Biology:Baculum
The baculum (pl.: bacula), also known as the penis bone, penile bone, os penis, os genitale,[1] or os priapi,[2] is a bone in the penis of many placental mammals. It is absent from the human penis, but present in the penises of some primates, such as the gorilla and the chimpanzee.[3][4] The baculum arises from primordial cells in soft tissues of the penis, and its formation is largely influenced by androgens.[5] The bone lies above the urethra,[6] and it aids sexual reproduction by maintaining stiffness during sexual penetration. The homologue to the baculum in female mammals is the baubellum (os clitoridis), a bone in the clitoris.[7][8][9]
Etymology
The word baculum means "stick" or "staff" in Latin and originates from Greek: βάκλον, baklon "stick".[10]
Function
The baculum is used for copulation and varies in size and shape by species. Its evolution may be influenced by sexual selection, and its characteristics are sometimes used to differentiate between similar species.[11] A bone in the penis allows a male to mate for a long time with a female,[12][13] which can be a distinct advantage in some mating strategies.[14][15] The length of the baculum may be related to the duration of copulation in some species.[16][17] In carnivorans[18] and primates, the length of the baculum appears to be influenced by postcopulatory sexual selection.[19] In some bat species, the baculum can also protect the urethra from compression.[20]
Presence in mammals
Mammals having a penile bone (in males) and a clitoral bone (in females) include various eutherians:
- Order Primates, although not in lorises,[21] humans, spider monkeys, or woolly monkeys[22]
- Order Rodentia (rodents),[23] though not in the related order Lagomorpha (rabbits, hares, etc.)[24]
- Order Eulipotyphla[25] (insectivores, including shrews and hedgehogs)
- Order Carnivora[9] (including members of many well-known families, such as ursids (bears),[26] canids (dogs),[1] pinnipeds (walruses, seals, sea lions),[6] procyonids (raccoons etc.),[27] mustelids (otters, weasels, skunks and others)).[28] The baculum is usually longer in the Canoidea than in the Feloidea, although fossas have long bacula and giant pandas have short bacula.[9]
- Order Chiroptera (bats).[29][30][31]
It is absent in humans, ungulates (hoofed mammals),[32] elephants, monotremes (platypus, echidna),[33] marsupials,[34] lagomorphs,[24] hyenas,[35] binturongs,[25] sirenians,[6] and cetaceans (whales, dolphins, and porpoises),[6] among others.
Evidence suggests that the baculum was independently evolved 9 times and lost in 10 separate lineages.[25] The baculum is an exclusive characteristic of placentals and closely related eutherians, being absent in other mammal clades, and it has been speculated to be derived from the epipubic bones more widely spread across mammals, but notoriously absent in placentals.[34]
Among the primates, marmosets,[clarification needed] weighing around 500 grams (18 oz), have a baculum measuring around 2 millimetres (0.079 in), while the tiny 63 g (2.2 oz) galago has one around 13 millimetres (0.51 in) long. The great apes, despite their size, tend to have very small penis bones, and humans are the only ones to have lost them altogether.[15]
In some mammalian species, such as badgers[36][37] and raccoons (Procyon lotor), the baculum can be used to determine relative age. If a raccoon's baculum tip is made up of uncalcified cartilage, has a porous base, is less than 1.2 g (0.042 oz) in mass, and measures less than 90 mm (3.5 in) long, then the baculum belongs to a juvenile.[27]
Absence in humans
Unlike most primates, humans lack an os penis,[38][39] but the bone is present, although much reduced, among other great apes. In many ape species, it is a relatively insignificant 10–20 mm (0.39–0.79 in) structure. Cases of human penis ossification following trauma have been reported,[40] and one case was reported of a congenital os penis surgically removed from a 5-year-old boy, who also had other developmental abnormalities, including a cleft scrotum.[41] Clellan S. Ford and Frank A. Beach in Patterns of Sexual Behavior (1951), p. 30 say, "Both gorillas and chimpanzees possess a penile bone. In the latter species, the os penis is located in the lower part of the organ and measures approximately three-quarters of an inch in length."[4] In humans, the rigidity of the erection is provided entirely through blood pressure in the corpora cavernosa. An "artificial baculum" or penile implant is sometimes used to treat erectile dysfunction in humans.[42]
In The Selfish Gene, Richard Dawkins[43] proposed honest advertising as the evolutionary explanation for the loss of the baculum. The hypothesis states that if erection failure is a sensitive early warning of ill health (physical or mental), females could have gauged the health of a potential mate based on his ability to achieve erection without the support of a baculum.
