The Coevolution of Human Immunity and Helminthic Parasites

The Coevolution of Human Immunity and Helminthic Parasites Most multicellular organisms, both vertebrate and invertebrate, have an evolutionary history of infestation by extracellular parasitic worms known as helminths. The immune systems of these species have adapted to the stress of helminth infection, or helmnithiasis, through the development of mechanisms to modulate worm load in chronically infested individuals. Most marsupials and mammals, including humans, use a particular immune response mediated by IgE antibodies molecules that identify and neutralize foreign objects to defend against helminths (Poulsen Hummelshoj 2007). In parts of the world where helminthiasis is still prevalent, there is a selective advantage for genes that increase production of IgE antibodies. However, the adaptation to the stress of chronic worm infestation accounts for the maladaptive response to innocuous substance allergy upon removal of the stress. The presence of parasites triggers the production of molecules known as interleukin-10 (IL-10) cytokines that damped the inflammation response. However, in the absence of parasites, IgE antibodies target harmless proteins and the lack of IL-10 production results in a potentially dangerous inflammatory response. In addition to the removal of the early evolutionary stress of helminths, many technology and infrastructure changes in developed countries have increased human exposure to allergens, thus increasing hypersensitivity to seemingly innocuous substances.   Many species, including humans, evolved under the stress of helminth infestation. As early hominids expanded their ecological niche and encountered new foods, they became hosts for an increased number of helminth species, which would have, in turn, evolved with the hominids. Today, humans act as hosts for more than 25 species of helminths (Warren et al 1990). The shift from the hunter-gatherer lifestyle to agriculture increased sedentism and disease, including parasitic infection. Such disease was spread through contact with animals, other humans, and their wastes. The development of agricultural methods such as irrigation and the use fertilizer would have increased the exposure of early humans to soil-transmitted helminths (Cockburn 1971). For contemporary humans, parasitic and infectious diseases are controlled in some areas of the world, while chronic, noninfectious, degenerative diseases are on the rise. Although new technology has allowed some human populations to benefit from the control of infectious disease, many individuals throughout the world are still affected by infection and parasites. Globally, more than two billion people are chronically infected with soil-transmitted helminths such as schistosomes and hookworms (Florh et al. 2008). These numbers indicate there is still selection for protective mechanisms against helminthiasis in a large proportion of the contemporary human population. Given the prolonged mammalian history with parasites, the immune system has evolved protective mechanisms to safeguard the heath of a host in the event of a parasitic infection. When a helminth enters a host, antigens from the parasite diffuse across the hosts internal membranes. Two types of white blood cells, B cells and T cells, recognize antigens in the blood stream. B cells are released into the blood and carried to capillary beds serving the tissues and organs of the lymphatic system a system of vessels and organs that helps balance the fluid content of blood and the surrounding tissues while participating in the bodys defense against invading disease organisms (Russel et al. 2008). T cells are released into the blood and carried to the thymus, an organ of the lymphatic system. The adaptive immune responses are regulated by two mechanisms: antibody-mediated immunity and cell-mediated immunity. During antibody-mediated immunity, derivatives of B cells known as plasma cells secrete antibodies that circulate throughout the blood and lymphatic fluid, recognizing, binding, and removing antigens. Each plasma cell is specific for at least one particular antigen, but some are capable of recognizing any antigen, even if it has never before been encountered. Plasma cells are capable of secreting are five major classes of antibodies. These antibodies are, in order of decreasing concentration, IgG, IgA, IgM, IgD, and IgE (Barnes et al. 1999). Each type of antibody has a specific function in the immune system and IgE is most relevant in combating infection by parasitic worms and mediating many allergic responses such as hay fever, asthma, and