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Medieval university
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This article is about Western European institutions. For other early centers of higher learning, see ancient higher-learning institutions. For an overview of medieval foundations, see List of medieval universities.
For a broader coverage of this topic, see History of universities.
Illustration from a 14th-century manuscript showing a meeting of doctors at the University of Paris.
A medieval university is a corporation organized during the Middle Ages for the purposes of higher learning.
The first Western European institutions generally considered universities were established in the Kingdom of Italy, then part of the Holy Roman Empire, the Kingdom of England, the Kingdom of France, the Kingdom of Spain, and the Kingdom of Portugal between the 11th and 15th centuries for the study of the Arts and the higher disciplines of Theology, Law, and Medicine.[1] These universities evolved from much older Christian cathedral schools and monastic schools, and it is difficult to define the exact date when they became true universities, though the lists of studia generalia for higher education in Europe held by the Vatican are a useful guide.
The word universitas originally applied only to the scholastic guilds—that is, the corporation of students and masters—within the studium, and it was always modified, as universitas magistrorum, universitas scholarium, or universitas magistrorum et scholarium. Eventually, however, probably in the late 14th century, the term began to appear by itself to exclusively mean a self-regulating community of teachers and scholars recognized and sanctioned by civil or ecclesiastical authority.[2]
From the early modern period onwards, this Western-style organizational form gradually spread from the medieval Latin west across the globe, eventually replacing all other higher-learning institutions and becoming the preeminent model for higher education everywhere.[3]
Contents
[hide]
1 Antecedents
2 Establishment
3 Characteristics
4 The students
4.1 Course of study
4.2 Legal status
5 See also
6 Notes
7 References
8 External links
Antecedents[edit]
A map of medieval universities.
The university is generally regarded as a formal institution that has its origin in the Medieval Christian setting.[4][5] Prior to the establishment of universities, European higher education took place for hundreds of years in Christian cathedral schools or monastic schools (scholae monasticae), in which monks and nuns taught classes. Evidence of these immediate forerunners of the later university at many places dates back to the 6th century AD.[6]
With the increasing growth and urbanization of European society during the 12th and 13th centuries, a demand grew for professional clergy. Before the 12th century, the intellectual life of Western Europe had been largely relegated to monasteries, which were mostly concerned with performing the liturgy and prayer; relatively few monasteries could boast true intellectuals. Following the Gregorian Reform's emphasis on canon law and the study of the sacraments, bishops formed cathedral schools to train the clergy in Canon law, but also in the more secular aspects of religious administration, including logic and disputation for use in preaching and theological discussion, and accounting to more effectively control finances. Pope Gregory VII was critical in promoting and regulating the concept of modern university as his 1079 Papal Decree ordered the regulated establishment of cathedral schools that transformed themselves into the first European universities.[7]
Learning became essential to advancing in the ecclesiastical hierarchy, and teachers also gained prestige. Demand quickly outstripped the capacity of cathedral schools, each of which was essentially run by one teacher. In addition, tensions rose between the students of cathedral schools and burghers in smaller towns. As a result, cathedral schools migrated to large cities, like Bologna, Rome and Paris.
Some scholars such as Syed Farid Alatas have noted some parallels between Madrasahs and early European colleges and have thus inferred that the first universities in Europe were influenced by the Madrasahs in Islamic Spain and the Emirate of Sicily.[8] Other scholars such as George Makdisi, Toby Huff and Norman Daniel, however, have questioned this, citing the lack of evidence for an actual transmission from the Islamic world to Christian Europe and highlighting the differences in the structure, methodologies, procedures, curricula and legal status of the "Islamic college" (madrasa) versus the European university.[9][10][11]
Establishment[edit]
Teaching at Paris, in a late 14th-century Grandes Chroniques de France: the tonsured students sit on the floor.
