept of Proletarian Nationalism, supposed to legitimize Italy's imperialism by a mixture of socialism with nationalism:

text: We must start by recognizing the fact that there are proletarian nations as well as proletarian classes; that is to say, there are nations whose living conditions are subject...to the way of life of other nations, just as classes are. Once this is realised, nationalism must insist firmly on this truth: Italy is, materially and morally, a proletarian nation.[19]

text: The Second Italo-Abyssinian War (1935–36), ordered by the Fascist Benito Mussolini, would actually be one of the last colonial wars (that is, intended to colonise a foreign country, as opposed to wars of national liberation), occupying Ethiopia—which had remained the last independent African territory, apart from Liberia.

text: Crises prior to World War I[edit]

text: Colonisation of the Congo[edit]

text: David Livingstone's explorations, carried on by Henry Morton Stanley, excited imaginations with Stanley's grandiose ideas for colonisation; but these found little support owing to the problems and scale of action required, except from Léopold II of Belgium, who in 1876 had organised the International African Association (the Congo Society). From 1869 to 1874, Stanley was secretly sent by Léopold II to the Congo region, where he made treaties with several African chiefs along the Congo River and by 1882 had sufficient territory to form the basis of the Congo Free State. Léopold II personally owned the colony from 1885 and used it as a source of ivory and rubber.

text: While Stanley was exploring Congo on behalf of Léopold II of Belgium, the Franco-Italian marine officer Pierre de Brazza travelled into the western Congo basin and raised the French flag over the newly founded Brazzaville in 1881, thus occupying today's Republic of the Congo. Portugal, which also claimed the area due to old treaties with the native Kongo Empire, made a treaty with Britain on 26 February 1884 to block off the Congo Society's access to the Atlantic.

text: By 1890 the Congo Free State had consolidated its control of its territory between Leopoldville and Stanleyville, and was looking to push south down the Lualaba River from Stanleyville. At the same time, the British South Africa Company of Cecil Rhodes was expanding north from the Limpopo River, sending the Pioneer Column (guided by Frederick Selous) through Matabeleland, and starting a colony in Mashonaland.

text: To the west, in the land where their expansions would meet, was Katanga, site of the Yeke Kingdom of Msiri. Msiri was the most militarily powerful ruler in the area, and traded large quantities of copper, ivory and slaves — and rumors of gold reached European ears. The scramble for Katanga was a prime example of the period. Rhodes and the BSAC sent two expeditions to Msiri in 1890 led by Alfred Sharpe, who was rebuffed, and Joseph Thomson, who failed to reach Katanga. Leopold sent four CFS expeditions. First, the Le Marinel Expedition could only extract a vaguely worded letter. The Delcommune Expedition was rebuffed. The well-armed Stairs Expedition was given orders to take Katanga with or without Msiri's consent. Msiri refused, was shot, and the expedition cut off his head and stuck it on a pole as a "barbaric lesson" to the people. The Bia Expedition finished the job of establishing an administration of sorts and a "police presence" in Katanga.

text: Thus, the half million square kilometers of Katanga came into Leopold's possession and brought his African realm up to 2,300,000 square kilometres (890,000 sq mi), about 75 times larger than Belgium. The Congo Free State imposed such a terror regime on the colonised people, including mass killings and forced labor, that Belgium, under pressure from the Congo Reform Association, ended Leopold II's rule and annexed it in 1908 as a colony of Belgium, known as the Belgian Congo.

text: The brutality of King Leopold II of Belgium in his former colony of the Congo Free State,[20][21] now the Democratic Republic of the Congo, was well documented; up to 8 million of the estimated 16 million native inhabitants died between 1885 and 1908.[22] According to the former British diplomat Roger Casement, this depopulation had four main causes: "indiscriminate war", starvation, reduction of births and diseases.[23] Sleeping sickness ravaged the country and must also be taken into account for the dramatic decrease in population; it has been estimated that sleeping sickness and smallpox killed nearly half the population in the areas surrounding the lower Congo River.[24]

text: Estimates of the total death toll vary considerably. As the first census did not take place until 1924, it is difficult to quantify the population loss of the period. Casement's report set it at three million.[25] William Rubinstein wrote: "More basically, it appears almost certain that the population figures given by Hochschild are inaccurate. There is, of course, no way of ascertaining the population of the Congo before the twentieth century, and estimates like 20 million are purely guesses. Most of the interior of the Congo was literally unexplored if not inaccessible."[26] See Congo Free State for further details including numbers of victims.

text: A similar situation occurred in the neighbouring French Congo. Most of the resource extraction was run by concession companies, whose brutal methods, along with the introduction of disease, resulted in the loss of up to 50 percent of the indigenous population.[27] The French government appointed a commission, headed by de Brazza, in 1905 to investigate the rumoured abuses in the colony. However, de Brazza died on the return trip, and his "searingly critical" report was neither acted upon nor released to the public.[28] In the 1920s, about 20,000 forced laborers died building a railroad through the French territory.[29]

text: Suez Canal[edit]

text: French diplomat Ferdinand de Lesseps had obtained many concessions from Isma'il Pasha, the Khedive of Egypt and Sudan, in 1854–56, to build the Suez Canal. Some sources estimate the workforce at 30,000,[30] but others estimate that 120,000 workers died over the ten years of construction due to malnutrition, fatigue and disease, especially cholera.[31] Shortly before its completion in 1869, Khedive Isma'il borrowed enormous sums from British and French bankers at high rates of interest. By 1875, he was facing financial difficulties and was forced to sell his block of shares in the Suez Canal. The shares were snapped up by Britain, under its Prime Minister, Benjamin Disraeli, who sought to give his country practical control in the management of this strategic waterway. When Isma'il repudiated Egypt's foreign debt in 1879, Britain and France seized joint financial control over the country, forcing the Egyptian ruler to abdicate, and installing his eldest son Tewfik Pasha in his place. The Egyptian and Sudanese ruling classes did not relish foreign intervention.