hives (Russell et al 2008). Cell-mediated immunity serves as the primary mechanism for killing parasite larvae. During cell-media ted immunity, a subset of T cells becomes activated and, with other cells of the immune system, attacks and kills foreign cells directly. These two mechanisms interact to defend the host against extracellular parasites. Parasitic antigens are first detected by plasma cells in the membranes of the gastrointestinal and respiratory tracts, triggering the production of two types of IgE antibodies: those that are specific for a particular parasite and those that are nonspecific (Grant et al 2008). These antibodies bind to mast cells. Mast cells are a particular type of cell found within many body tissues that contain granules of molecules such as histamine. The mast cells are activated to degranulate when the antigen binds to the attached IgE antibody, causing the internal histamine to be released. The release of histamine causes various physiological changes associated with inflammation (Flohr et al. 2008). The cascade of reactions functions to damage and expel the parasite (Barnes et al. 1999). Inflammation is complex biological process that occurs in vascular tissues as a response to pathogens (such as helminths), damaged cells, or irritants. In an inflammatory response, an individual may experience bronchial constriction, vascular dilation, and an increase in mucous secretions, which lead to the associated symptoms of wheezing, coughing, itching, sneezing, and vomiting. During anaphylaxis, a severe form of inflammation, there is an intense generation of mast cells and release of their mediators. Such a response has effects on various organs and may be fatal. Examples of anaphylaxis-inducing antigens include antibiotics, foods, and foreign proteins, such as venom. Thus the inflammatory response to the presence of a particular antigen may in an of itself harmful to the organism (Florh et al. 2008) Once an immune reaction has run its course and the invading parasites have been eliminated, long-lived T helper cells, derived from the encounter with the antigen remain in an inactive state in the lymphatic system and provide an immunological memory of the foreign antigen (Poulsen Hummelshoj 2007). When a foreign antigen enters the body for a subsequent time, a secondary immune response is triggered. The helper T cells recognize the antigen and secrete small proteins known as cytokines that regulate or assist in an immune response. Helper T cells can be divided into TH1 and TH2 subsets that fulfill separate functions in regulating response to infection. TH1 cells produce the response to intracellular infections while TH2 cells produce responses to extracellular infections and allergens. During helminth infections, the number of TH2 cells is greater than the number of TH1 cells. When TH2 cells detect previously recognized parasitic antigens, they secrete a particular cytokine, or known interleukin-4 (IL-4) (Barnes et al 1999). IL-4 promotes parasite-specific IgE antibody, helper T cell, and mast cell production. The adaptation of the IgE antibody immune response is beneficial during helminth infection. High levels of IgE minimize the number of parasites that infest a host during chronic exposure (Dunne et al. 1992). Individuals infected with helminths may have IgE antibody levels that are up to 100 times greater than the normal level, which typically decrease after anti-helminth treatment (Poulsen Hummelshoj 2007). Additionally, the type of IgE antibody produced may change throughout a humans life to better target a particular parasite. Studies have shown that humans acquire a natural immunity to schistosome infection in adolescence (Grant et al 2008). This natural immunity corresponds to increased levels of IgE from schistosome-specific antigens and decreased production of non-specific IgE. For young children, the greater nonspecific component in IgE production occurs at the expense of schistosome-specific IgE, resulting in a less protective antibody-mediated immune response when compared to adolescents and adults. To establish long-term immunity and because contacts between vector an agent that transmits an infectious disease and host may be infrequent, it is important for the both the host and parasite to maintain chronic infections. Most human parasitic infections last for years and must therefore not overwhelm the host. Parasites produce self-limiting infections that allow the host to defend against lethal infection while maintaining a viable population. One strategy is through concomitant immunity, a response seen in adult schistosomes, where