Hastings Rashdall set out the modern understanding[12] of the medieval origins of the universities, noting that the earliest universities emerged spontaneously as "a scholastic Guild, whether of Masters or Students... without any express authorisation of King, Pope, Prince or Prelate."[13]
Among the earliest universities of this type were the University of Bologna (1088), University of Paris (teach. mid-11th century, recogn. 1150), University of Oxford (teach. 1096, recogn. 1167), University of Modena (1175), University of Palencia (1208), University of Cambridge (1209), University of Salamanca (1218), University of Montpellier (1220), University of Padua (1222), University of Toulouse (1229), University of Orleans (1235), University of Siena (1240), University of Valladolid (1241) University of Northampton (1261), University of Coimbra (1288), University of Pisa (1343), Charles University in Prague (1348), Jagiellonian University (1364), University of Vienna (1365), Heidelberg University (1386) and the University of St Andrews (1413) begun as private corporations of teachers and their pupils.[14][15]
In many cases universities petitioned secular power for privileges and this became a model. Emperor Frederick I in Authentica Habita (1158) gave the first privileges to students in Bologna. Another step was when Pope Alexander III in 1179 "forbidding masters of the church schools to take fees for granting the license to teach (licentia docendi), and obliging them to give license to properly qualified teachers".[16] Hastings Rashdall considered that the integrity of a university was only preserved in such an internally regulated corporation, which protected the scholars from external intervention. This independently evolving organization was absent in the universities of southern Italy and Spain, which served the bureaucratic needs of monarchs—and were, according to Rashdall, their artificial creations.[17]
The University of Paris was formally recognized when Pope Gregory IX issued the bull Parens scientiarum (1231).[16] This was a revolutionary step: studium generale (university) and universitas (corporation of students or teachers) existed even before, but after the issuing of the bull, they attained autonomy. "[T]he papal bull of 1233, which stipulated that anyone admitted as a teacher in Toulouse had the right to teach everywhere without further examinations (ius ubique docendi), in time, transformed this privilege into the single most important defining characteristic of the university and made it the symbol of its institutional autonomy . . . By the year 1292, even the two oldest universities, Bologna and Paris, felt the need to seek similar bulls from Pope Nicholas IV."[16]
Mob Quad, in Merton College, Oxford. This group of buildings was constructed in three phases and concluded in c. 1378.
By the 13th century, almost half of the highest offices in the Church were occupied by degreed masters (abbots, archbishops, cardinals), and over one-third of the second-highest offices were occupied by masters. In addition, some of the greatest theologians of the High Middle Ages, Thomas Aquinas and Robert Grosseteste, were products of the medieval university.
The development of the medieval university coincided with the widespread reintroduction of Aristotle from Byzantine and Arab scholars. In fact, the European university put Aristotelian and other natural science texts at the center of its curriculum,[18] with the result that the "medieval university laid far greater emphasis on science than does its modern counterpart and descendent."[19]
Although it has been assumed that the universities went into decline during the Renaissance due to the scholastic and Aristotelian emphasis of its curriculum being less popular than the cultural studies of Renaissance humanism, Toby Huff has noted the continued importance of the European universities, with their focus on Aristotle and other scientific and philosophical texts into the early modern period, arguing that they played a crucial role in the Scientific Revolution of the 16th and 17th centuries. As he puts it "Copernicus, Galileo, Tycho Brahe, Kepler, and Newton were all extraordinary products of the apparently procrustean and allegedly Scholastic universities of Europe... Sociological and historical accounts of the role of the university as an institutional locus for science and as an incubator of scientific thought and arguments have been vastly understated."[20]
Characteristics[edit]
Diagrams, in a volume of treatises on natural science, philosophy, and mathematics. This 1300 manuscript is typical of the sort of book owned by medieval university students.
Initially medieval universities did not have physical facilities such as the campus of a modern university. Classes were taught wherever space was available, such as churches and homes. A university was not a physical space but a collection of individuals banded together as a universitas. Soon, however, universities began to rent, buy or construct buildings specifically for the purposes of teaching.[21]
Universities were generally structured along three types, depending on who paid the teachers. The first type was in Bologna, where students hired and paid for the teachers. The second type was in Paris, where teachers were paid by the church. Oxford and Cambridge were predominantly supported by the crown and the state, which helped them survive the Dissolution of the Monasteries in 1538 and the subsequent removal of all principal Catholic institutions in England. These structural differences created other characteristics. At the Bologna university the students ran everything—a fact that often put teachers under great pressure and disadvantage. In Paris, teachers ran the school; thus Paris became the premiere spot for teachers from all over Europe. Also, in Paris the main subject matter was theology, so control of the qualifications awarded was in the hands of an external authority - the Chancellor of the diocese. In Bologna, where students chose more secular studies, the main subject was law.
