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Penis

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Penis
Penis of anAsian elephant
Details
PrecursorGenital tubercle(amniotes)
SystemReproductive system,sometimes with thegenitourinary system
Identifiers
Latinpenis
Anatomical terminology

Apenis(/ˈpnɪs/;pl.:penisesorpenes) is a malesex organthat is used toinseminatefemale orhermaphroditeanimals duringcopulation.[1][2]Such organs occur in bothvertebratesandinvertebrates,but not in all male animals.

The termpenisapplies to manyintromittent organs,but not to all. As an example, the intromittent organ of mostCephalopodais thehectocotylus,a specialized arm, and male spiders use theirpedipalps.Even within theVertebrata,there are morphological variants with specific terminology, such ashemipenes.

Etymology

The word "penis" is taken from theLatinword for "tail".Some derive that fromIndo-European*pesnis,and the Greek word πέος = "penis" from Indo-European*pesos.Prior to the adoption of the Latin word in English, the penis was referred to as a "yard". TheOxford English Dictionarycites an example of the wordyardused in this sense from 1379,[3]and notes that in hisPhysical Dictionaryof 1684,Steven Blankaartdefined the wordpenisas "the Yard, made up of two nervous Bodies, the Channel, Nut, Skin, and Fore-skin, etc."[4]According toWiktionary,this term meant (among other senses) "rod" or "bar".

As with nearly any aspect of the body involved in sexual orexcretoryfunctions, the penis is the subject of manyslangwords andeuphemismsfor it, a particularly common and enduring one being "cock". SeeWikiSaurus:penisfor a list of alternative words for penis.

The Latin word"phallus"(fromGreekφαλλος) is sometimes used to describe the penis, although "phallus" originally was used to describerepresentations,pictorial or carved, of the penis.[5]

Evolution and function

The external genital organs appeared in theDevonian,about 410 million years ago, whentetrapodsbegan to abandon the aquatic environment.[6]In fact, the necessity to overcome the absence of a liquid phase in which to release the gametes was achieved through the transition tointernal fertilization.

Among amniotes, the development of an erectile penis occurred independently formammals,squamates(lizardsandsnakes),testudines(turtles), andarchosaurs(crocodilesandbirds).

Over time, birds have lost this organ, with the exception ofPaleognathaeandAnseriformes.[7]

The penis is anintromittent organused to transfersperminto the femalegenital tractfor potentialfertilizationand, in the case ofplacentals,also for the excretion ofurine.[8][9]The penises of different animal groups are nothomologouswith each other, but were created several times independently of each other in the course of evolution.

Anerectionis the stiffening and rising of the penis, which occurs duringsexual arousal,though it can also happen in non-sexual situations. Duringejaculation,a series of muscular contractions delivers semen, containing malegametesknown as sperm cells orspermatozoa,from the penis. Ejaculation is usually accompanied byorgasm.

The last common ancestor of all livingamniotes(mammals, birds and reptiles) likely possessed a penis.[10]

Vertebrates

Birds

Maleduckshave a corkscrew-shaped penis while female ducks have corkscrew vaginas with many blind pockets designed for difficult penetration and to prevent becoming pregnant.[11]This reduced the likelihood of fertilization by unwanted aggressors in favor of fitter mates.

Most male birds (e.g.,roostersandturkeys) have acloaca(also present on the female), but not a penis. Among bird species with a penis arepaleognaths(tinamousandratites)[12]andAnatidae(ducks, geese and swans).[13]A bird penis is different in structure from mammal penises, being an erectile expansion of the cloacal wall (in ducks) and being erected bylymph,not blood.[14]It is usually partially feathered and in some species features spines and brush-like filaments, and in a flaccid state, curls up inside the cloaca.

Mammals

Penis of ahorse
Penis of acat
Penis of adog(Great Dane)

As with any other bodily attribute, the length and girth of the penis can be highly variable betweenmammalsof differentspecies.[15][16]In many mammals, the penis is retracted into aprepucewhen not erect. Mammals have eithermusculocavernouspenises, which expand while erect, orfibroelasticpenises, which become erect by straightening without expanding.[17]Preputial glandsare present in some prepuces. The penis bearsdistalpart of theurethrainplacental mammals.[8]Theperineumoftesticondmammals separates theanusand the penis.