text: During the 1870s, European initiatives against the slave trade caused an economic crisis in northern Sudan, precipitating the rise of Mahdist forces.[32] In 1881, the Mahdist revolt erupted in Sudan under Muhammad Ahmad, severing Tewfik's authority in Sudan. The same year, Tewfik suffered an even more perilous rebellion by his own Egyptian army in the form of the Urabi Revolt. In 1882, Tewfik appealed for direct British military assistance, commencing Britain's administration of Egypt. A joint British-Egyptian military force ultimately defeated the Mahdist forces in Sudan in 1898. Thereafter, Britain (rather than Egypt) seized effective control of Sudan.

text: Berlin Conference (1884–85)[edit]

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text: The occupation of Egypt, and the acquisition of the Congo were the first major moves in what came to be a precipitous scramble for African territory. In 1884, Otto von Bismarck convened the 1884–85 Berlin Conference to discuss the African problem. The diplomats put on a humanitarian façade by condemning the slave trade, prohibiting the sale of alcoholic beverages and firearms in certain regions, and by expressing concern for missionary activities. More importantly, the diplomats in Berlin laid down the rules of competition by which the great powers were to be guided in seeking colonies. They also agreed that the area along the Congo River was to be administered by Léopold II of Belgium as a neutral area, known as the Congo Free State, in which trade and navigation were to be free. No nation was to stake claims in Africa without notifying other powers of its intentions. No territory could be formally claimed prior to being effectively occupied. However, the competitors ignored the rules when convenient and on several occasions war was only narrowly avoided.[33]

text: Britain's administration of Egypt and South Africa[edit]

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text: Britain's administration of Egypt and the Cape Colony contributed to a preoccupation over securing the source of the Nile River. Egypt was overrun by British forces in 1882 (although not formally declared a protectorate until 1914, and never an actual colony); Sudan, Nigeria, Kenya and Uganda were subjugated in the 1890s and early 20th century; and in the south, the Cape Colony (first acquired in 1795) provided a base for the subjugation of neighboring African states and the Dutch Afrikaner settlers who had left the Cape to avoid the British and then founded their own republics. Theophilus Shepstone annexed the South African Republic (or Transvaal) in 1877 for the British Empire, after it had been independent for twenty years. In 1879, after the Anglo-Zulu War, Britain consolidated its control of most of the territories of South Africa. The Boers protested, and in December 1880 they revolted, leading to the First Boer War (1880–81). British Prime Minister William Gladstone signed a peace treaty on 23 March 1881, giving self-government to the Boers in the Transvaal. The Jameson Raid of 1895 was a failed attempt by the British South Africa Company and the Johannesburg Reform Committee to overthrow the Boer government in the Transvaal. The Second Boer War, fought between 1899 and 1902, was about control of the gold and diamond industries; the independent Boer republics of the Orange Free State and the South African Republic (or Transvaal) were this time defeated and absorbed into the British Empire.

text: The French thrust into the African interior was mainly from the coasts of West Africa (modern day Senegal) eastward, through the Sahel along the southern border of the Sahara, a huge desert covering most of present-day Senegal, Mali, Niger, and Chad. Their ultimate aim was to have an uninterrupted colonial empire from the Niger River to the Nile, thus controlling all trade to and from the Sahel region, by virtue of their existing control over the Caravan routes through the Sahara. The British, on the other hand, wanted to link their possessions in Southern Africa (modern South Africa, Botswana, Zimbabwe, Lesotho, Swaziland, and Zambia), with their territories in East Africa (modern Kenya), and these two areas with the Nile basin. The Sudan (which in those days included most of present-day Uganda) was the key to the fulfillment of these ambitions, especially since Egypt was already under British control. This "red line" through Africa is made most famous by Cecil Rhodes. Along with Lord Milner, the British colonial minister in South Africa, Rhodes advocated such a "Cape to Cairo" empire, linking the Suez Canal to the mineral-rich Southern part of the continent by rail. Though hampered by German occupation of Tanganyika until the end of World War I, Rhodes successfully lobbied on behalf of such a sprawling African empire.

text: If one draws a line from Cape Town to Cairo (Rhodes's dream), and one from Dakar to the Horn of Africa (now Ethiopia, Eritrea, Djibouti and Somalia), (the French ambition), these two lines intersect somewhere in eastern Sudan near Fashoda, explaining its strategic importance. In short, Britain had sought to extend its East African empire contiguously from Cairo to the Cape of Good Hope, while France had sought to extend its own holdings from Dakar to the Sudan, which would enable its empire to span the entire continent from the Atlantic Ocean to the Red Sea.

text: A French force under Jean-Baptiste Marchand arrived first at the strategically located fort at Fashoda, soon followed by a British force under Lord Kitchener, commander in chief of the British Army since 1892. The French withdrew after a standoff and continued to press claims to other posts in the region. In March 1899, the French and British agreed that the source of the Nile and Congo Rivers should mark the frontier between their spheres of influence.[citation needed]

text: Moroccan Crisis[edit]

text: Although the 1884–85 Berlin Conference had set the rules for the Scramble for Africa, it had not weakened the rival imperialists. The 1898 Fashoda Incident, which had seen France and the British Empire on the brink of war, ultimately led to the signature of the Entente Cordiale of 1904, which countered the influence of the European powers of the Triple Alliance. As a result, the new German Empire decided to test the solidity of such influence, using the contested territory of Morocco as a battlefield.

text: Thus, Kaiser Wilhelm II visited Tangiers on 31 March 1905 and made a speech in favor of Moroccan independence, challenging French influence in Morocco. France's influence in Morocco had been reaffirmed by Britain and Spain in 1904. The Kaiser's speech bolstered French nationalism, and with British support the French foreign minister, Théophile Delcassé, took a defiant line. The crisis peaked in mid-June 1905, when Delcassé was forced out of the ministry by the more conciliation-minded premier Maurice Rouvier. But by July 1905 Germany was becoming isolated and the French agreed to a conference to solve the crisis. Both France and Germany continued to posture up until the conference, with Germany mobilizing reserve army units in late December and France actually moving troops to the border in January 1906.