It was also characteristic of teachers and scholars to move around. Universities often competed to secure the best and most popular teachers, leading to the marketisation of teaching. Universities published their list of scholars to entice students to study at their institution. Students of Peter Abelard followed him to Melun, Corbeil, and Paris,[22] showing that popular teachers brought students with them.
The students[edit]
Students attended the medieval university at different ages—from 14 if they were attending Oxford or Paris to study the Arts, to their 30s if they were studying Law in Bologna. During this period of study, students often lived far from home and unsupervised, and as such developed a reputation, both among contemporary commentators and modern historians, for drunken debauchery. Students are frequently criticised in the Middle Ages for neglecting their studies for drinking, gambling and sleeping with prostitutes.[23]
Course of study[edit]
Universitas Istropolitana (A former university building in present-day Bratislava)
University studies took six years for a Master of Arts degree (a Bachelor of Arts degree was awarded after completing the third or fourth year). Studies for this were organized by the faculty of arts, where the seven liberal arts were taught: arithmetic, geometry, astronomy, music theory, grammar, logic, and rhetoric.[24][25] All instruction was given in Latin and students were expected to converse in that language.[26] The trivium comprised the three subjects that were taught first: grammar, logic, and rhetoric. These three subjects were the most important of the seven liberal arts for medieval students.[27] The curriculum came also to include the three Aristotelian philosophies: physics, metaphysics and moral philosophy.[27]
Much of medieval thought in philosophy and theology can be found in scholastic textual commentary because scholasticism was such a popular method of teaching. Aelius Donatus' Ars grammatica was the standard textbook for grammar; also studied were the works of Priscian and Graecismus by Eberhard of Béthune.[28] Cicero's works were used for the study of rhetoric.[27] Studied books on logic included Porphyry's introduction to Aristotelian logic, Gilbert de la Porrée's De sex principiis and Summulae Logicales by Petrus Hispanus (later Pope John XXI).[29] The standard work of astronomy was Tractatus de sphaera.[29]
Once a Master of Arts degree had been conferred, the student could leave the university or pursue further studies in one of the higher faculties, law, medicine, or theology, the last one being the most prestigious. A popular textbook for theological study was called the Sentences (Quattuor libri sententiarum) of Peter Lombard; theology students as well as masters were required to write extensive commentaries on this text as part of their curriculum.[citation needed] Studies in the higher faculties could take up to twelve years for a master's degree or doctorate (initially the two were synonymous), though again a bachelor's and a licentiate's degree could be awarded along the way.[30]
Courses were offered according to books, not by subject or theme. For example, a course might be on a book by Aristotle, or a book from the Bible. Courses were not elective: the course offerings were set, and everyone had to take the same courses. There were, however, occasional choices as to which teacher to use.[31]
A university class, Bologna (1350s).
Students often entered the University at fourteen to fifteen years of age, though many were older.[32] Classes usually started at 5:00 or 6:00 AM.
Legal status[edit]
Further information: Tenure (academic)
Students were afforded the legal protection of the clergy. In this way no one was allowed to physically harm them; they could only be tried for crimes in an ecclesiastical court, and were thus immune from any corporal punishment. This gave students free rein in urban environments to break secular laws with impunity, which led to many abuses: theft, rape and murder were not uncommon among students, who did not face serious consequences[33] and students were known to engage in drunkenness.
This led to uneasy tensions with secular authorities—the demarcation between town and gown. Masters and students would sometimes "strike" by leaving a city and not returning for years. This happened at the University of Paris strike of 1229 after a riot left a number of students dead. The University went on strike and they did not return for two years. As students had the legal status of clerics, which Canon Law prohibited for women, women were not admitted into universities.
Most universities in Europe were recognised by the Holy See as a Studium Generale, testified by a papal bull. Members of these institutions were encouraged to disseminate their knowledge across Europe, often lecturing at a different Studia Generales. Indeed, one of the privileges the papal bull confirmed was the right to confer the Ius ubique docendi, the right to teach everywhere.[34]
See also[edit]
List of oldest universities in continuous operation
Renaissance of the 12th century
Studium Generale
Ancient universities of Scotland
Town and gown
Nation (university)
University
Notes[edit]
Jump up ^ de Ridder-Symoens 1992, pp. 47–55
Jump up ^ Encyclopædia Britannica: History of Education. The development of the universities.