A bone called thebaculumis present in most placentals but absent in humans, cattle and horses.

In mammals, the penis is divided into three parts:[18]

The internal structures of the penis consist mainly of cavernous,erectile tissue,which is a collection ofblood sinusoidsseparated by sheets ofconnective tissue(trabeculae).

Genitorinary system of a maleraccoon(Procyon lotor)

Canine peniseshave a structure at the base called thebulbus glandis.[19][20]During copulation, thespotted hyenainserts his penis throughthe female's pseudo-penisinstead of directly through thevagina,which is blocked by the falsescrotum.Thepseudo-penisclosely resembles the male hyena's penis, but can be distinguished from the male's genitalia by its greater thickness and more roundedglans.[21]Domestic catshave barbed penises, with about 120–150 one millimetre longbackwards-pointing spines.[22]

Marsupialsusually have bifurcated penises[23]that are retracted into a preputial sheath in the male'surogenital sinuswhen not erect.[24]Monotremesandmarsupial molesare the only mammals in which the penis is located inside the cloaca.[25][26]

Reptiles

Maleturtlesandcrocodileshave a penis, while male specimens of the reptile orderSquamata,which aresnakesandlizards,have two paired organs calledhemipenes.Tuatarasmust use their cloacae for reproduction.[27]Due toevolutionary convergence,turtle and mammal penises have a similar structure.[28]

Fish

In some fish, the gonopodium, andropodium, andclaspersare intromittent organs (to introduce sperm into the female) developed from modified fins.[29]

Invertebrates

The spine-covered penis ofCallosobruchus analis,abean weevil

In maleinsects,the structure analogous to a penis is known asaedeagus.The male copulatory organ of various lower invertebrate animals is often called thecirrus.[30]

In 2010, entomologist Charles Linehard described a new genus ofbarkfliescalledNeotrogla.Species of this genus have sex-reversed genitalia: females have penis-like organs called gynosomes that are inserted into vagina-like openings of males during mating.[31]A similar female structure has also been described in the closely relatedAfrotrogla.[32]Scientists who study these insects have occasionally called the gynosome a "female penis"[33][34]and insisted to drop the definition of penis as "the male copulatory organ".[35]Motivations for using the term “female penis” include that such a term "is easier to understand and much more eye-catching"[36]and that the gynosome have "analogous features" with male penises.[35]Meanwhile, critics have argued that it does not fit the intromittent organ definition of "a structure that enters the female genital tract and deposits sperm".[37]

Heraldry

Pizzles are represented inheraldry,where the adjectivepizzled(orvilené[38]) indicates that part of an animatecharge's anatomy, especially if coloured differently.