an immune response is induced to limit, but not eliminate, subsequent infections of the host by infective larvae, without causing the rejection of the adult worms (Sher Ottensen 1988). Schistosomes and hookworms also trigger the production of the anti-inflammatory cytokine interleukin-10 (IL-10) in parasite-induced T cells. IL-10 protects the host from extreme mast cell degranulation and the initiation of intense inflammation (Florh et al. 2008). T he level of IL-10 decreases after anti-helminth treatments once the parasite is no longer present to induce production. In summary, the immune system of most mammalian and marsupial hosts is highly adapted to battle parasitic disease. The generation of parasite-specific IgE antibodies by plasma cells initiates an inflammatory response and killer cell activity. During subsequent encounters with an antigen, the synthesis of IgE is controlled by TH2 cells and up-regulated by the cytokine IL-4. The inflammatory response is, however modulated by the release of anti-inflammatory IL-10 cytokines, in order to protect the host from the dangerous effects of intense mast cell degranulation. Through these mechanisms, the more successful human host will produce higher levels of parasite-specific IgE antibodies with which to prevent overwhelming worm infestation. Hosts less proficient at producing sufficiently high levels of parasite-specific IgE antibodies are more likely to succumb to greater worm loads. Allergy is hypersensitivity to a typically innocuous substance. Allergy begins after sensitization of a specific allergen, an antigen that elicits an allergic response. Similar to a helminthic infestation, plasma cells generate IgE antibodies during sensitization that are specific to the allergens to which an individual has been exposed. These IgE antibodies bind to receptors on mast cells. The binding of the allergen to an IgE antibody triggers a cascade of events resembling the immune response to helmthiasis (Zanders et al. 1992). The mast cells degranulate to release mediators, including histamine. Unlike in helminthiasis, in which IgE antibodies are directed at the worm and its by-produces, the allergic response is directed at seemingly innocuous substances. Also, the allergens are not capable of initiating the parasite-induced production of IL-10 that protects the host from the potentially harmful effects of the inflammatory response (Flohr et al. 2008). Thus, humans have adapte d to respond to the outside world in the presence of helminths and in their absence we are unable to modulate the maladaptive inflammatory response that may result in annoying or dangerous symptoms. In industrialized countries, the prevalence of allergies and conditions such as asthma have increased over the last three or four decades (Poulsen Hummelshoj 2007). These countries have better-developed infrastructures that have resulted in the elimination of helminths and an increase in noninfectious disease. Similarly, allergic disease prevalence is increasing in industrializing countries such as India and China (Flohr et al. 2008). Besides the removal of helminths, a significant consequence of modernization is the creation of a microenvironment that increases our exposure to domestic arthropods, such as dust mites, and other pests. Research has shown that there is a positive correlation between level of infestation of household pests and the degree of urbanization (Barnes et al. 1999). Evidence suggests that allergic reactions are less pronounced in individuals infected with helminths. Thus, areas where helminthic infection is endemic typically have lower levels of allergic disease when compared with areas free of helminths (Grant et al. 2008). Studies have consistently found that most helminths investigated imbue their hosts with protective effects during skin prick tests (SPT) tests used to diagnose allergies by eliciting a small, controlled allergic response. However, while all helminths increase the level of IgE antibody produced by a host, infection by schistosomes and hookworms parasites found to trigger the production of IL-10 have the strongest association with protection against allergy and asthma (Flohr et al. 2008). Individuals are more likely to develop asthma during the absence of helminthiasis, or during mild helminthiasis a time during which less parasite-specific IgE antibody is produced than during a chronic infection, and less IL-10 is produce d to reduce inflammation (Lynch 1992). Therefore, helminthiasis and allergy are not likely mutually exclusive, but allergy is much less likely to occur in severe helminthic disease than in mild helminthic