text: The 1906 Algeciras Conference was called to settle the dispute. Of the thirteen nations present, the German representatives found their only supporter was Austria-Hungary. France had firm support from Britain, the US, Russia, Italy and Spain. The Germans eventually accepted an agreement, signed on 31 May 1906, whereby France yielded certain domestic changes in Morocco but retained control of key areas.

text: However, five years later the Second Moroccan Crisis (or Agadir Crisis) was sparked by the deployment of the German gunboat Panther to the port of Agadir on 1 July 1911. Germany had started to attempt to surpass Britain's naval supremacy—the British navy had a policy of remaining larger than the next two naval fleets in the world combined. When the British heard of the Panther's arrival in Morocco, they wrongly believed that the Germans meant to turn Agadir into a naval base on the Atlantic.

text: The German move was aimed at reinforcing claims for compensation for acceptance of effective French control of the North African kingdom, where France's pre-eminence had been upheld by the 1906 Algeciras Conference. In November 1911 a convention was signed under which Germany accepted France's position in Morocco in return for territory in the French Equatorial African colony of Middle Congo (now the Republic of the Congo).

text: France and Spain subsequently established a full protectorate over Morocco (30 March 1912), ending what remained of the country's formal independence. Furthermore, British backing for France during the two Moroccan crises reinforced the Entente between the two countries and added to Anglo-German estrangement, deepening the divisions that would culminate in the First World War.

text: Dervish resistance[edit]

text: Following the Berlin Conference at the end of the 19th century, the British, Italians, and Ethiopians sought to claim lands owned by the Somalis such as the Warsangali Sultanate, the Ajuran Sultanate and the Gobroon Dynasty.

text: The Dervish State was a state established by Mohammed Abdullah Hassan, a Somali religious leader who gathered Muslim soldiers from across the Horn of Africa and united them into a loyal army known as the Dervishes. This Dervish army enabled Hassan to carve out a powerful state through conquest of lands sought after by the Ethiopians and the European powers. The Dervish State successfully repulsed the British Empire four times and forced it to retreat to the coastal region.[34] Due to these successful expeditions, the Dervish State was recognized as an ally by the Ottoman and German empires. The Turks also named Hassan Emir of the Somali nation,[35] and the Germans promised to officially recognize any territories the Dervishes were to acquire.[36]

text: After a quarter of a century of holding the British at bay, the Dervishes were finally defeated in 1920 as a direct consequence of Britain's use of aircraft.[37]

text: Herero Wars and the Maji-Maji Rebellion[edit]

text: Between 1904 and 1908, Germany's colonies in German South-West Africa and German East Africa were rocked by separate, contemporaneous native revolts against their rule. In both territories the threat to German rule was quickly defeated once large-scale reinforcements from Germany arrived, with the Herero rebels in German South-West Africa being defeated at the Battle of Waterberg and the Maji-Maji rebels in German East Africa being steadily crushed by German forces slowly advancing through the countryside, with the natives resorting to guerrilla warfare. German efforts to clear the bush of civilians in German South-West Africa then resulted in a genocide of the population.

text: In total, as many as 65,000 Herero (80% of the total Herero population), and 10,000 Namaqua (50% of the total Namaqua population) either starved, died of thirst, or were worked to death in camps such as Shark Island Concentration Camp between 1904 and 1908. Characteristic of this genocide was death by starvation and the poisoning of the population's wells whilst they were trapped in the Namib Desert.

text: Colonial encounter[edit]

text: Colonial consciousness and exhibitions[edit]

text: Colonial lobby[edit]

text: In its earlier stages, imperialism was generally the act of individual explorers as well as some adventurous merchantmen. The colonial powers were a long way from approving without any dissent the expensive adventures carried out abroad. Various important political leaders, such as Gladstone, opposed colonisation in its first years. However, during his second premiership between 1880 and 1885 he could not resist the colonial lobby in his cabinet, and thus did not execute his electoral promise to disengage from Egypt. Although Gladstone was personally opposed to imperialism, the social tensions caused by the Long Depression pushed him to favor jingoism: the imperialists had become the "parasites of patriotism" (John A. Hobson).[38] In France, then Radical politician Georges Clemenceau also adamantly opposed himself to it: he thought colonisation was a diversion from the "blue line of the Vosges" mountains, that is revanchism and the patriotic urge to reclaim the Alsace-Lorraine region which had been annexed by the German Empire with the 1871 Treaty of Frankfurt. Clemenceau actually made Jules Ferry's cabinet fall after the 1885 Tonkin disaster. According to Hannah Arendt in The Origins of Totalitarianism (1951), this expansion of national sovereignty on overseas territories contradicted the unity of the nation state which provided citizenship to its population. Thus, a tension between the universalist will to respect human rights of the colonised people, as they may be considered as "citizens" of the nation state, and the imperialist drive to cynically exploit populations deemed inferior began to surface. Some, in colonising countries, opposed what they saw as unnecessary evils of the colonial administration when left to itself; as described in Joseph Conrad's Heart of Darkness (1899)—published around the same time as Kipling's The White Man's Burden—or in Louis-Ferdinand Céline's Journey to the End of the Night (1932).

text: Colonial lobbies emerged to legitimize the Scramble for Africa and other expensive overseas adventures. In Germany, France, and Britain, the middle class often sought strong overseas policies to ensure the market's growth. Even in lesser powers, voices like Enrico Corradini claimed a "place in the sun" for so-called "proletarian nations", bolstering nationalism and militarism in an early prototype of fascism.

text: Colonial propaganda and jingoism[edit]

text: Colonial exhibitions[edit]