Jump up ^ Rüegg, Walter (ed.): Geschichte der Universität in Europa, 3 vols., C.H. Beck, München 1993, ISBN 3-406-36956-1
Jump up ^ Rüegg, Walter: "Foreword. The University as a European Institution", in: A History of the University in Europe. Vol. 1: Universities in the Middle Ages, Cambridge University Press, 1992, ISBN 0-521-36105-2, pp. XIX–XX
Jump up ^ Verger 1999
Jump up ^ Riché, Pierre (1978): "Education and Culture in the Barbarian West: From the Sixth through the Eighth Century", Columbia: University of South Carolina Press, ISBN 0-87249-376-8, pp. 126-7, 282-98
Jump up ^ Thomas Oestreich (1913). "Pope St. Gregory VII". In Herbermann, Charles. Catholic Encyclopedia. New York: Robert Appleton Company.
Jump up ^ Alatas, S. F. (2006), "From Jami`ah to University: Multiculturalism and Christian–Muslim Dialogue" (PDF), Current Sociology, 54 (1): 112–132 [123–4], doi:10.1177/0011392106058837, archived (PDF) from the original on 2017-09-23
Jump up ^ George Makdisi: "Madrasa and University in the Middle Ages", in: Studia Islamica, Vol. 32 (1970), S. 255-264 (264):
Thus the university, as a form of social organization, was peculiar to medieval Europe. Later, it was exported to all parts of the world, including the Muslim East; and it has remained with us down to the present day. But back in the middle ages, outside of Europe, there was nothing anything quite like it anywhere.
Jump up ^ The scholarship on these differences is summarized in Toby Huff, Rise of early modern science, 2nd ed. p. 149-159; p. 179-189.
Jump up ^ Norman Daniel: Review of "The Rise of Colleges. Institutions of Learning in Islam and the West by George Makdisi", Journal of the American Oriental Society, Vol. 104, No. 3 (Jul. - Sep., 1984), pp. 586-588 (587)
Jump up ^ Pryds, Darleen (2000), "Studia as Royal Offices: Mediterranean Universities of Medieval Europe", in Courtenay, William J.; Miethke, Jürgen; Priest, David B., Universities and Schooling in Medieval Society, Education and Society in the Middle Ages and Renaissance, 10, Leiden: Brill, p. 83, ISBN 90-04-11351-7, ISSN 0926-6070, In his magisterial work on European universities, Hastings Rashdall [considered that] the integrity of a university is preserved only when the institution evolved into an internally regulated corporation of scholars, be they students or masters.
Jump up ^ Rashdall, Hastings (1895), The Universities of Europe in the Middle Ages, 1, Oxford: Clarendon Press, pp. 17–18, retrieved February 2012, The University was originally a scholastic Guild, whether of Masters or Students. Such Guilds sprang into existence, like other Guilds, without any express authorisation of King, Pope, Prince, or Prelate. They were spontaneous products of the instinct of association that swept over the towns of Europe in the course of the eleventh and twelfth centuries. Check date values in: |accessdate= (help)
Jump up ^ "10 of the Oldest Universities in the World". Top Universities. 2016-09-16. Archived from the original on 2017-02-11. Retrieved 2017-05-30.
Jump up ^ Seelinger, Lani. "The 13 Oldest Universities In The World". Culture Trip. Archived from the original on 2017-10-01. Retrieved 2017-05-30.
^ Jump up to: a b c Kemal Gürüz, Quality Assurance in a Globalized Higher Education Environment: An Historical Perspective Archived 2008-02-16 at the Wayback Machine., Istanbul, 2007, p. 5
Jump up ^ Pryds, Darleen (2000), "Studia as Royal Offices: Mediterranean Universities of Medieval Europe", in Courtenay, William J.; Miethke, Jürgen; Priest, David B., Universities and Schooling in Medieval Society, Education and Society in the Middle Ages and Renaissance, 10, Leiden: Brill, pp. 83–99, ISBN 90-04-11351-7, ISSN 0926-6070
Jump up ^ Toby Huff, Rise of early modern science 2nd ed. p. 180-181
Jump up ^ Edward Grant, "Science in the Medieval University", in James M. Kittleson and Pamela J. Transue, ed., Rebirth, Reform and Resilience: Universities in Transition, 1300-1700, Columbus: Ohio State University Press, 1984, p. 68
Jump up ^ Toby Huff, Rise of Early Modern science, 2nd ed., p. 344.