disease (Barnes et al. 1999). The removal of helminths from infected populations in Venezuela, Vietnam, and Gabon has shown a resultant increase in allergic skin sensitization during SPT (Florh et al. 2008). Marsh et al. (1980) found that non-European descendents living in developed countries have a higher propensity for allergic response. These results are expected because those individuals likely had a greater genetic propensity to produce IgE, resulting in an increased inflammation response, a decrease in IL-10 production to modulate inflammation, and an increased exposure to inhalant allergens. There is a selective advantage for a predisposition to produce high levels of IgE, as this antibodies serves as a key regulator in the maintenance of helminthic infection in populations that are chronically exposed to parasites. Additionally, it has been found that certain levels of allergens affect people with family histories of allergy, but do not trigger an allergic response in most other people (Sporik et al. 1990). These finding imply that allergy and asthma reactions occur only in genetically susceptible individuals after adequate or persistent exposure to specific allergens. While the total level of serum IgE does not appear to directly reflect natural immunity against asthma in helminth infection-endemic populations, linkage studies have implicated a particular chromosome locus, or region, in controlling asthma and intensity of schistosomiasis in Brazilian and Senegalese populations. Because this same locus is identified with both helminthic infection and for allergy suscept ibility in a number of independent studies, there may be a common genetic basis for host protection against helminthic infection and susceptibility allergic disease (Grant et al. 2008). Through modernization, populations acquire objects that promote allergens such as upholstered furniture, carpeting and domestic pets. The introduction of such objects has been correlated to a rapidly increase the prevalence of asthma in populations with either high or low helminthiasis prevalence. An example in a study by Dowse et al. (1985) showed that asthma incidence increased over ten years within Eastern Highland villages of Papua New Guinea that was attributed to the introduction of wool blankets to the villagers and the sudden and profound exposure to house dust mites within the blankets. Barnes et al. (1997) found that house dust mite allergen concentrations in Barbados were higher in better-built homes, likely because the plumbing contributed to a higher humidity levels that were more conducive to dust mite proliferation than the drier wood homes. During the process of modernization, in addition to the acquisition of homes and objects that increase allergen exposure, the red uction or elimination of helminthiasis, increases the risk of allergic disease more dramatically. Adaptation often results in trade-offs that may compromise an individuals adjustment to his or her environment. The coevolution of helminths and humans shaped the immune response to be highly sensitive to parasitic antigens. This response, which is beneficial to host and parasites, is modulated by many mechanisms. TH2 activation stimulates the production of IL-4 cytokines that trigger production of IgE antibodies. IgE mediate an immune response targeted the antigens released by parasites as well as allergens. Parasites presence triggers anti-inflammatory IL-10 cytokines production by specialized T cells that reduce the inflammatory effects of mast cell degranulation. Removal of the stress helminthiasis also removes the modulation of the inflammatory response through IL-10. Under these conditions, the maladapted response of IgE antibodies reacting to harmless allergens is allergy in the form of disproportioned, potentially dangerous inflammation event. Although levels of IgE are highe st during a parasitic infection or an allergic response, levels are also affected by genetic predisposition. Selective pressures maintain high levels of IgE expression in regions of the world with high helminthiasis prevalence. Through modernization, the stress of helminthiasis has been removed while the stress of allergen exposure has increased. Activation of IgE by innocuous allergens triggers the maladaptive response of an allergic reaction. Individuals who are not infested by helminths with a genetic propensity for high IgE antibody expression are most susceptible to allergic hypersensitivity. In developed countries, decreased helminthiasis prevalence in junction with increased allergen exposure are responsible for the increase in allergic disease prevalence.

