Está en la página 1de 7

Eel life history

viviparus was the “mother of eels” (the translation of the
German name "Aalmutter").

1 Past studies of eels
In 1777, the Italian Carlo Mondini located an eel’s gonads
and demonstrated that eels are a kind of fish. In 1876,
as a young student in Austria, Sigmund Freud dissected
hundreds of eels in search of the male sex organs. He had
to concede failure in his first major published research
paper, and turned to other issues in frustration.[1][2][3][4]
Larval eels — transparent, leaflike two-inch (five-cm)
creatures of the open ocean — were not generally recognized as such until 1893; instead, they were thought
to be a separate species, Leptocephalus brevirostris (from
the Greek leptocephalus meaning “thin- or flat-head”). In
1886, however, the French zoologist Yves Delage discovered the truth when he kept leptocephali alive in a laboratory tank in Roscoff until they matured into eels, and in
1896 Italian zoologist Giovanni Battista Grassi confirmed
the finding when he observed the transformation of a
Leptocephalus into a round glass eel in the Mediterranean
Sea. (He also observed that salt water was necessary to
support the maturation process.) Although the connection between larval eels and adult eels is now well understood, the name leptocephalus is still used for larval eel.

Distribution and size of leptocephali larvae of the European eel,
Anguilla anguilla

2 Search for the spawning grounds

Distribution and size of leptocephali larvae of the American eel,
Anguilla rostrata

The eel is a long, thin bony fish of the order
Anguilliformes. Because fishermen never caught anything they recognized as young eels, the life cycle of the
eel was a mystery for a very long period of scientific
history. Although more than 6500 publications mention
eels, much of their life history remains an enigma.
The European eel (Anguilla anguilla) was historically
the one most familiar to Western scientists, beginning
with Aristotle, who wrote the earliest known inquiry
into the natural history of eels. He speculated that they Leptocephalus larva of an ocean eel
were born of “earth worms”, which he believed were
formed of mud, growing from the “guts of wet soil”
rather than through sexual reproduction. Many centuries 2.1 European eel
passed before scientists were able to demonstrate that
such spontaneous generation does not occur in nature.
The Danish professor Johannes Schmidt, beginning in
Other early scientists believed that the eelpout Zoarces 1904, led a series of expeditions into the Mediterranean
1