See also

References

Citations

  1. ^Janet Leonard; Alex Cordoba-Aguilar R (18 June 2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press.ISBN978-0-19-971703-3.Archivedfrom the original on 11 October 2013.Retrieved20 July2013.
  2. ^Schmitt, V.; Anthes, N.; Michiels, N. K. (2007)."Mating behaviour in the sea slug Elysia timida (Opisthobranchia, Sacoglossa): hypodermic injection, sperm transfer and balanced reciprocity".Frontiers in Zoology.4:17.doi:10.1186/1742-9994-4-17.PMC1934903.PMID17610714.
  3. ^Basu, S. C. (2011).Male Reproductive Dysfunction.JP Medical Ltd. p. 101.ISBN9789350252208.
  4. ^Simpson, John,ed. (1989)."penis,n.".Oxford English Dictionary(second ed.).Oxford University Press.ISBN978-0-19-861186-8.[dead link]
  5. ^"Online Etymology Dictionary".Etymonline.com.Archivedfrom the original on 2011-06-06.Retrieved2011-05-28.
  6. ^Dunlop, Jason A.; Penney, David.Fossil Arachnids.Siri Scientific Press. p. 44.ISBN978-0-95677-954-0.
  7. ^Cordoba-Aguilar, Alex; Leonard, Janet (2010).The Evolution of Primary Sexual Characters in Animals.Oxford University Press. pp. 216–221.ISBN978-0-19971-703-3.
  8. ^abMarvalee H. Wake (15 September 1992).Hyman's Comparative Vertebrate Anatomy.University of Chicago Press. p. 583.ISBN978-0-226-87013-7.Archivedfrom the original on 31 December 2013.Retrieved23 July2013.
  9. ^Feldhamer, George A. (2007-09-07).Mammalogy: Adaptation, Diversity, Ecology.JHU Press.ISBN978-0-8018-8695-9.
  10. ^Sanger TJ, Gredler ML, Cohn MJ (October 2015)."Resurrecting embryos of the tuatara, Sphenodon punctatus, to resolve vertebrate phallus evolution".Biology Letters.11(10): 20150694.doi:10.1098/rsbl.2015.0694.PMC4650183.PMID26510679.
  11. ^Brennan, Patricia L. R.; Clark, Christopher J.; Prum, Richard O. (2010-05-07)."Explosive eversion and functional morphology of the duck penis supports sexual conflict in waterfowl genitalia".Proceedings of the Royal Society B: Biological Sciences.277(1686): 1309–1314.doi:10.1098/rspb.2009.2139.ISSN0962-8452.PMC2871948.PMID20031991.
  12. ^Julian Lombardi (1998).Comparative Vertebrate Reproduction.Springer.ISBN978-0-7923-8336-9.Archivedfrom the original on 26 March 2014.Retrieved5 December2012.
  13. ^MobileReference (15 December 2009).The Illustrated Encyclopedia of European Birds: An Essential Guide to Birds of Europe.MobileReference.ISBN978-1-60501-557-6.Archived fromthe originalon 26 March 2014.Retrieved5 December2012.
  14. ^Frank B. Gill (6 October 2006).Ornithology.Macmillan. pp. 414–.ISBN978-0-7167-4983-7.Archivedfrom the original on 7 January 2014.Retrieved5 December2012.
  15. ^Tim Birkhead (2000).Promiscuity: An Evolutionary History of Sperm Competition.Harvard University Press. p. 102.ISBN978-0-674-00666-9.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.
  16. ^Virginia Douglass Hayssen; Ari Van Tienhoven (1993).Asdell's Patterns of Mammalian Reproduction: A Compendium of Species-Specific Data.Cornell University Press.ISBN978-0-8014-1753-5.Retrieved23 July2013.
  17. ^Chenoweth, Peter J.; Lorton, Steven P. (2022-02-03).Manual of Animal Andrology.CABI.ISBN978-1-78924-350-5.
  18. ^William O. Reece (2009-03-04).Functional Anatomy and Physiology of Domestic Animals.John Wiley & Sons.ISBN9780813814513.Archivedfrom the original on 2018-03-20.
  19. ^Susan Long (2006).Veterinary Genetics and Reproductive Physiology.Churchill Livingstone Elsevier.ISBN978-0-7506-8877-2.Archivedfrom the original on 2014-03-26.Retrieved2013-11-08.
  20. ^R. F. Ewer (1998).The Carnivores.Cornell University Press. p. 116.ISBN978-0-8014-8493-3.Archivedfrom the original on 26 March 2014.Retrieved23 July2013.