Jump up ^ A. Giesysztor, Part II, Chapter 4, page 136: University Buildings, in A History of the University In Europe, Volume I: Universities in the Middle Ages, W. Ruegg (ed.), Cambridge University Press, 1992.
Jump up ^ James M. Kittleson, Rebirth, reform and resilience: Universities in transition 1300-1700, (Columbus, Ohio State University Press, 1984), p. 164.
Jump up ^ Skoda, Hannah (21 February 2013). "Medieval Violence: Physical Brutality in Northern France, 1270-1330". OUP Oxford. Archived from the original on 15 May 2018 – via Google Books.
Jump up ^ H. Rashdall, The Universities of Europe in the Middle Ages, 3 Volumes, F.M. Powicke, A.B. Emden (Eds. of 2nd Edition), Oxford University Press, 1936.
Jump up ^ G. Leff and J. North, Chapter 10: The Faculty of Arts, in A History of the University in Europe, Volume I: Universities in the Middle Ages, W. Ruegg (ed.), Cambridge University Press, 1992.
Jump up ^ Rait, R.S. 1912. Life in the Medieval University, p. 133
^ Jump up to: a b c Rait, R.S. 1912. Life in the Medieval University, p. 138
Jump up ^ Rait, R.S. 1912. Life in the Medieval University, pp. 138-139
^ Jump up to: a b Rait, R.S. 1912. Life in the Medieval University, p. 139
Jump up ^ O. Pedersen, The First Universities - Studium Generale and the Origins of University Education in Europe, Cambridge University Press, 1997
Jump up ^ Pedersen, o.c., Chapter 10: Curricula and intellectual trends.
Jump up ^ H. Rashdall, o.c., Volume 3, page 352.
Jump up ^ H. Rashdall, o.c., Volume 3, page 360.
Jump up ^ Rashdall, o.c., Chapter I, page 8.
References[edit]
Cobban, Alan B. English University Life in the Middle Ages Columbus: Ohio State University Press, 1999. ISBN 0-8142-0826-6
Ferruolo, Stephen: The Origins of the University: The Schools of Paris and their Critics, 1100-1215 Stanford: Stanford University Press, 1998. ISBN 0-8047-1266-2
Haskins, Charles Homer: The Rise of Universities. Ithaca, New York: Cornell University Press, 1972. ISBN 0-87968-379-1
Rashdall, Hastings, rev. by F. M. Powicke, and A. B. Emden: The Universities of Europe in the Middle Ages, 3 vols. Oxford: Clarendon Press, 1895; 1987, ISBN 0-19-821431-6
Ridder-Symoens, Hilde de (ed.): A History of the University in Europe. Vol. I: Universities in the Middle Ages, Cambridge University Press, 1992, ISBN 0-521-36105-2
Rait, Robert S.: Life in the Medieval University, Cambridge University Press, 1931, ISBN 0-527-73650-3
Pedersen, Olaf, The First Universities: Studium Generale and the Origins of University Education in Europe, Cambridge University Press, 1997.
Seybolt, Robert Francis, trans.: The Manuale Scholarium: An Original Account of Life in the Mediaeval University, Cambridge: Harvard University Press, 1921
Thorndike, Lynn, trans. and ed.: University Records and Life in the Middle Ages, New York: Columbia University Press, 1975, ISBN 0-393-09216-X
Verger, Jacques (1999), "Universität", Lexikon des Mittelalters, 8, Stuttgart: J.B. Metzler, cols:1249–1255
External links[edit]
The Shift of Medical Education into the Universities
The Educational Legacy of Mediaeval and Renaissance Traditions.
From Manuscript to Print: Evolution of the Mediaeval Book.
Life of the Students at Paris.
Mediaeval History: A Mediaeval Atlas
Cambridge, A Brief History: The Mediaeval University.
Mediaeval Science, the Church, and Universities
Quality Assurance In A Globalized Higher Education Environment: An Historical Perspective (DOC file)
The Rise of Universities (classic), Charles Homer Haskins, 1923
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Host (biology)
From Wikipedia, the free encyclopedia
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The black rat is a reservoir host for bubonic plague: the oriental rat fleas that infest these rats are vectors for the disease.