The tactile stimulation hypothesis proposes that the loss of the baculum in humans is linked to the female choice for tactile stimulation: a boneless penis would be more flexible, facilitating a larger range of copulatory positions and whole body movement, giving females greater general physical stimulation.[44]
The mating system shift hypothesis proposes that the shift towards monogamy as the dominant reproductive strategy may have reduced the intensity of copulatory and post-copulatory sexual selection, and made the baculum obsolete.[45][46]
Humans "evolved a mating system in which the male tended to accompany a particular female all the time to try to ensure paternity of her children"[15][better source needed] which allows for frequent matings of short duration. Observation suggests that primates with a baculum only infrequently encounter females, but engage in longer periods of copulation that the baculum makes possible, thereby maximizing their chances of fathering the female's offspring. Human females exhibit concealed ovulation, also known as hidden estrus, meaning it is almost impossible to tell when the female is fertile (unless the cervical mucus is examined),[47] so frequent matings would be necessary to ensure paternity.[15][48][49]
Strengths and weaknesses of these hypotheses were revised in a 2021 study, which also proposed an alternative hypothesis: that conspecific aggression, in combination with the development of self-awareness, may have played a role in the loss. If the presence of a baculum exacerbated the prevalence and severity of penile injuries resulting from blunt trauma to a flaccid penis, increasing ability to foresee the consequences of their actions would also enable hominins to realise that these injuries are a useful tool in male-male competition. This behavioural innovation, planned conspecific aggression with the goal of temporary exclusion of competitors from the breeding pool, would create an environment in which a genetic mutation for a penis without a baculum (or with an unossified baculum) would strongly increase the fitness of the mutant phenotype. Along with the hominin propensity for social learning and cultural transmission, this hypothetical scenario may explain why this phenotype became fixed in all human populations.[50]
An alternative view is that its loss in humans is an example of neoteny during human evolution; late-stage fetal chimpanzees lack a baculum.[51]
Cultural significance
It has been argued that the "rib" (Hebrew צֵלׇע ṣēlāʿ, also translated "flank" or "side") in the story of Adam and Eve is actually a mistranslation of a Biblical Hebrew euphemism for baculum, and that its removal from Adam in the Book of Genesis is a creation story to explain this absence (as well as the presence of the perineal raphe – as a resultant "scar") in humans.[52]
In Hoodoo, the folk magic of the American South, the raccoon baculum is sometimes worn as an amulet for love or luck.[53]
Oosik
Oosik (Iñupiaq: usuk or uzuk) is a term used in Alaska Native cultures to describe the bacula of walruses, seals, sea lions and polar bears. Sometimes as long as 60 cm (24 in), fossilized bacula are often polished and used as a handle for knives and other tools. The oosik is a polished and sometimes carved baculum of these large northern carnivores.
Oosiks are also sold as tourist souvenirs. In 2007, a 4.5 ft-long (1.4 m) fossilized penis bone from an extinct species of walrus, believed by the seller to be the largest in existence, was sold for $8,000.[54]
United States Congressman for Alaska, Don Young, was known for possessing an 18-inch walrus oosik, and once brandished it like a sword during a congressional hearing.[55]
See also
- Argumentum ad baculum – Latin expression describing an argument based on the use of force.
- Mammal penis
- Penile spines
References
- ↑ 1.0 1.1 Howard E. Evans; Alexander de Lahunta (7 August 2013). Miller's Anatomy of the Dog. Elsevier Health Sciences. ISBN 978-0-323-26623-9. https://books.google.com/books?id=6eBOAQAAQBAJ. Retrieved 27 October 2015.
- ↑ Dollé, P.; Izpistia-Belmonte, J.-C.=; Brown, J.M.; Tickle, C.; Duboule, D. (1991). "HOX-4 genes and the morphogenesis of mammalian genitalia". Genes & Development 5 (10): 1767–1776. doi:10.1101/gad.5.10.1767. PMID 1680771. Archived from the original on 20 July 2018. https://web.archive.org/web/20180720204753/http://genesdev.cshlp.org/content/5/10/1767.full.pdf.