Impact of the Collision of the Old and New World on Europeans, Africans, and the Indians Essay

The collision of the New and Old World impacted the Europeans, Africans, and Indigenous people immensely. When the two worlds were introduced to each other they set up trade routes, such as the Columbian Exchange. Although it was new for all three of them, they adjusted well to the changes over time. The introduction of the new foods, animals, diseases, religious matters, etc. made all three groups forced to accommodate. In 1492, Africans along with the Europeans and the Indians, created the Columbian Exchange after Christopher Columbus discovered them. The Africans didn’t have a choice on whether they wanted to come to the New World or not, they were shipped here on tiny boats and due to the death of the Indians the Europeans forced them to do the hard labor without any pay. They worked in farms and/or plantations for their â€Å"masters† who provided them with limited or no rights and they lived in horrible conditions. In return, the Africans received different varieties of crops, for example, potatoes, corn, beans, tomatoes, and many other crops. Europeans also tried to convert both Africans and Indians to Christianity and succeeded. In 1519, Hernan Cortez, plus one hundred men, landed on the island of Cozumel. The men who landed had been enslaved for many years by the Mayans, but finally away. As the Indians settled, they were introduced to a new way of life, soon becoming a â€Å"wide-range hunting society† in which they roamed the open land for buffalo. The Indians had to adjust to slave work as the Europeans had them work in the sugar mills and cane fields. The Europeans brought along a variety of diseases, which soon killed a large group of the Indians, the population started at 1 million and ended at 200. This caused the Indians into taking revenge against the Europeans. They did so by injecting the first sexually transmitted diseases such as syphillis into Europe. Revenge was the only feasible option for the natives because how cruel the Europeans were to them. After the invasion of Hernan Cortes in 1519 and Pizarro in 1533, the Europeans forced the Indians to give them their land and crops. The Europeans used the lands for agriculture and as sources of precious metals, and were able to successfully plant sugar and tobacco plantations. With such great demand for these crops back in Europe there was a necessity for large amounts of labor. Being that most of the indigenous population was either familiar with the land and could easily run away or was wiped out by disease, they couldn’t enslave them. To fill the void left by the indigenous people, the Europeans decided to enslave millions of Africans instead. The Africans weren’t likely to run away because of the unfamiliarity of the land and they wouldn’t know how to feed themselves. In 1545, Hernando de Soto, Francisco Vasquez de Coronado, and Alvar Nunes Cabeza de Vaca foun d silver in Potosoi. The Spanish wealth and power was based on the New World’s gold and silver mines, all the gold and silver made the Spanish in power for about 100 years. The creation of the New World and the Columbian Exchange impacted three ethnicities: the Europeans, the indigenous, and the Africans. Despite the few minor setbacks the Europeans had to go through, such as the â€Å"starving time† and the spread of syphillis, the Europeans were impacted in a very positive way. They were able to find gold and silver and plant many crops, and as a result were able to create the 13 colonies. But the indigenous people did not benefit, most died from disease brought by the Europeans or were tortured and killed. Out of all three groups the Africans had the most negative impact, thousands were forced to work and they were left with no rights. The troubles from the New World still impact America to this day: Africans weren’t granted their rights back until many years later and unfortunately there is still racism, the few indigenous people left live in reservations, and because of the Europeans we have America.

Milton Paradise Lost Commentary

The Renaissance era represents a complete break with the Middle Ages on a political, philosophical, scientifical and theological scale. Indeed, the discovery of new territories and the expeditions of explorers such as Francis Drake, the resurgence of classic writtings, the new scientific outbreaks of Copernicus, Newton and Galilei as well as the Protestant reformation led by Luther and his 95 Theses, triggered a will for knowledge and a quest for truth, thus putting an end to the intellectual hegemony of the Roman Catholic Church in Western Europe. Intellectuals of this era began to think on mankind, looking for answers outside of the Religious constraints. Written during this context of political and religious upheaval, Paradise Lost, an epic poem published in 1667 in ten books, reflects in a way the great changes of the Renaissance. This poem, written by John Milton (December 1608 – November 1674), an English poet, polemicist and civil servant, tell the Christian story of the creation of the Earth, the fall of Satan and the Fall