In biology and medicine, a host is an organism that harbours a parasitic, a mutualistic, or a commensalist guest (symbiont), the guest typically being provided with nourishment and shelter. Examples include animals playing host to parasitic worms (e.g. nematodes), cells harbouring pathogenic (disease-causing) viruses, a bean plant hosting mutualistic (helpful) nitrogen-fixing bacteria. More specifically in botany, a host plant supplies food resources to micropredators, which have an evolutionarily stable relationship with their hosts similar to ectoparasitism. The host range is the collection of hosts that an organism can use as a partner.
Contents
1 Symbiosis
2 Hosts to parasites
2.1 Definitive and secondary hosts
2.2 Paratenic hosts
2.3 Dead-end hosts
2.4 Plant hosts of micropredators
2.5 Host range
3 Non-parasitic associations
3.1 Mutualistic hosts
3.2 Hosts in cleaning symbiosis
3.3 Commensal hosts
4 History
5 See also
6 References
Symbiosis[edit]
Further information: Symbiosis
Symbiosis spans a wide variety of possible relationships between organisms, differing in their permanence and their effects on the two parties. If one of the partners in an association is much larger than the other, it is generally known as the host.[1] In parasitism, the parasite benefits at the host's expense.[2] In commensalism, the two live together without harming each other,[3] while in mutualism, both parties benefit.[4]
Most parasites are only parasitic for part of their life cycle. By comparing parasites with their closest free-living relatives, parasitism has been shown to have evolved on at least 233 separate occasions. Some organisms live in close association with a host and only become parasitic when environmental conditions deteriorate.[5]
A parasite may have a long term relationship with its host, as is the case with all endoparasites. The guest seeks out the host and obtains food or another service from it, but does not usually kill it.[6] In contrast, a parasitoid spends a large part of its life within or on a single host, ultimately causing the host's death, with some of the strategies involved verging on predation. Generally the host is kept alive until the parasitoid is fully grown and ready to pass on to its next life stage.[7] A guest's relationship with its host may be intermittent or temporary, perhaps associated with multiple hosts, making the relationship equivalent to the herbivory of a wild-living animal. Another possibility is that the host-guest relationship may have no permanent physical contact, as in the brood parasitism of the cuckoo.[6]
Hosts to parasites[edit]
Micropredator, parasite, parasitoid, and predator strategies compared. Their interactions with their hosts form a continuum. Micropredation and parasitoidism are now considered to be evolutionary strategies within parasitism.[2]
See also: Parasitism
Parasites follow a wide variety of evolutionary strategies, placing their hosts in an equally wide range of relationships.[2] Parasitism implies host–parasite coevolution, including the maintenance of gene polymorphisms in the host, where there is a trade-off between the advantage of resistance to a parasite and a cost such as disease caused by the gene.[8]
Definitive and secondary hosts[edit]
There are several kinds of host from a parasite's point of view.
A definitive or primary host is one in which a parasite reaches maturity and reproduces sexually. An intermediate host is one in which the parasite does not do this. A virus is an obligate parasite, acting as a living thing only to the extent that when it is in a host cell, the machinery of that cell makes copies of the virus.[9] A reservoir host can harbour a pathogen indefinitely with no ill effects, with important implications for disease control. A single reservoir host may be reinfected several times.[10] A host of predilection is the one preferred by a parasite.[11] An amplifying host is one in which the level of pathogen can become high enough that a vector such as a mosquito that feeds on it will probably become infectious.[12]
A secondary or intermediate host harbors a parasite only for a short transition period, during which (usually) some developmental stage is completed. For trypanosomes, the cause of sleeping sickness, strictly, humans are the secondary host, while the tsetse fly is the primary host, given that it has been shown that reproduction occurs in the insect.[13] Tapeworms and other parasitic flatworms have complex lifecycles, in which specific developmental stages are completed in a sequence of several different hosts.[14]
In trichinosis, the same host (such as humans or pigs) is both intermediate and definitive.