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text: However, by the end of World War I the colonial empires had become very popular almost everywhere in Europe: public opinion had been convinced of the needs of a colonial empire, although most of the metropolitans would never see a piece of it. Colonial exhibitions had been instrumental in this change of popular mentalities brought about by the colonial propaganda, supported by the colonial lobby and by various scientists. Thus, the conquest of territories were inevitably followed by public displays of the indigenous people for scientific and leisure purposes. Karl Hagenbeck, a German merchant in wild animals and a future entrepreneur of most Europeans zoos, thus decided in 1874 to exhibit Samoa and Sami people as "purely natural" populations. In 1876, he sent one of his collaborators to the newly conquered Egyptian Sudan to bring back some wild beasts and Nubians. Presented in Paris, London, and Berlin these Nubians were very successful. Such "human zoos" could be found in Hamburg, Antwerp, Barcelona, London, Milan, New York City, Paris, etc., with 200,000 to 300,000 visitors attending each exhibition. Tuaregs were exhibited after the French conquest of Timbuktu (visited by René Caillié, disguised as a Muslim, in 1828, thereby winning the prize offered by the French Société de Géographie); Malagasy after the occupation of Madagascar; Amazons of Abomey after Behanzin's mediatic defeat against the French in 1894. Not used to the climatic conditions, some of the indigenous exposed died, such as some Galibis in Paris in 1892.[39]

text: Geoffroy de Saint-Hilaire, director of the Parisian Jardin d'acclimatation, decided in 1877 to organize two "ethnological spectacles", presenting Nubians and Inuit. The public of the Jardin d'acclimatation doubled, with a million paying entrances that year, a huge success for these times. Between 1877 and 1912, approximately thirty "ethnological exhibitions" were presented at the Jardin zoologique d'acclimatation.[40] "Negro villages" would be presented in Paris' 1878 and 1879 World's Fair; the 1900 World's Fair presented the famous diorama "living" in Madagascar, while the Colonial Exhibitions in Marseilles (1906 and 1922) and in Paris (1907 and 1931) would also display human beings in cages, often nudes or quasi-nudes.[41] Nomadic "Senegalese villages" were also created, thus displaying the power of the colonial empire to all the population.

text: In the US, Madison Grant, head of the New York Zoological Society, exposed Pygmy Ota Benga in the Bronx Zoo alongside the apes and others in 1906. At the behest of Grant, a prominent scientific racist and eugenicist, zoo director Hornaday placed Ota Benga in a cage with an orangutan and labeled him "The Missing Link" in an attempt to illustrate Darwinism, and in particular that Africans like Ota Benga are closer to apes than were Europeans. Other colonial exhibitions included the 1924 British Empire Exhibition and the successful 1931 Paris "Exposition coloniale".

text: Countering disease[edit]

text: From the beginning of the 20th century onward, the elimination or control of disease in tropical countries became a driving force for all colonial powers.[42] The sleeping sickness epidemic in Africa was arrested due to mobile teams systematically screening millions of people at risk.[43] In the 20th century, Africa saw the biggest increase in its population due to lessening of the mortality rate in many countries due to peace, famine relief, medicine, and above all, the end or decline of the slave trade.[44] Africa's population has grown from 120 million in 1900[45] to over 1 billion today.[46]

text: Colonialism leading to World War I[edit]

text: During the New Imperialism period, by the end of the 19th century, Europe added almost 9,000,000 square miles (23,000,000 km2) – one-fifth of the land area of the globe – to its overseas colonial possessions. Europe's formal holdings now included the entire African continent except Ethiopia, Liberia, and Saguia el-Hamra, the latter of which would be integrated into Spanish Sahara. Between 1885 and 1914, Britain took nearly 30% of Africa's population under its control; 15% for France, 11% for Portugal, 9% for Germany, 7% for Belgium and 1% for Italy.[citation needed] Nigeria alone contributed 15 million subjects, more than in the whole of French West Africa or the entire German colonial empire. It was paradoxical that Britain, the staunch advocate of free trade, emerged in 1914 with not only the largest overseas empire thanks to its long-standing presence in India, but also the greatest gains in the "scramble for Africa", reflecting its advantageous position at its inception. In terms of surface area occupied, the French were the marginal victors but much of their territory consisted of the sparsely populated Sahara.

text: The political imperialism followed the economic expansion, with the "colonial lobbies" bolstering chauvinism and jingoism at each crisis in order to legitimize the colonial enterprise. The tensions between the imperial powers led to a succession of crises, which finally exploded in August 1914, when previous rivalries and alliances created a domino situation that drew the major European nations into World War I. Austria-Hungary attacked Serbia to avenge the murder by Serbian agents of Austrian crown prince Francis Ferdinand, Russia would mobilize to assist allied Serbia, Germany would intervene to support Austria-Hungary against Russia. Since Russia had a military alliance with France against Germany, the German General Staff, led by General von Moltke decided to realize the well prepared Schlieffen Plan to invade and quickly knock France out of the war before turning against Russia in what was expected to be a long campaign. This required an invasion of Belgium which brought Britain into the war against Germany, Austria-Hungary and their allies. German U-Boat campaigns against ships bound for Britain eventually drew the United States into what had become World War I. Moreover, using the Anglo-Japanese Alliance as an excuse, Japan leaped onto this opportunity to conquer German interests in China and the Pacific to become the dominating power in the Western Pacific, setting the stage for the Second Sino-Japanese War (starting in 1937) and eventually World War II.

text: African colonies listed by colonising power[edit]

text: Belgium[edit]

text:
Congo Free State and Belgian Congo (today's Democratic Republic of the Congo)
Ruanda-Urundi (comprising modern Rwanda and Burundi, 1922–62)

text: France[edit]

text:
French West Africa :

Mauritania
Senegal
Albreda (1681–1857, now part of Gambia)
French Sudan (now Mali)
French Guinea (now Guinea)
Ivory Coast
Niger
French Upper Volta (now Burkina Faso)
French Dahomey (now Benin)
French Togoland (1916–60, now Togo)
Enclaves of Forcados and Badjibo (in modern Nigeria)

French Equatorial Africa :

Gabon
French Cameroun (1922–60)
French Congo (now Republic of the Congo)
Oubangui-Chari (now Central African Republic)
Chad

French North Africa :

French Algeria
French Protectorate of Tunisia
French Morocco
Fezzan-Ghadames (1943–1951) (administration given by the UNO after its conquest by Charles de Gaulle)
Egypt (ownership (1798–1801)) (Condominium of France and the United Kingdom) (1876–1882)[47]