- ↑ Dixson, Alan F. (26 January 2012). Primate Sexuality: Comparative Studies of the Prosimians, Monkeys, Apes, and Humans. OUP Oxford. ISBN 978-0-19-150342-9. https://books.google.com/books?id=khzhd2nXWM0C. Retrieved 13 February 2018.
- ↑ 4.0 4.1 Patterns of Sexual Behavior Clellan S. Ford and Frank A. Beach, published by Harper & Row, New York in 1951. ISBN:0-313-22355-6
- ↑ Nasoori, Alireza (2020). "Formation, structure, and function of extra‐skeletal bones in mammals". Biological Reviews 95 (4): 986–1019. doi:10.1111/brv.12597. PMID 32338826. https://archive.org/details/formation-structure-and-function-of-extra-skeletal-bones-in-mammals.
- ↑ 6.0 6.1 6.2 6.3 William F. Perrin; Bernd Wursig; J. G.M. Thewissen (26 February 2009). Encyclopedia of Marine Mammals. Academic Press. pp. 68–. ISBN 978-0-08-091993-5. https://books.google.com/books?id=2rkHQpToi9sC&q=pinniped+baculum&pg=PA68. Retrieved 30 October 2020.
- ↑ Best, Troy L.; Granai, Nancy J. (2 December 1994). "Tamius merriami". Mammalian Species (476): 1–9. doi:10.2307/3504203. http://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/i0076-3519-476-01-0001.pdf. Retrieved 15 January 2010.
- ↑ Burrows, Harold (1945). Biological Actions of Sex Hormones. Cambridge University Press. p. 264. ISBN 9780521043946. https://books.google.com/books?id=MFw4AAAAIAAJ&pg=PA264. Retrieved 4 August 2012.
- ↑ 9.0 9.1 9.2 Ewer, R. F. (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. https://books.google.com/books?id=IETMd3-lSlkC. Retrieved 16 December 2012.
- ↑ Liddell, Henry George; Scott, Robert. "βάκλον". Tufts University. https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0058:entry=ba/klon.
- ↑ Ramm, Steven A. "Sexual selection and genital evolution in mammals: a phylogenetic analysis of baculum length ." The American Naturalist 169.3 (2007): 360–369.
- ↑ Dixson, A. F. "Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)." Journal of Zoology 235.1 (1995): 67–76.
- ↑ DIXSON33, Alan, N. YHOL T. Jenna, and Matt Anderson. "A positive relationship between baculum length and prolonged intromission patterns in mammals ." 动物学报 50.4 (2004): 490–503.
- ↑ H Ferguson, Steven, and Serge Lariviere. "Are long penis bones an adaption to high latitude snowy environments? ." Oikos 105.2 (2004): 255–267.
- ↑ 15.0 15.1 15.2 15.3 "Godinotia". Walking With Beasts. ABC – BBC. 2002. pp. Question: How do we know how Godinotia (the primate in program 1) mated?. http://www.abc.net.au/beasts/evidence/prog1/page4_2.htm.
- ↑ Dixson, A. F. (1987). "Observations on the evolution of the genitalia and copulatory behaviour in male primates". Journal of Zoology 213 (3): 423–443. doi:10.1111/j.1469-7998.1987.tb03718.x.
- ↑ Stockley, Paula (2012). "The baculum". Current Biology 22 (24): R1032–R1033. doi:10.1016/j.cub.2012.11.001. PMID 23257184.
- ↑ Brassey, Charlotte A.; Behnsen, Julia; Gardiner, James D. (2020). "Postcopulatory sexual selection and the evolution of shape complexity in the carnivoran baculum". Proceedings of the Royal Society B: Biological Sciences 287 (1936). doi:10.1098/rspb.2020.1883. PMID 33049172.
- ↑ Brindle, Matilda, and Christopher Opie. "Postcopulatory sexual selection influences baculum evolution in primates and carnivores ." Proc. R. Soc. B. Vol. 283. No. 1844. The Royal Society, 2016.
- ↑ Herdina, Anna Nele; Kelly, Diane A.; Jahelková, Helena; Lina, Peter H. C.; Horáček, Ivan; Metscher, Brian D. (2015). "Testing hypotheses of bat baculum function with 3D models derived from microCT". Journal of Anatomy 226 (3): 229–235. doi:10.1111/joa.12274. PMID 25655647.