It is not always easy or even possible to identify which host is definitive and which secondary. As the life cycles of many parasites are not well understood, sometimes the subjectively more important organism is arbitrarily labelled as definitive, and this designation may continue even after it is found to be incorrect. For example, sludge worms are sometimes considered "intermediate hosts" for salmonid whirling disease, even though the myxosporean parasite reproduces sexually inside them.[15] In trichinosis, a disease caused by roundworms, the host has reproductive adults in its digestive tract and immature juveniles in its muscles, and is therefore both an intermediate and a definitive host.[16]
Paratenic hosts[edit]
A paratenic host is similar to an intermediate host, except that it is not needed for the parasite's development cycle to progress. Paratenic hosts serve as "dumps" for non-mature stages of a parasite in which they can accumulate in high numbers. The trematode Alaria americana may serve as an example: the so-called mesocercarial stages of this parasite reside in tadpoles, which are rarely eaten by the definitive canine host. The tadpoles are more frequently preyed on by snakes, in which the mesocercariae may not undergo further development. However, the parasites may accumulate in the snake paratenic host and infect the definitive host once the snake is consumed by a canid.[17] The nematode Skrjabingylus nasicola is another example, with slugs as the intermediate hosts, shrews and rodents as the paratenic hosts, and mustelids as the definitive hosts.[18]
Dead-end hosts[edit]
A dead-end or incidental host is an intermediate host that generally does not allow transmission to the definitive host, thereby preventing the parasite from completing its development. For example, humans and horses are dead-end hosts for West Nile virus, whose life-cycle is normally between culicine mosquitoes and birds.[12] People and horses can become infected, but the level of virus in their blood does not become high enough to pass on the infection to mosquitoes that bite them.[12]
Plant hosts of micropredators[edit]
Buff ermine moth caterpillar, a polyphagous micropredator
Micropredation is an evolutionarily stable strategy within parasitism, in which a small predator lives parasitically on a much larger host plant, eating parts of it.[2]
The range of plants on which a herbivorous insect feeds is known as its host range. This can be wide or narrow, but it never includes all plants. A small number of insects are monophagous, feeding on a single plant. The silkworm larva is one of these, with mulberry leaves being the only food consumed. More often, an insect with a limited host range is oligophagous, being restricted to a few closely related species, usually in the same plant family.[19] The diamondback moth is an example of this, feeding exclusively on brassicas,[20] and the larva of the potato tuber moth feeds on potatoes, tomatoes and tobacco, all members of the same plant family, Solanaceae.[21] Herbivorous insects with a wide range of hosts in various different plant families are known as polyphagous. One example is the buff ermine moth whose larvae feed on alder, mint, plantain, oak, rhubarb, currant, blackberry, dock, ragwort, nettle and honeysuckle.[22]
Influenza virus can change by genetic reassortment as it travels between different hosts in its range.
Plants often produce toxic or unpalatable secondary metabolites to deter herbivores from feeding on them. Monophagous insects have developed specific adaptations to overcome those in their specialist hosts, giving them an advantage over polyphagous species. However, this puts them at greater risk of extinction if their chosen hosts suffer setbacks. Monophagous species are able to feed on the tender young foliage with high concentrations of damaging chemicals on which polyphagous species cannot feed, having to make do with older leaves. There is a trade off between offspring quality and quantity; the specialist maximises the chances of its young thriving by paying great attention to the choice of host, while the generalist produces larger numbers of eggs in sub-optimal conditions.[23]
Some insect micropredators migrate regularly from one host to another. The hawthorn-carrot aphid overwinters on its primary host, a hawthorn tree, and migrates during the summer to its secondary host, a plant in the carrot family.[24]
Host range[edit]
The host range is the set of hosts that a parasite can use as a partner. In the case of human parasites, the host range influences the epidemiology of the parasitism or disease. For instance, the production of antigenic shifts in Influenza A virus can result from pigs being infected with the virus from several different hosts (such as human and bird). This co-infection provides an opportunity for mixing of the viral genes between existing strains, thereby producing a new viral strain. An influenza vaccine produced against an existing viral strain might not be effective against this new strain, which then requires a new influenza vaccine to be prepared for the protection of the human population.[25]
Non-parasitic associations[edit]
Mutualistic hosts[edit]
Further information: Mutualism (biology)
Mycorrhiza, a mutualistic interaction between a plant's roots and a fungus