French East Africa :

French Madagascar
Comoros
Scattered islands in the Indian Ocean
French Somaliland (now Djibouti)
Isle de France (1715-1810) (now Mauritius)

text: Germany[edit]

text:
German Kamerun (now Cameroon and part of Nigeria, 1884–1916)
German East Africa (now Rwanda, Burundi and most of Tanzania, 1885–1919)
German South-West Africa (now Namibia, 1884–1915)
German Togoland (now Togo and eastern part of Ghana, 1884–1914)

text: After the First World War, Germany's possessions were partitioned among Britain (which took a sliver of western Cameroon, Tanzania, western Togo, and Namibia), France (which took most of Cameroon and eastern Togo) and Belgium (which took Rwanda and Burundi).

text: Italy[edit]

text:
Italian Libya
Italian Eritrea
Italian Somaliland

text: Later, during the Interwar period, with the Second Italo-Ethiopian War Italy would annex Ethiopia, which formed together with Eritrea and Italian Somaliland the Italian East Africa (A.O.I., "Africa Orientale Italiana", also defined by the fascist government as L'Impero).

text: Portugal[edit]

text:
Portuguese West Africa
(now Angola)

Mainland Angola
Portuguese Congo
(now Cabinda Province of Angola)

Portuguese East Africa
(now Mozambique)
Portuguese Guinea
(now Guinea-Bissau)

Portuguese Cape Verde
Portuguese São Tomé and Príncipe

São Tomé Island
Príncipe Island
Fort of São João Baptista de Ajudá
(now Ouidah, in Benin)

text: Spain[edit]

text:
Spanish North Africa

Northern Spanish Morocco

Chefchaouen (Chauen)
Jebala (Yebala)
Kert
Loukkos (Lucus)
Rif

Spanish West Africa

Southern Spanish Morocco

Cape Juby
Ifni

Spanish Sahara
(now Western Sahara)

Río de Oro
Saguia el-Hamra

Spanish Guinea
(now Equatorial Guinea)

Fernando Pó
Río Muni
Annobón

text: United Kingdom[edit]

text: The British were primarily interested in maintaining secure communication lines to India, which led to initial interest in Egypt and South Africa. Once these two areas were secure, it was the intent of British colonialists such as Cecil Rhodes to establish a Cape-Cairo railway and to exploit mineral and agricultural resources. Control of the Nile was viewed as a strategic and commercial advantage.

text:
Egypt
Anglo-Egyptian Sudan (1899–1956)
British Somaliland (now part of Somalia)
British East Africa:

Kenya Colony
Uganda Protectorate
Tanzania :

Tanganyika Territory (1919–61)
Zanzibar

British Mauritius
Bechuanaland (now Botswana)
Southern Rhodesia (now Zimbabwe)
Northern Rhodesia (now Zambia)
British Seychelles
British South Africa

South Africa :

Transvaal Colony
Cape Colony
Colony of Natal
Orange River Colony

South-West Africa (from 1915, now Namibia)

British West Africa

Gambia Colony and Protectorate
Sierra Leone
Nigeria
British Togoland (1916–56, today part of Ghana)
Cameroons (1922–61, now parts of Cameroon and Nigeria)
Gold Coast (British colony) (now Ghana)

Nyasaland (now Malawi)
Basutoland (now Lesotho)
Swaziland

text: Independent states[edit]

text: Liberia was the only nation in Africa that was a colony and a protectorate of the United States. Liberia was founded, colonised, established, and controlled by the American Colonization Society, a private organization established in order to relocate freed African-American and Caribbean slaves from the United States and the Caribbean islands in 1821. Liberia declared its independence from the American Colonization Society on July 26, 1847. Liberia is Africa's oldest democratic republic, and the second-oldest black republic in the world (after Haiti).

text: Ethiopia maintained its independence from Italy after the Battle of Adwa which resulted in the Treaty of Addis Ababa. With the exception of the occupation between 1936 and 1941 by Benito Mussolini's military forces, Ethiopia is Africa's oldest independent nation.

text: Modern Scramble for Africa[edit]

text: The neutrality of this article is disputed. Relevant discussion may be found on the talk page. Please do not remove this message until conditions to do so are met. (July 2016) (Learn how and when to remove this template message)

text: The new scramble for Africa began with the emergence of the Afro-Neo-Liberal capitalist movement in Post-Colonial Africa.[48] When African nations began to gain independence during the Post World War II Era, their post colonial economic structures remained undiversified and linear. In most cases, the bulk of a nation’s economy relied on cash crops or natural resources. The decolonisation process kept independent African nations at the mercy of colonial powers due to structurally-dependent economic relations.[49] Structural Adjustment Programs lead to the privatization and liberalization of many African political and economic systems, forcefully pushing Africa into the global capitalist market.[50] The economic decline in the 1990s fostered democratization by the World Bank intervening in the political and economic affairs of Africa once again.[51] All of these factors led to Africa’s forced development under Western ideological systems of economics and politics.[52]

text: Petro-states[edit]

text: In the era of globalization, many African countries have emerged as petro-states (for example Sudan, Cameroon, Nigeria, Angola). These are nations with an economic and political partnership between transnational oil companies and the ruling elite class in oil-rich African nations.[53] Numerous countries have entered into a neo-imperial relationship with Africa during this time period. Mary Gilmartin notes that “material and symbolic appropriation of space [is] central to imperial expansion and control”; nations in the globalization era who invest in controlling land internationally are engaging in neo-imperialism.[54] Chinese (and other Asian countries) state oil companies have entered Africa’s highly competitive oil sector. China National Petroleum Corporation purchased 40% of Greater Nile Petroleum Operating Company. Furthermore, Sudan exports 50–60% of its domestically produced oil to China, making up 7% of China’s imports. China has also been purchasing equity shares in African oil fields, invested in industry related infrastructure development and acquired continental oil concessions throughout Africa.[55]


document_title: RNA splicing

question_text: which type of rna catalyzes the removal of introns from pre-mrna

root

text: RNA splicing

text: Splicing pathways[edit]

text: Several methods of RNA splicing occur in nature; the type of splicing depends on the structure of the spliced intron and the catalysts required for splicing to occur.