so they have to rely on stored Sea. The elvers grow larger and are referred to as yellow eels. turn into elvers (young eels). and hypothesized that they all must have de. a travel speed in the ocean of 15 km per day from completion of leptocephalus metamorphosis until can be assumed. In a 1922 expedition. he went. He also ob. As the glass eels enter fresh waters. in England. the skin is still transparent and the red gills and the heart are visible. and likely pass through a stage similar to the anguillid glass eels. he sailed as far as the Sargasso making feeding impossible. and then enter estuaries as glass eels and swim upstream to live in fresh water during their juvenile growth phase. Eels in this so-called “recruitment” developmen632-06389-0). length about 8 cm 2 SEARCH FOR THE SPAWNING GROUNDS tion of glass eels takes place (for deli food and stocking) is Epney. across the Atlantic Ocean. they are called yellow eels because of their golden pigmentation. length about 25 cm . first down parency of their bodies. According stage. Haiti. He noted that all the leptocephali he found were very similar. they have matured and grown to a length of 60 to 80 cm. These freshwater eels spawn in the ocean. thus colonising the continent. which are the juvenile stage of eels before their reproductive maturation begins. (Glass eels are a food item in Spain.2 Glass eels at the transition between ocean and fresh water. have been the peditions were largely financed by the Carlsberg Foundabasis of traditional fisheries with characteristic trapnets. and the sides of their bodies turn silvery. Marine eels of the order Anguilliformes also have a leptocephalus stage.energy alone.700 mi) open scended from a common ancestor species. and insects. they develop pigmentation. The exthrough the Baltic Sea in the Danish belts. eye pigments change for optimal vision in dim blue clear Although Schmidt did not directly observe eel spawning. At this stage.[6] The external features undergo other dramatic changes. In fresh water. and Puerto Rico remains unknown.ocean journey back to their spawning grounds north of served that the farther out to sea in the Atlantic Ocean the Antilles. Glass eels typically refers to the Baltic. he was able to decreate a countershading pattern which makes them difduce the following about the life history of the eel. and grow to 75–90 mm within The German fisheries biologist Friedrich Wilhelm Tesch. south of Bermuda. based ficult to see by predators during their long. their gut dissolves. These migrating eels are typically called “silThe larvae of European eels travel with the Gulf Stream ver eels” or “big eyes”. as well: the eyes start to enlarge. In July. overcoming various natural challenges — sometimes by piling up their bodies by the tens of thousands to climb over obstacles — and they reach even the smallest of creeks. They can propel themselves over wet grass and dig through wet sand to reach upstream headwaters and ponds. some individuals mature and migrate back towards the sea. in about 165 to 175 when leaving from the English ChanOne well-known location in which the large-scale collecJuvenile eels. they start to become pigmented and are typically referred to as elvers. but they are rarely seen in the ocean. By the time they leave the continent. open-ocean on the size distribution of the leptocephali he collected: migration. but finally the transmitter signals were lost at the between the leptocephalus stage and the juvenile (elver) continental shelf when the batteries ran out. Eel migration out of their freshwater growth habitats from various parts of Europe. they migrate up rivers and streams. crossing wet grasslands at night to reach rivers that lead to the sea.) Once they recruit to coastal areas.000 km (3. and feed on creatures such as small crustaceans.How the adults make the 6. the smaller the leptocephali he caught were. the larvae that had ever been seen. worms. After 10 to 14 years. Glass eels are defined as “all developmental stages to Schmidt. then along the coasts of Norway and Engan intermediary stage in the eel’s complex life history land. on the Severn. ocean light. where he caught the smallest eel. or Sea and the North Atlantic to investigate eels. one to three years. tion. before they reach the coasts of Euan eel expert and author of the book The Eel (ISBN 0rope.[5] The term typically refers to a transdays to reach the Sargasso Sea from around Scotland and parent glass eel of the family Anguillidae. to or even find ready-to-spawn adult eels. conducted many expeditions with hightal stage are known as glass eels because of the transtech instrumentation to follow eel migration. so a silver eel would need 140 to 150 full pigmentation”.