Some hosts participate in fully mutualistic interactions with both organisms being completely dependent on the other. For example, termites are hosts to the protozoa that live in their gut and which digest cellulose,[26] and the human gut flora is essential for efficient digestion.[27] Many corals and other marine invertebrates house zooxanthellae, single-celled algae, in their tissues. The host provides a protected environment in a well-lit position for the algae, while benefiting itself from the nutrients produced by photosynthesis which supplement its diet.[28] Lamellibrachia luymesi, a deep sea giant tubeworm, has an obligate mutualistic association with internal, sulfide-oxidizing, bacterial symbionts. The tubeworm extracts the chemicals that the bacteria need from the sediment, and the bacteria supply the tubeworm, which has no mouth, with nutrients.[29] Some hermit crabs place pieces of sponge on the shell in which they are living. These grow over and eventually dissolve away the mollusc shell; the crab may not ever need to replace its abode again and is well-camouflaged by the overgrowth of sponge.[30]
An important hosting relationship is mycorrhiza, a symbiotic association between a fungus and the roots of a vascular host plant. The fungus receives carbohydrates, the products of photosynthesis, while the plant receives phosphates and nitrogenous compounds acquired by the fungus from the soil. Over 95% of plant families have been shown to have mycorrhizal associations.[31] Another such relationship is between leguminous plants and certain nitrogen-fixing bacteria called rhizobia that form nodules on the roots of the plant. The host supplies the bacteria with the energy needed for nitrogen fixation and the bacteria provide much of the nitrogen needed by the host. Such crops as beans, peas, chickpeas and alfalfa are able to fix nitrogen in this way,[32] and mixing clover with grasses increases the yield of pastures.[33]
Cleaning symbiosis: a Hawaiian cleaner wrasse with its client, a yellowtail wrasse
Hosts in cleaning symbiosis[edit]
Further information: Cleaning symbiosis and Cleaner fish
Hosts of many species are involved in cleaning symbiosis, both in the sea and on land, making use of smaller animals to clean them of parasites. Cleaners include fish, shrimps and birds; hosts or clients include a much wider range of fish, marine reptiles including turtles and iguanas, octopus, whales, and terrestrial mammals.[4] The host appears to benefit from the interaction, but biologists have disputed whether this is a truly mutualistic relationship or something closer to parasitism by the cleaner.[34][35]
Nurse shark playing host to commensal remoras, which gain a free ride and which may serve as cleaners
Commensal hosts[edit]
Further information: Commensalism and Phoresis (biology)
Remoras (also called suckerfish) can swim freely but have evolved suckers that enable them to adhere to smooth surfaces, gaining a free ride (phoresis), and they spend most of their lives clinging to a host animal such as a whale, turtle or shark.[3] However, the relationship may be mutualistic, as remoras, though not generally considered to be cleaner fish, often consume parasitic copepods: for example, these are found in the stomach contents of 70% of the common remora.[36] Many molluscs, barnacles and polychaete worms attach themselves to the carapace of the Atlantic horseshoe crab; for some this is a convenient arrangement, but for others it is an obligate form of commensalism and they live nowhere else.[26]
History[edit]
Further information: Parasitism
The first host to be noticed in ancient times was human: human parasites such as hookworm are recorded from ancient Egypt from 3000 BC onwards, while in ancient Greece, the Hippocratic Corpus describes human bladder worm.[37] The medieval Persian physician Avicenna recorded human and animal parasites including roundworms, threadworms, the Guinea worm and tapeworms.[37] In Early Modern times, Francesco Redi recorded animal parasites, while the microscopist Antonie van Leeuwenhoek observed and illustrated the protozoan Giardia lamblia from "his own loose stools".[37]
Hosts to mutualistic symbionts were recognised more recently, when in 1877 Albert Bernhard Frank described the mutualistic relationship between a fungus and an alga in lichens.[38]
See also[edit]
PHI-base (Pathogen-Host Interaction database)
Host cell protein
References[edit]
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^ Jump up to: a b Jackson, John (30 November 2012). "How does the Remora develop its sucker?". National History Museum. Retrieved 19 October 2017.
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Jump up ^ "Potato tuberworm: Phthorimaea operculella". Featured Creatures. IFAS. Retrieved 18 October 2017.
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Jump up ^ Cressey, R.; Lachner, E. (1970). "The parasitic copepod diet and life history of diskfishes (Echeneidae)". Copeia. 1970 (2): 310–318. doi:10.2307/1441652. JSTOR 1441652.
^ Jump up to: a b c Cox, Francis E. G. (June 2004). "History of human parasitic diseases". Infectious Disease Clinics of North America. 18 (2): 173–174. doi:10.1016/j.idc.2004.01.001. PMID 15145374.
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Hoste – Catalan
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Hostitel – Czech
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Hosto – Ido
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Inang – Indonesian
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Selil ot – Haitian Creole
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