tag: p

text: Spliceosomal[edit]

text: Introns[edit]

text: The word intron is derived from the term intervening sequence, that is, a segment of DNA that "intervenes" between two exons of a gene. The term intron refers to both the DNA sequence within a gene and the corresponding sequence in the unprocessed RNA transcript. As part of the RNA processing pathway, introns are removed by RNA splicing either shortly after or concurrent with transcription.[1] Introns are found in the genes of most organisms and many viruses. They can be located in a wide range of genes, including those that generate proteins, ribosomal RNA (rRNA), and transfer RNA (tRNA).[2]

text: Spliceosomal introns often reside within the sequence of eukaryotic protein-coding genes. Within the intron, a donor site (5' end of the intron), a branch site (near the 3' end of the intron) and an acceptor site (3' end of the intron) are required for splicing. The splice donor site includes an almost invariant sequence GU at the 5' end of the intron, within a larger, less highly conserved region. The splice acceptor site at the 3' end of the intron terminates the intron with an almost invariant AG sequence. Upstream (5'-ward) from the AG there is a region high in pyrimidines (C and U), or polypyrimidine tract. Further upstream from the polypyrimidine tract is the branchpoint, which includes an adenine nucleotide involved in lariat formation.[3][4] The consensus sequence for an intron (in IUPAC nucleic acid notation) is: G-G-[cut]-G-U-R-A-G-U (donor site) ... intron sequence ... Y-U-R-A-C (branch sequence 20-50 nucleotides upstream of acceptor site) ... Y-rich-N-C-A-G-[cut]-G (acceptor site).[5] However, it is noted that the specific sequence of intronic splicing elements and the number of nucleotides between the branchpoint and the nearest 3’ acceptor site affect splice site selection.[6][7] Also, point mutations in the underlying DNA or errors during transcription can activate a cryptic splice site in part of the transcript that usually is not spliced. This results in a mature messenger RNA with a missing section of an exon. In this way, a point mutation, which might otherwise affect only a single amino acid, can manifest as a deletion or truncation in the final protein.

text: Formation and activity[edit]

text: Splicing is catalyzed by the spliceosome, a large RNA-protein complex composed of five small nuclear ribonucleoproteins (snRNPs, pronounced 'snurps'). Assembly and activity of the spliceosome occurs during transcription of the pre-mRNA. The RNA components of snRNPs interact with the intron and are involved in catalysis. Two types of spliceosomes have been identified (major and minor) which contain different snRNPs.

text: The major spliceosome splices introns containing GU at the 5' splice site and AG at the 3' splice site. It is composed of the U1, U2, U4, U5, and U6 snRNPs and is active in the nucleus. In addition, a number of proteins including U2 small nuclear RNA auxiliary factor 1 (U2AF35), U2AF2 (U2AF65)[8] and SF1 are required for the assembly of the spliceosome.[4][9] The spliceosome forms different complexes during the splicing process:[10]

tag: ul

text:
Complex E

The U1 snRNP binds to the GU sequence at the 5' splice site of an intron;
Splicing factor 1 binds to the intron branch point sequence;
U2AF1 binds at the 3' splice site of the intron;
U2AF2 binds to the polypyrimidine tract;[11]

text:
Complex A (pre-spliceosome)

The U2 snRNP displaces SF1 and binds to the branch point sequence and ATP is hydrolyzed;

text:
Complex B (pre-catalytic spliceosome)

The U5/U4/U6 snRNP trimer binds, and the U5 snRNP binds exons at the 5' site, with U6 binding to U2;

text:
Complex B*

The U1 snRNP is released, U5 shifts from exon to intron, and the U6 binds at the 5' splice site;

text:
Complex C (catalytic spliceosome)

U4 is released, U6/U2 catalyzes transesterification, making the 5'-end of the intron ligate to the A on intron and form a lariat, U5 binds exon at 3' splice site, and the 5' site is cleaved, resulting in the formation of the lariat;

text:
Complex C* (post-spliceosomal complex)

U2/U5/U6 remain bound to the lariat, and the 3' site is cleaved and exons are ligated using ATP hydrolysis. The spliced RNA is released, the lariat is released and degraded,[12] and the snRNPs are recycled.

text:
The minor spliceosome is very similar to the major spliceosome, but instead it splices out rare introns with different splice site sequences. While the minor and major spliceosomes contain the same U5 snRNP, the minor spliceosome has different but functionally analogous snRNPs for U1, U2, U4, and U6, which are respectively called U11, U12, U4atac, and U6atac.[14]
Trans-splicing is a form of splicing that joins two exons that are not within the same RNA transcript.[15]

text: Recursive splicing[edit]

text: In most cases, splicing removes introns as single units from precursor mRNA transcripts. However, in some cases, especially in mRNAs with very long introns, splicing happens in steps, with part of an intron removed and then the remaining intron is spliced out in a following step. This has been found first in the Ultrabithorax (Ubx) gene of the fruit fly, Drosophila melanogaster, and a few other Drosophila genes, but cases in human have been reported as well.[16][17]

text: Self-splicing[edit]

text: Self-splicing occurs for rare introns that form a ribozyme, performing the functions of the spliceosome by RNA alone. There are three kinds of self-splicing introns, Group I, Group II and Group III. Group I and II introns perform splicing similar to the spliceosome without requiring any protein. This similarity suggests that Group I and II introns may be evolutionarily related to the spliceosome. Self-splicing may also be very ancient, and may have existed in an RNA world present before protein.

text: Two transesterifications characterize the mechanism in which group I introns are spliced:

text:
3'OH of a free guanine nucleoside (or one located in the intron) or a nucleotide cofactor (GMP, GDP, GTP) attacks phosphate at the 5' splice site.
3'OH of the 5' exon becomes a nucleophile and the second transesterification results in the joining of the two exons.