depth.000 a kilogram at times when $1. After spawning in the Sargasso Sea and moving to the west. 2. bicolor bicolor. However. but beginning in the mid1980s. First it was believed European and American eels were the same species due to their similar appearance and behavior.[7] Knowledge of what happens to individual silver eels after they leave the continental shelf is based solely on the study of three eels found in the stomachs of deepsea fishes and whales — caught off the coasts of Ireland and the Azores — and on laboratory research into the physiology of eels. A. but this has not been confirmed. marmorata in the West Mariana Ridge.000 would buy the same amount of American glass eels at their catching sites. with A. Japanese scientists discovered and caught matured adult eels of A. but they differ in chromosome count and various molecular genetic markers. European demand for eels could not be met for the first time ever. float up toward the surface.[11] Such a kilogram. dealers from China alone placed advance orders for more than 250. and broadcast their position. and with some spawning by the American eel possibly even occurring off the Yucatán Peninsula off the Gulf of Mexico.[9] Southern Africa’s four species of freshwater eels (A. Data from Maine and other North American coasts showed similar declines. even today.000 kg. anguilla. rostrata 103 to 110. up in some of the Southern African river systems and then [10] Tesch — like Schmidt — kept trying to persuade spon.3 Japanese eel The spawning area of the Japanese eel. A.100 per kg. The spawning grounds for the two species are in an overlapping area of the southern Sargasso Sea. bengalensis labiata. In New Jersey. although not as drastic. The traditional European stocking programs could not compete any longer: each week. Even before the 1997 generation hit the coasts of Europe. glass eel arrival in the spring dropped drastically — in Germany to 10% and in France to 14% of their previous levels — from even conservative estimates. and A. only preliminary experiments along these lines have ever been performed. rostrata apparently being more westward than A. Furthermore. and dealers from Asia bought all they could.back again to the ocean off Madagascar. some bidding more than $1. marmorata) have a very interesting migratory pattern. consisting of 5000 glass eels. Anguilla japonica. A. In 1997. and temperature to satellite receivers. Glass eel on the online in situ microscope at the LEO project 2. sors to provide more funding for expeditions. His proposal was to release 50 silver eels from Danish waters. It takes them on a long journey from their spawning grounds in the Indian Ocean north of Madagascar to high Glass eel No one yet knows the reasons. Migration was mapped in 2016. Asian elvers have sold in Hong Kong for as much as $5.000 after they leave an Asian fish farm.000 and as much as $150. the price for a kilogram of glass eel went up another US$30. anguilla counting 110 to 119 and A. mossambica. with transmitters that would detach from the eels each 3 Decline of the glass eels second day.2 American eel Another Atlantic eel species is known: the American eel.000 to $6. He also suggested that countries on the western side of the Atlantic could perform a similar release experiment at the same time. and in the number of vertebrae. has also been precisely located to be to the west of the Suruga seamount[8] and their leptocephali are then transported to the west to East Asia by the North Equatorial Current. Anguilla rostrata. japonica and A.3 nel. may bring at least $60. the leptocephali of the American eel exit the Gulf Stream earlier than the European eel and begin migrating into the estuaries along the east coast of North America between February and late April at an age around one year and a length around 60 mm. in June and August 2008. over 2000 licences for glass eel catch were issued and reports of 38 kg per .