of Man. As a puritan, Milton, who had already attacked the Church for its corruption in Lycidas (1637), give this classical Christian tale puritan traits (such as the protrayal of Eve as a dedicated worker). The passage under study is taken from Book 9 of Paradise lost. In this book, the narrator focuses on the disobedience of Adam and Eve. Satan returns to the Garden of Eden eight days after his banishment by Gabriel to avenge himself. Adam and Eve , preparing for their labors, decide to work separatly , even if Adam is anxious that eve would be an easy prey to Satan’s temptation . In this passage we can see the approach of the Satan , (the snake) and the begining of his temptation that will lead Eve to eat the prohibited fruit. We can clearly see that Milton here uses a mix of Classical and Judeo-christian mythology to tell this tale. We can thus wonder In which way he uses these influences, and why he uses them ? In a first part we will see that the Serpent is a symbol of temptation, and in a second part we wll analyse Milton’s uses of Classical mythology. First, Milton depicts the snake which Satan is possessing, as an incredibly beautiful animal. The main strategy of Satan to try to corrupt the naive mind of Eve is to appear as a magnificent snake. According to the narrator, his physical appearance is so pleasing that never a snake will be as beautiful (on verse 504 â€Å"pleasing was his shape, and lovely, never since a serpent lovelier†). To describe the snake, the narrator use the lexical field of precious material: â€Å"carbuncle his eyes† on verse 500, â€Å"neck of verdent gold† on verse 501. Here Satan seems to have chosen a snake whose appearance can attract Eve only because his similitude with precious material. In a kind of materialist temptation, he tries to appeal to Eve inner sin of cupidity to seduce her. But there is something paradoxical in the way the snake is depicted as a beautiful creature. Indeed, the snake is often assimilated with death, the venom of some species of snakes such as the cobra can kill a man in less than one hour. It also an animal that generate instinctive fear in most human (and occurence of Ophidiophobia can be found everywhere). Here death and beauty become assimilated in one animal. It can thus be perceived as personification of the treachery of satan: behind the beaty of his rethoric, of his discourse hides a terrible fates, the banishment of Man from Heaven. This can also underline the interest of humans for macabre, dark topics. We can see since the dawn of age exemples of this morbid attraction of the Human race: public executions had always (and still do) gathered a lot of people around the death of a single man as well as we can see every day the impact (and also its capacity of generating profits) of a news involving the mysterious death of a movie star, of the murder of a whole family by its patriarch. As we know, Satan has chosen this animal, maybe he relies on this fascination to attract eve attention. We can also notice that the serpent can be interpreted as a phallic symbol. On verse 498 to 502 the snake is described as a â€Å"circular base that towered folds above folds a surprising maze [†¦ ] with burnished neck [†¦ ] erect amidst his circling spires†. Thus the snake appears as a symbol of virility, a phallic form erecting from the ground and can be understood as another strategy of Satan to cajole Eve into trusting him. On the other hand this links made by Milton between sensuality and religion can be seen as a hint of the influence of the metaphysical poetry on his writting: this link beeing one of the interest of the metaphysical poets who were contemporary with Milton. Then the snake can be analysed as a symbol of corruption and of perfidy. Satan uses a disguise, a snake to talk to eve. He uses his beauty, his eloquence to seduce her. His wave movements, looking like a dance, and his attitude (â€Å"Fawning, and licking the ground whereon she strod† on verse 526) can be interpreted as a nuptial parade with the snake laying on the primal instinct (which are restricted by religion) of Man to attract Eve in his trap. It is also interesting to note that the occurence of the word Satan is, as Satan himself, disguised in this passage. Here Satan is refered to as â€Å"the snake† or â€Å"the enemy of mankind† on verse 494. The word Satan can only be seen if the reader looks closely at the poem: an acrostic is hidden from verse 510 to 515. This clever technic reflects the vicious strategy of the Devil in the passage. Satan’s main strategy to talk Eve into beliving that she has the right to eat the apple is flattery. The Devil choose to tempt Eve because he was afraid of Adam’s intelligence which could have be an obstacle to his scheme. Thus he constantly celebrate eve beauty and superiority of others God’s creature to put her in a position where she can feel free to act as she wants. From verse 532 to the end of the passage the snake states that eve is â€Å"sole wonder†, that she is a â€Å"universally admired† â€Å"celestial beauty† that should be â€Å" a goddes among gods, adored and served by angels numberless†. The snake thus seems to