tag: ol

text: The mechanism in which group II introns are spliced (two transesterification reaction like group I introns) is as follows:

text:
The 2'OH of a specific adenosine in the intron attacks the 5' splice site, thereby forming the lariat
The 3'OH of the 5' exon triggers the second transesterification at the 3' splice site, thereby joining the exons together.

text: tRNA splicing[edit]

text: tRNA (also tRNA-like) splicing is another rare form of splicing that usually occurs in tRNA. The splicing reaction involves a different biochemistry than the spliceosomal and self-splicing pathways.

text: In the yeast Saccharomyces cerevisiae, a yeast tRNA splicing endonuclease heterotetramer, composed of TSEN54, TSEN2, TSEN34, and TSEN15, cleaves pre-tRNA at two sites in the acceptor loop to form a 5'-half tRNA, terminating at a 2',3'-cyclic phosphodiester group, and a 3'-half tRNA, terminating at a 5'-hydroxyl group, along with a discarded intron.[18] Yeast tRNA kinase then phosphorylates the 5'-hydroxyl group using adenosine triphosphate. Yeast tRNA cyclic phosphodiesterase cleaves the cyclic phosphodiester group to form a 2'-phosphorylated 3' end. Yeast tRNA ligase adds an adenosine monophosphate group to the 5' end of the 3'-half and joins the two halves together.[19] NAD-dependent 2'-phosphotransferase then removes the 2'-phosphate group.[20][21]

text: Evolution[edit]

text: Splicing occurs in all the kingdoms or domains of life, however, the extent and types of splicing can be very different between the major divisions. Eukaryotes splice many protein-coding messenger RNAs and some non-coding RNAs. Prokaryotes, on the other hand, splice rarely and mostly non-coding RNAs. Another important difference between these two groups of organisms is that prokaryotes completely lack the spliceosomal pathway.

text: Because spliceosomal introns are not conserved in all species, there is debate concerning when spliceosomal splicing evolved. Two models have been proposed: the intron late and intron early models (see intron evolution).

text:
Splicing diversity

Eukaryotes
Prokaryotes

Spliceosomal
+
−

Self-splicing
+
+

tRNA
+
+

text: Biochemical mechanism[edit]

text: Spliceosomal splicing and self-splicing involve a two-step biochemical process. Both steps involve transesterification reactions that occur between RNA nucleotides. tRNA splicing, however, is an exception and does not occur by transesterification.[22]

text: Spliceosomal and self-splicing transesterification reactions occur via two sequential transesterification reactions. First, the 2'OH of a specific branchpoint nucleotide within the intron, defined during spliceosome assembly, performs a nucleophilic attack on the first nucleotide of the intron at the 5' splice site, forming the lariat intermediate. Second, the 3'OH of the released 5' exon then performs a nucleophilic attack at the first nucleotide following the last nucleotide of the intron at the 3' splice site, thus joining the exons and releasing the intron lariat.[23]

text: Alternative splicing[edit]

text: In many cases, the splicing process can create a range of unique proteins by varying the exon composition of the same mRNA. This phenomenon is then called alternative splicing. Alternative splicing can occur in many ways. Exons can be extended or skipped, or introns can be retained. It is estimated that 95% of transcripts from multiexon genes undergo alternative splicing, some instances of which occur in a tissue-specific manner and/or under specific cellular conditions.[24] Development of high throughput mRNA sequencing technology can help quantify the expression levels of alternatively spliced isoforms. Differential expression levels across tissues and cell lineages allowed computational approaches to be developed to predict the functions of these isoforms.[25][26] Given this complexity, alternative splicing of pre-mRNA transcripts is regulated by a system of trans-acting proteins (activators and repressors) that bind to cis-acting sites or "elements" (enhancers and silencers) on the pre-mRNA transcript itself. These proteins and their respective binding elements promote or reduce the usage of a particular splice site. The binding specificity comes from the sequence and structure of the cis-elements, e.g. in HIV-1 there are many donor and acceptor splice sites. Among the various splice sites, ssA7, which is 3' acceptor site, folds into three stem loop structures, i.e. Intronic splicing silencer (ISS), Exonic splicing enhancer (ESE), and Exonic splicing silencer (ESSE3). Solution structure of Intronic splicing silencer and its interaction to host protein hnRNPA1 give insight into specific recognition.[27] However, adding to the complexity of alternative splicing, it is noted that the effects of regulatory factors are many times position-dependent. For example, a splicing factor that serves as a splicing activator when bound to an intronic enhancer element may serve as a repressor when bound to its splicing element in the context of an exon, and vice versa.[28] In addition to the position-dependent effects of enhancer and silencer elements, the location of the branchpoint (i.e., distance upstream of the nearest 3’ acceptor site) also affects splicing.[6] The secondary structure of the pre-mRNA transcript also plays a role in regulating splicing, such as by bringing together splicing elements or by masking a sequence that would otherwise serve as a binding element for a splicing factor.[29][30]

text: Splicing response to DNA damage[edit]

text: DNA damage affects splicing factors by altering their post-translational modification, localization, expression and activity.[31] Furthermore, DNA damage often disrupts splicing by interfering with its coupling to transcription. DNA damage also has an impact on the splicing and alternative splicing of genes intimately associated with DNA repair.[31] For instance, DNA damages modulate the alternative splicing of the DNA repair genes Brca1 and Ercc1.

text: Experimental manipulation of splicing[edit]

text: Splicing events can be experimentally altered[32][33] by binding steric-blocking antisense oligos such as Morpholinos or Peptide nucleic acids to snRNP binding sites, to the branchpoint nucleotide that closes the lariat,[34] or to splice-regulatory element binding sites.[35]

text: Splicing errors and variation[edit]

text: It has been suggested that one third of all disease-causing mutations impact on splicing.[28] Common errors include:

text:
Mutation of a splice site resulting in loss of function of that site. Results in exposure of a premature stop codon, loss of an exon, or inclusion of an intron.
Mutation of a splice site reducing specificity. May result in variation in the splice location, causing insertion or deletion of amino acids, or most likely, a disruption of the reading frame.
Displacement of a splice site, leading to inclusion or exclusion of more RNA than expected, resulting in longer or shorter exons.