doi:10. Karen (1993) Think of an Eel. [3] FH. [7] “Empirical observations of the spawning migration of European eels: The long and dangerous road to the Sargasso Sea”. 4 Threats to eels Strong concerns exist that the European eel population might be devastated by a new threat: Anguillicola crassus. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. The demand for adult eels has continued to grow. Retrieved 2012-01-04. [4] “Sigmund Freud und der Aal” (in German).. (2009). Mochioka. S. Jutta Birmele. April ladders have been constructed in North America and Eu2004 rope to help the fish bypass obstructions. more than 100 million kg were consumed in 1996. In New Jersey. 1996 (LEO) site.. In Europe. 2004-10-20. NYU Press.org. Berlinonline.[12] As open ocean voyagers. 2002. 1995. “Der Aal im Nationalsozialismus”. The eel. Mathematisch-Naturwissenschaftliche Classe. more funding becomes available. M.de. "Beobachtungen über Gestaltung und feineren Bau der als Hoden beschriebenen Lappenorgane des Aals" ["Observations on the configuration and finer structure of the lobed organs in eels described as testes"]. New high-tech eel aquaculture plants are appearing in Asia. p. Web. Published by Blackwell Science. 75: 257– 259. A.. Wno. Since 1995. 1–22. . Valerie D Greenberg (eds. it will be possible to log in to the system via a Longterm Ecological Observatory [12] Wuertz et al. worldwide interest in American eels has increased dramatically. “Discovery of mature freshwater eels in the open ocean”.de. 7 Further reading 5 See also • Eel ladder • Fish migration • Tesch.). 419 (1877). Sigmund.408pp.new tools in environmental education. pp. As soon as nal on December 24. 408 pages [6] Piper. Tsukamoto. [8] Tsukamoto K. and each female can produce 2 to 10 million eggs. 2003.. Archived from the original on December 17. H. but in Japan alone. Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. turkurier. 2011. Nature 439(7079): 929 [9] Chow.Children’s picture book describing the life cycle of the eel. Retrieved 2013-07-16. Ross (2007). 75. Blackwell Science.Berliner Zeitung” (in German).org. 6 References [1] Freud. Since the 1970s. Kaji. [2] “Was dachten Nazis über den Aal? | Archiv . Retrieved 2012-01-04. Retrieved 11 October 2016. an ongoing project monitors the glass eel [11] “Demand for Baby Eels Brings High Prices and Limits”. This parasite from East Asia (the original host is A. Fisheries Science. japonica) appeared in European eel populations in the early 1980s. Vol. Reading Freud’s Reading. Traditional eel aquaculture operations rely on wild-caught elvers..archive. Jay Geller. Kurogi. • Wallace. an increasing number of eel [10] Jim Cambray: African freshwater eels . Oceanic biology: spawning of eels near a seamount. japonica. Freud’s study was in response to Szymon Syrski's book Ueber die Reproductions-Organe der Aale (1874). with detrimental effects on the native Japanese eel. Retrieved 2013-07-16. see Ursula Reidel-Schrewe “Freud’s Début in the Sciences” in: Sander L. 25 million kg are consumed each year. As the European eels become less available. Recently. Oxford (UK).1007/s12562-008-0017-5 Because the eels are catadromous (living in fresh water but spawning in the sea). it also appeared in the United States (Texas and South Carolina). most likely due to uncontrolled aquaculture eel shipments. (23 February 2006). Website: Science in Africa. K.4 7 FURTHER READING night and fisherman have been made. 1 . Walker Books (UK) .. 2000-12-03. this parasite was shown to inhibit the function of the swimbladder as a hydrostatic organ. F-W (2003) The eel. although the average catch is closer to 1 kg. eels need the carrying capacity of the swimbladder (which makes up 3–6% of the eel’s body weight) to cross the ocean on stored energy alone. Germany imported more than $50 million worth of eels in 2002. Archived from the origimigration with an online in situ microscope. S. a foreign parasitic nematode. Greenwood Press. dams and other river obstructions can block their ability to reach inland feeding grounds. Gilman. as of 2003. eel populations are already from 30% to 100% infected with the nematode. but experimental hormone treatments in Japan have led to artificially spawned eels. In Europe.W. Kul- [5] Tesh F. third edition. Eggs from these treated eels have a diameter of about 1 mm. N. Okazaki.