- ↑ Ronald M. Nowak; Ernest Pillsbury Walker (28 October 1999). Walker's Primates of the World. JHU Press. ISBN 978-0-8018-6251-9. https://archive.org/details/walkersprimateso0000nowa. "baculum."
- ↑ Harvey, Suzanne. "How Did Man Lose His Penis Bone?". University College London, Researchers in Museums blog, 26 November 2012. http://blogs.ucl.ac.uk/researchers-in-museums/2012/11/26/how-did-man-lose-his-penis-bone/.
- ↑ Harkness, John E.; Turner, Patricia V.; VandeWoude, Susan; Wheler, Colette L. (2 April 2013). Harkness and Wagner's Biology and Medicine of Rabbits and Rodents. John Wiley & Sons. ISBN 978-1-118-70907-8. https://books.google.com/books?id=GJIOPzAVNCoC. Retrieved 27 October 2015.
- ↑ 24.0 24.1 George A. Feldhamer; Lee C. Drickamer; Stephen H. Vessey; Joseph F. Merritt; Carey Krajewski (19 February 2015). Mammalogy: Adaptation, Diversity, Ecology. Johns Hopkins University Press. ISBN 978-1-4214-1589-5. https://books.google.com/books?id=CAq5BgAAQBAJ. Retrieved 13 February 2018.
- ↑ 25.0 25.1 25.2 Schultz, Nicholas G.; Lough-Stevens, Michael; Abreu, Eric; Orr, Teri; Dean, Matthew D. (2016-06-01). "The Baculum was Gained and Lost Multiple Times during Mammalian Evolution" (in en). Integrative and Comparative Biology 56 (4): 644–56. doi:10.1093/icb/icw034. ISSN 1540-7063. PMID 27252214.
- ↑ Dyck, Markus G.; Bourgeois, Jackie M.; Miller, Edward H. (2004). "Growth and variation in the bacula of polar bears (Ursus maritimus) in the Canadian Arctic". Journal of Zoology 264 (1): 105–110. doi:10.1017/S0952836904005606. http://journals.cambridge.org/action/displayAbstract;jsessionid=3332AADA7ADA71727F83E689232BAB80.tomcat1?fromPage=online&aid=241981. Retrieved 15 January 2010.
- ↑ 27.0 27.1 Nova J. Silvy (7 February 2012). The Wildlife Techniques Manual: Volume 1: Research. Volume 2: Management 2-vol. Set. JHU Press. ISBN 978-1-4214-0159-1. https://books.google.com/books?id=PL2IHTdzSeAC. Retrieved 27 October 2015.
- ↑ Baryshnikov, Gennady F.; Bininda-Emonds, Olaf R.P.; Abramov, Alexei V. (2003). "Morphological variability and evolution of the baculum (os penis) in Mustelidae (Carnivora)". Journal of Mammalogy 84 (2): 673–690. doi:10.1644/1545-1542(2003)084<0673:mvaeot>2.0.co;2.
- ↑ Hosken, D., et al. "Is the bat os penis sexually selected? ." Behavioral Ecology and Sociobiology 50.5 (2001): 450–460.
- ↑ Lüpold, S., A. G. McElligott, and D. J. Hosken. "Bat genitalia: allometry, variation and good genes ." Biological Journal of the Linnean Society 83.4 (2004): 497–507.
- ↑ Elizabeth G. Crichton; Philip H. Krutzsch (12 June 2000). Reproductive Biology of Bats. Academic Press. pp. 103–. ISBN 978-0-08-054053-5. https://books.google.com/books?id=f1aNgZwGsYoC&q=bat+(%22os+penis%22+OR+baculum)&pg=PA103. Retrieved 30 October 2020.
- ↑ Ronald M. Nowak (7 April 1999). Walker's Mammals of the World. JHU Press. pp. 1007–. ISBN 978-0-8018-5789-8. https://books.google.com/books?id=7W-DGRILSBoC&q=ungulate+baculum&pg=PA1007. Retrieved 30 October 2020.
- ↑ Grützner, F.; Nixon, B.; Jones, R.C. (2008). "Reproductive biology in egg-laying mammals". Sexual Development 2 (3): 115–127. doi:10.1159/000143429. PMID 18769071. https://www.researchgate.net/publication/23233860. Retrieved 3 June 2020.