  21. ^Glickman, SE; Cunha, GR; Drea, CM; Conley, AJ; Place, NJ (2006)."Mammalian sexual differentiation: lessons from the spotted hyena"(PDF).Trends Endocrinol Metab.17(9): 349–356.doi:10.1016/j.tem.2006.09.005.PMID17010637.S2CID18227659.Archived(PDF)from the original on 2013-02-22.
  22. ^Aronson, L. R.; Cooper, M. L. (1967)."Penile spines of the domestic cat: their endocrine-behavior relations"(PDF).Anat. Rec.157(1): 71–8.doi:10.1002/ar.1091570111.PMID6030760.S2CID13070242.Archived fromthe original(PDF)on 2017-06-20.
  23. ^Tyndale-Biscoe, C. Hugh; Renfree, Marilyn (1987-01-30).Reproductive Physiology of Marsupials.Cambridge University Press.ISBN978-0-521-33792-2.
  24. ^Armati, Patricia J.; Dickman, Chris R.; Hume, Ian D. (2006-08-17).Marsupials.Cambridge University Press.ISBN978-1-139-45742-2.
  25. ^Gadow, H. On the systematic position of Notoryctes typhlops. Proc. Zool. Soc. London 1892, 361–370 (1892).
  26. ^Riedelsheimer, B., Unterberger, P., Künzle, H. and U. Welsch. 2007. Histological study of the cloacal region and associated structures in the hedgehog tenrec Echinops telfairi. Mammalian Biology 72(6): 330-341.
  27. ^Lutz, Dick (2005), Tuatara: A Living Fossil, Salem, Oregon: DIMI PRESS,ISBN0-931625-43-2
  28. ^Kelly, D. A. (2004)."Turtle and mammal penis designs are anatomically convergent".Proceedings. Biological Sciences.271(Suppl 5): S293–S295.doi:10.1098/rsbl.2004.0161.PMC1810052.PMID15503998.
  29. ^Ruschenberger, William Samuel Waithman (1846).Elements of Herpetology, and of Ichthyology: Prepared for the Use of Schools and Colleges.Grigg & Elliot. pp. 129–145.
  30. ^"Penis | Description, Anatomy, & Physiology | Britannica".Encyclopædia Britannica.January 2024.
  31. ^Lienhard, Charles; Oliveira do Carmo, Thais; Lopes Ferreira, Rodrigo (2010)."A new genus of Sensitibillini from Brazilian caves (Psocodea: 'Psocoptera': Prionoglarididae)".Revue Suisse de Zoologie.117(4): 611–635.doi:10.5962/bhl.part.117600.ISSN0035-418X.Archivedfrom the original on 2014-11-03.
  32. ^Yoshizawa K, Ferreira R.L., Yao I, Lienhard C & Kamimura Y. "Independent origins of female penis and its coevolution with male vagina in cave insects (Psocodea: Prionoglarididae)".Biology Letters14(11):doi.org/10.1098/rsbl.2018.0533
  33. ^Yoshizawa, Kazunori; Ferreira, Rodrigo L.; Kamimura, Yoshitaka; Lienhard, Charles (2014)."Female Penis, Male Vagina, and Their Correlated Evolution in a Cave Insect".Current Biology.24(9): 1006–10.doi:10.1016/j.cub.2014.03.022.hdl:2115/56857.ISSN0960-9822.PMID24746797.
  34. ^Yoshizawa K, Ferreira R.L., Lienhard C & Kamimura Y. (2019). "Why Did a Female Penis Evolve in a Small Group of Cave Insects?".BioEssays41(6):doi.org/10.1002/bies.201900005
  35. ^abYoshizawa K, Ferreira R.L., Kamimura Y & Lienhard C."A Transgender Brazilian Cave Insect".The Winnower3/9/2014
  36. ^Hollier J & Hollier A. (2020)."The retired taxonomist and the gynosome – the discovery of the female penis".Antenna44(3): p. 122-125
  37. ^Hayssen V. (2020). "Misconceptions about Conception and Other Fallacies: Historical Bias in Reproductive Biology".Integrative and Comparative Biology60(3): p. 683-791:doi.org/10.1093/icb/icaa035
  38. ^Rietstap, J. B. (1884)."Armorial général; précédé d'un Dictionnaire des termes du blason".G. B. van Goor zonen: XXXI.Vilené: se dit un animal qui a la marque du sexe d'un autre émail que le corps{{cite journal}}:Cite journal requires|journal=(help)

General and cited references

Horses

Marsupials

Other animals

External links