ix + 386. Findley. A field guide to Atlantic coast fishes of North America. p. A field guide to freshwater fishes of North America north of Mexico. et al. Claro (ed. Patrimoines Naturels. James R. Washington. In W. F. Ecology of the Marine Fishes of Cuba.R. (1995). In W. Bond.C. and John A. 65 s. 3. Brooker.M. Wolstehnolmen and M. USA. and James D. Houghton Mifflin Company. 195 p. • Murdy. Burr (1991). no. 162-176. 9. 2905. U.) Ecología de . Fishes of Chesapeake Bay. R. U. • Food and Agriculture Organization (1992). Fischer (ed. Birdsong. Sounds of Western North Atlantic fishes. Croix. Ser. Courtney. Smithsonian Institution Press. James R. Greenwood Press.S.Y. the Bahamas. • Kenny. DC. World record game fishes.G. A List of Common and Scientific Names of Fishes from the United States and Canada. Rodolfo. Edwin J.L. Reeve M.. National Audubon Society field guide to tropical marine fishes of the Caribbean. volume 1. 70:(105):647 p. B. An annotated list of the fishes of St. Amer. Aquaculture production statistics 1986–1995. • Nielsen. Yntema. American Fisheries Society. Chapter 2: The Marine Ichthyofauna of Cuba. Ray S. London. Vol. 2001. USA.S. Bond. Nuuk. Florida. FAO Stat.. USA. and W. xi + 324. R. and I. A List of Common and Scientific Names of Fishes from the United States and Canada. et al. Bethesda. (1997). • Claro. Joseph. 1980. Rev. American Fisheries Society. vol 13. 1980. eds.W. USA. Musick 1997. 212-230. and Bermuda. • Greenfield. Bethesda. Fisk i grønlandske farvande. University Press of Florida. and E. Clavijo (1975). Soc. Ser. Robert N. • Smith. Brooker. Fish. Rodolfo. Gilbert. Lachner. Ernest A. 2004.M. 354 p. Maracas. Ray (1986). and B. 29. Lloyd T. Jr. 432 p. with special reference to those kept in the New York Aquarium. Parenti / Claro. FAO Fish. [pag. International Game Fish Association.R.. no. In G. St. Longevity of fishes in captivity. Richard C.5 • Wenner. • Lim. (eds. • Page.. and Noël. California. Jr. Atlas des poissons et des crustacés d'eau douce de la Martinique. • Robins. J. • Piper. FAO yearbook 1990. Fishery statistics. A reference file of biological underwater sounds. Mowbray (1970). • Nigrelli. Houghton Mifflin Company. • International Game Fish Association (1991).F. • Ogden. 98 p. P.. Smithsonian Institution Press. Bertelsen (1992). Fishes of the continental waters of Belize. Spec. Fourth Edition. L. ISBN 0-940228-47-5. C. and L. USA.. Stauffer. Bailey. Atuakkiorfik. and Lynne R. 815. The Johns Hopkins Press. and Mexico. ISBN 1-56098-985-8. 21-57. California Academy of Sciences.E. 174. and G. Maryland.A. USA.M. Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. ed. D. Catches and landings. no. C. Churchill. Characterísticas generales de la ictiofauna. • Eschmeyer. • Robins. Maryland. Lea. Exotic fishes in Puerto Rico. Virgin Islands. (1959). p. Boston. Meunier. (1984). Joseph S.W and J. R. p. Kenny.J. Florida.. ISBN 1-888569-61-1. • Robins. American Fisheries Society Special Publication. Baltimore. Carl E.. FAO Fish. (2002). Common and scientific names of fishes from the United States. San Francisco. Ernest A. USA. American Fisheries Society Special Publication. Fourth Edition. Parenti. Williams.A. USA. Keith.S. Catalog of Fishes. 12. (1994). 1998. Paris: MNHN. Trinidad and Tobago.C.) Distribution.) FAO species identification sheets for fishery purposes. M.S. • Fish. Special Publication of the Center for Biodiversity Research and Information. • Nelson. no. D. Carter R. (1978). Héctor Espinosa-Pérez. J. (38). (1987). Crossman. P. Sixth Edition.P. American Fisheries Society Special Publication. Rome.) Ciba Foundation Colloquium on Ageing: the life span of animals. West Atlantic (Fishing Area 31). eds. New York.R.H. American Fisheries Society. and J. • Erdman. • Jessop. ISBN 1-56098-638-7. Maryland. 720 p. R (2007).A. the Gulf of Mexico. Alfred A. Bethesda. Canada. Inc. Lachner. Views from the bridge: a memoir on the freshwater fishes of Trinidad. 311 p. 174. Anguillidae. biology and management of exotic fishes. DC. Kenyon C.. Florida. 51: 120p.E Thomerson (1997). 116:161170. Knopf. Lindeman. Boston. Carl E. J. O'Connor (eds.. FAO. Reeve M.]. P. Baltimore. J. • Claro. 12. Publ. 5. Bailey. In R. Circ. Washington. var. C. 1. Julian S. No. Trans. William N. Migrating American eels in Nova Scotia. 55-70. Johns Hopkins University Press. Edward O. • FAO (1997). Richard C.