text: Although many splicing errors are safeguarded by a cellular quality control mechanism termed nonsense-mediated mRNA decay (NMD),[36] a number of splicing-related diseases also exist, as suggested above.[37]

text: Allelic differences in mRNA splicing are likely to be a common and important source of phenotypic diversity at the molecular level, in addition to their contribution to genetic disease susceptibility. Indeed, genome-wide studies in humans have identified a range of genes that are subject to allele-specific splicing.

text: Protein splicing[edit]

text: In addition to RNA, proteins can undergo splicing. Although the biomolecular mechanisms are different, the principle is the same: parts of the protein, called inteins instead of introns, are removed. The remaining parts, called exteins instead of exons, are fused together. Protein splicing has been observed in a wide range of organisms, including bacteria, archaea, plants, yeast and humans.[38]


document_title: Mann (film)

question_text: nasha ye pyar ka nasha copied from which italian song

text: Mann (film)

text: Plot[edit]

text: Dev Karan Singh (Aamir Khan), a casanova and ambitious painter deep in debt, agrees to marry Anita (Deepti Bhatnagar), the daughter of Singhania (Dalip Tahil), a rich tycoon. Priya (Manisha Koirala), a music teacher for children, is engaged to Raj (Anil Kapoor), whom she has agreed to marry because he had helped her when she was in need. Priya and Dev meet on a cruise and fall in love. However, due to them already being engaged to other people, they agree to work everything out and meet in 6 months on Valentine's Day to get married.

text: During the 6 months, Dev breaks off his engagement with Anita and starts working hard. Driven by his love, he creates and auctions beautiful paintings, and becomes very successful. On the other hand, Priya realizes that leaving Raj will be wrong and sadly writes a letter to Dev, explaining everything. When Raj gets the letter instead, he supports Priya and convinces her to go to Dev. Things take a bad turn when, on her way to meet him, Priya gets hit by a car and gets her legs amputated. Dev, who waited all night for Priya, believes she has rejected him, as he does not know of her accident. Priya forbids Raj from telling Dev, not wanting to become a burden on him. However, both Dev and Priya still love and pine for each other.

text: Though heartbroken, Dev continues his career and goes on to become a famous painter. One day, Priya attends his art exhibition and wishes to buy a painting of his, a sentimental picture of her speaking to his beloved grandmother, whom Dev had introduced to her during the cruise. Dev had said that he would not sell the painting as it was only for Priya, but upon hearing that the girl who wanted it understands the emotion behind the picture and is disabled, he tells the host of the exhibition to give the painting to her for free.

text: One day, he visits Priya to give her an anklet that his grandmother, who recently died, had wished for Priya to have when she became Dev's bride. Initially unaware of Priya's condition, Dev realizes the truth when he sees the painting in another room. The two embrace tearfully and he assures Priya that he will love her no matter what. Dev and Priya then get married and live happily ever after.

text: Cast[edit]

text:
Aamir Khan as Dev Karan Singh
Manisha Koirala as Priya Verma
Sharmila Tagore as Dev's grandmother
Anil Kapoor as Raj
Deepti Bhatnagar as Anita Singhania
Dalip Tahil as Pratap Rai Singhania
Shama Sikander[2] as Kamini
Neeraj Vora as Nattu
Satyen Kappu as Raj's uncle
Anant Mahadevan as Creditor
Paresh Ganatra
Rani Mukerji ... Special Appearance in the song 'Kali Nagin Ke Jaisi Zulfain'
Ketki Dave ... as Madhu, Priya's friend (special appearance)
Amrieshwar Kumar as child actor in school.

text: Box office[edit]

text: The film was an average grosser at box office. [3]

text: Casting[edit]

text: Originally, Aishwarya Rai was offered the lead role of Priya and after she had declined due to prior date commitments, Manisha Koirala was roped in. Interestingly, the movie Hum Dil De Chuke Sanam starring Aishwarya was released at the same time and became an unexpected hit.

text: Soundtrack[edit]

text: The music for the movie was composed by the composer duo Sanjeev Darshan while lyrics were by Sameer.The songs were very popular among the masses. It remains third selling soundtrack of the year 1999. The songs like 'Nasha yeh pyar ka' 'Khushiyon aur gham' 'Mera nann' were chartbusters . Most of the songs were copied from another singers famous songs without their copyright also not credited like The song 'Nasha yeh pyar ka' is based on the melody of the Italian song L'Italiano by Toto Cutugno and "Tinak Tin Tana" Lifted from song "Yang Sedang Sedang Saja" by Malaysian singer Iwan.

text: Track listing[edit]

text:
#
Title
Singer(s)
Note(s)

1
"Chaha Hai Tujhko"
Udit Narayan, Anuradha Paudwal
Lifted from the song "Edho Oru Paatu" from Unnidathil Ennai Koduthen (1998)

2
"Dance Music"
Instrumental

3
"Kehna Hai Tumse"
Udit Narayan, Hema Sardesai
Lifted from song "Liquid" by Jars of Clay (1995)

4
"Kali Nagin Ke Jaisi"
Udit Narayan, Kavita Krishnamurthy
Composition from song "Ya Rayah" by Algerian singer Rachid Taha (1997).

5
"Khushiyan Aur Gham"
Udit Narayan, Anuradha Paudwal
Lifted from song "Come Vorrei" by Italian pop group Ricchi & Poveri (1981)

6
"Kyon Chhupate Ho"
Udit Narayan, Anuradha Paudwal

7
"Mera Mann Kyon Tumhe Chahe"
Udit Narayan, Alka Yagnik

8
"Nasha Yeh Pyar Ka"
Udit Narayan
Composition from the 1983 Italian pop song "L'Italiano" by Toto Cutugno.

9
"Tinak Tin Tana"
Udit Narayan, Alka Yagnik
Lifted from song "Yang Sedang Sedang Saja" by Malaysian singer Iwan

10
"Tumhare Baghair Jeena Kya (Dialogue)"
Aamir Khan
Dialogue