contradict Gods Hierarchy of creation, The Great Chain of Being. According to him, Eve is higher than all the other creations of god, and should be sitting just next to god, just as the Holy Spirit and Jesus. In this passage the snake is the voices that question god authority and urges Eve to more freedom which appears, in the eyes of God as an Heresy. Hence a parallel can be drawn betwen the questioning of God’s Hierarchy by the snakes and the questioning of God’s Hierarchy by the new theories about cosmos expressed by scientists such as Copernic who were at the time seen as Heretics, devilish prophets and treated as such. Now that we have seen in which way the snakes represents a symbol of temptation embodying as well as the Biblical symbol of sin, the Renaissance questioning of God’s Hierarchy, we are going to see and analyse Milton’s use of Classic Mythology in this passage. First Milton’s use of Classic reference in his text can be interpreted as a perversions of those myths. In this passage he uses three myth to illustrate the action. He first refers to the tale of Hermione and Cadmus taken from Metamorphoses written by Ovid in AD 8. According to this classic of Latin literature they were both transformed into snakes in an act of vengence from the Gods. Then he refers to transformation of â€Å"Amonian Jove† and â€Å"Capitoline† into snakes to seduce and â€Å"mate† with Olympias and the mother of Scipio to give birth of two of the most famous and glorious war chief of both hellenistic and Roman era: Alexander the Great and Scipio Africanus. We can notice that Milton cleverly used myth in which the snakes as a role close to the one it has in the Bible. Concerning Ovid, the snakes is an object of temptation for which Hermione yields and is thus punished by being transformed in a snake. Concerning the myth of the birth of Alexender and Scipio, the snakes, just as in the Bible, is in fact a deity in disguise, seducing a woman. But if we look closely, these myth appeared to be perverted. In the classical myth, the deity or characters who are transformed into a snake are great and nobles characters: Cadmus is the founder of Thebes, and Amonian Jove and Capitoline are Jupiter. In paradise lost, the snake is Satan, the personification of rebellion against God’s will and of all the Human Sins. In this antithetic linking of diverse mythology, Milton wants to contrast Satan perfidy with classical characters nobility while underlining common themes. He thus appeals to a particular knowledge of his readers. On the other hand , the author’s use of classical mythology in this text can be seen as another influence of the metaphysical poetry on the Milton. Mixing diverse sources of knowledge to build his imagery is a technic used by metaphysiacal poets such as John Donne. But when others metaphysical poets mixed scientific, geographical, astronomical discoveries with theology and philospy, Milton just mixed Judeo-christian religious myths with classical myths. Even if this influence is small, it is sufficient to guess that Milton, contemporaneous with Donne, Herbert, Henry or Crashaw, has drawn some inspiration from the work of these poets. Moreover, Milton use of Classical mythology is representative of the Renaissance. Indeed, it clearly illustrates the resurgence of classical works during this era. Every arts had been affected by this tendency, in paintings, the classical style was a new fashion, the subjects taken from greek mythology began to be more and more numerous, Michaelangelo being a good exemple in both the sculpture and painting field. This resurgence has also an impact on architecture (with the building of houses, mansions, official buildign following the principle of the straight line) , on philosophy ( the â€Å"raison† and the re-reading of the platonician and socratic philosophy) as well as on literature. Thus Milton appears as an exemple of this new interest ofr the classical literrature, and htis work, Paradise lost, embodies this feature of the Renaissance. Finally, as we have seen, Milton uses his main influence, the Bible, to depicts the snake as a symbol of temptation and perfidy and uses his other influence, the classical mythology, in a perversion of the myths to develop his imagery. Furthermore, analysing this text, we can see that Milton was also influenced by the Metaphysical Poets in his uses of mixed sources as well as the use of the theme of the link between sensuality and religion. This book, which is now one of the most famous piece of english poetry, had a great influence on following artists such as Blake who considered Milton as the major English Poet and Edmond Burke and the Romantic theory, asserting â€Å"No person seems better to have understood the secret of heightening, or of setting terrible things, if I may use the expression, in their strongest light, by the force of a judicious obscurity than Milton. † , valuating his exploration of blank verse. Milton also had a great impact on the English language by creating neologism that are still used today such as pand?monium, enslaved, satanic and self-esteem.

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