K Glass Eels — a large commercial firm’s website. • Böhlke..S.S. Starnes 2003. B. • Bussing. C. W. 7(1):17-31. R. J. R. 8 External links • The Maine Eel and Elver Fishery Maine Department of Marine Resources • Fishbase entry for Anguilla anguilla • Fishbase entry for Anguilla rostrata • ICES report about eel stock collapse • U. Gardner. L. the U. A.M. Checklist of Vertebrates of the United States. Territories. San José Costa Rica: Editorial de la Universidad de Costa Rica. 2nd Ed. (1998). J. Instituto de Oceanología Academia de Ciencias de Cuba and Centro de Investigaciones de Quintana Roo. Fishes of the Bahamas and adjacent tropical waters. University of Texas Press. W. 2nd edition.S.A. Chaplin (1993). McDiarmid. 468 p.6 8 los peces marinos de Cuba.G. and Canada. C. and W. A list of Barbadian fishes.C. with history and fact pages • Projekt eelBASE EXTERNAL LINKS . Austin. • Butsch. (1939). • Banks.E. Peces de las aguas continentales de Costa Rica [Freshwater fishes of Costa Rica]. and C. R.H.

jpg License: Public domain Contributors: ? Original artist: ? • File:Conger_conger_Gervais. donated to the Wikimedia Foundation Original artist: Pearson Scott Foresman • File:Pacific_oysters_01.org/wikipedia/commons/9/98/Glasseelkils. and licenses 9. Stan Shebs. Tcr25. Piano non troppo.org/wikipedia/commons/8/80/LeptocephalusConger.18.wikimedia.org/wikipedia/commons/7/76/Conger_conger_Gervais.wikimedia.jpg Source: https://upload.0 Contributors: ? Original artist: Uwe Kils • File:Atlantic_cod. contributors. Fred Condo. SmackBot. Athaenara.0 Contributors: Transferred from en. Kils.jpg License: CC BY 2. Original artist: Kils at English Wikipedia • File:Glasseelskils.1 Text • Eel life history Source: https://en. Nilmerg.0 Contributors: Transferred from en.jpg Source: https://upload. Lightbot.jpg License: CC-BY-SA-3. Tannin. LucasVB. Scimitar. Edgar181. Bender235. Heidijane. Based on Image:Question book. CapnPrep. Calabe1992. Thehelpfulbot.0 Contributors: Transferred from en. Hayleybumm. MJMiller33.org/wikipedia/commons/6/6b/Anguillamuk. Hebrides.org/wikipedia/en/9/99/Question_book-new. Gadfium. Lotje. Mieliestronk.wikimedia. JohnOwens.wikipedia. Dmmaus. mod. Original artist: Kils at English Wikipedia • File:Lobster_(PSF). Ahoerstemeier. Fwb44.wikimedia.0 . Fishmac13. Fredrik.wikimedia.jpg Source: https://upload. Jeltz. Viriditas.0 Contributors: Created from scratch in Adobe Illustrator. Jeremy89442. Yobot. Thincat. VoABot II.wikimedia.jpg License: CC-BY-SA-3.wikimedia. Epipelagic.0 Contributors: Transferred from en. Olivier. Reatlas.jpg Source: https://upload. Shoyochow~enwiki. Pinethicket.0 Contributors: ? Original artist: ? • File:LeptocephalusConger. Reaper Eternal. Gdrbot. Guanabot.5 Contributors: Wikipedia Commons Original artist: David Monniaux. Cyberbot II. Rondonia9.org/wikipedia/commons/a/a3/Atlantic_cod. Robbot. RudolfRed.org/wikipedia/commons/a/ad/Rostratamuk. Wiae. ClueBot NG.wikipedia to Commons.wikipedia English Wikipedia Original artist: Original uploader was Kils at en. ChrisGualtieri. Hans~enwiki. Caroline1981.org/wikipedia/commons/3/3d/Pacific_oysters_01. VolkovBot.org/wiki/Eel_life_history?oldid=743802920 Contributors: PierreAbbat.svg License: Cc-by-sa-3.org/wikipedia/commons/8/8c/Lobster_%28PSF%29. Xezbeth. Matthead. Original artist: Gervais et Boulart • File:Glasseelkils.png created by User:Equazcion Original artist: Tkgd2007 • File:Rostrata. GreenC bot and Anonymous: 67 9. Matrix1919. Herugrim. Mandrake76.wikimedia. Bgwhite. TGCP. Addbot. by Peter Gugerell • File:Question_book-new.3 Content license • Creative Commons Attribution-Share Alike 3. Closedmouth. Favonian.gif Source: https://upload. Louperibot. BG19bot. Keilana. Nik42.jpg License: CC-BY-SA-3. Martpol.wikimedia. ZéroBot.png License: Public domain Contributors: Archives of Pearson Scott Foresman.wikimedia.jpg Source: https://upload.png Source: https://upload. Eyster.wikimedia. JNShutt.jpg Source: https://upload. Creidieki. AnomieBOT. Gilliam. Timwi.wikipedia to Commons.jpg License: CCBY-SA-3. Yerpo.svg Source: https://upload. NickPenguin. Ferrbott.wikimedia. Dthomsen8. H. Ian (Wiki Ed). Rich Farmbrough. Cuchullain. Menchi.0 Contributors: Photo by Uwe Kils Original artist: Uwe Kils • File:Rostrataluk. Wwoods.wikipedia to Commons by JohnnyMrNinja using CommonsHelper. Omnipaedista. RussBot.jpg Source: https://upload.jpg License: CC-BY-SA-3. Hoss. Fraggle81. CForrer. Zoicon5. Angel caboodle. Ptbotgourou. Philippe.gif License: CC-BY-SA-3.jpg License: CC-BY-SA-3. Le Anh-Huy. Original artist: The original uploader was Kils at English Wikipedia • File:Rostratamuk.wikipedia English Wikipedia 9.2 Images • File:Anguillamuk.jpg Source: https://upload.jpg License: Public domain Contributors: Les poissons Gervais. Bobauvergne.org/wikipedia/commons/3/3e/Rostrataluk. Anna Frodesiak.jpg Source: https://upload. BattyBot.org/wikipedia/commons/4/40/Glasseelskils. John of Reading. Discospinster.org/wikipedia/commons/9/94/Rostrata.7 9 Text and image sources. CatherineMunro. ClueBot.