- ↑ 34.0 34.1 Frederick S. Szalay (11 May 2006). Evolutionary History of the Marsupials and an Analysis of Osteological Characters. Cambridge University Press. pp. 293–. ISBN 978-0-521-02592-8. https://books.google.com/books?id=HmQTTChULOMC&q=marsupial+baculum&pg=PA293. Retrieved 30 October 2020.
- ↑ Richard Estes (1991). The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates. University of California Press. pp. 323–. ISBN 978-0-520-08085-0. https://archive.org/details/isbn_0520080858. Retrieved 12 December 2012. "boneless."
- ↑ Abramov, Alexei V. "Variation of the baculum structure of the Palaearctic badger (Carnivora, Mustelidae, Meles) ." Russian Journal of Theriology 1.1 (2002): 57–60.
- ↑ Ahnlund, H. "Age determination in the European badger, Meles meles L. " Zeitschrift für Säugetierkunde 41.1 (1976): 119–125.
- ↑ Martin, Robert D. (2007). "The evolution of human reproduction: A primatological perspective". American Journal of Physical Anthropology 134: 59–84. doi:10.1002/ajpa.20734. PMID 18046752.
- ↑ Friderun Ankel-Simons (27 July 2010). Primate Anatomy: An Introduction. Elsevier. ISBN 978-0-08-046911-9. https://books.google.com/books?id=Mwl3M6c5KzoC&q=pensi.
- ↑ Sarma, Deba; Thomas Weilbaecher (1990). "Human os penis". Urology 35 (4): 349–350. doi:10.1016/0090-4295(90)80163-H. PMID 2108520.
- ↑ Champion, RH; J Wegrzyn (1964). "Congenital os penis". Journal of Urology 91 (6): 663–4. doi:10.1016/S0022-5347(17)64197-1. PMID 14172255.
- ↑ Carrion, Hernan, et al. "A history of the penile implant to 1974." Sexual medicine reviews 4.3 (2016): 285–293.
- ↑ Dawkins R (2006) The Selfish Gene. Oxford University Press, Oxford, UK.
- ↑ Cormier LA, Jones SR (2015) The Domesticated Penis: How Womanhood Has Shaped Manhood. University of Alabama Press, Tuscaloosa, USA.
- ↑ Brennan PLR (2016) The evolution of genitalia. In: Shackelford TK, Weekes-Shackelford VA (eds) Encyclopedia of Evolutionary Psychological Science, 1–4. Springer International Publishing, Cham, Switzerland
- ↑ London, University College. "Study sheds light on the function of the penis bone in male competition" (in en). https://phys.org/news/2016-12-function-penis-bone-male-competition.html.
- ↑ Ilyich, Iryna. "Dr". https://flo.health/menstrual-cycle/health/vaginal-discharge/discharge-fluid-mucus.
- ↑ "Scientists have answered one of the biggest questions people have about their penis" (in en-GB). The Independent. 2016-12-14. https://www.independent.co.uk/life-style/health-and-families/health-news/penis-bone-humans-baculum-sex-doesnt-last-long-enough-ucl-why-a7473401.html.
- ↑ "ABC - Science - Beasts - Evidence - Programme 1 - Godinotia". 2014-04-29. http://www.abc.net.au/beasts/evidence/prog1/page4_2.htm.
- ↑ Jakovlić, Ivan (2021) “The Missing Human Baculum: A Victim of Conspecific Aggression and Budding Self-Awareness?” Mammal Review. https://doi.org/10.1111/mam.12237
- ↑ Bednarik, R. G. (2011). The Human Condition. doi:10.1007/978-1-4419-9353-3. ISBN 978-1-4419-9352-6. (page 134), cited by:
Achrati, Ahmed (November 2014). "Neoteny, female hominin and cognitive evolution". Rock Art Research 31 (1): 232–238.
"In humans, neoteny is manifested in the resemblance of many physiological features of a human to a late-stage foetal chimpanzee. These foetal characteristics include hair on the head, a globular skull, ear shape, vertical plane face, absence of penal bone (baculum) in foetal male chimpanzees, the vagina pointing forward in foetal ape, the presence of hymen in neonate ape, and the structure of the foot. 'These and many other features', Bednarik says, 'define the anatomical relationship between ape and man as the latter's neoteny'" - ↑ Gilbert, S. F.; Zevit, Z. (2001). "Congenital human baculum deficiency: The generative bone of Genesis 2:21-23". American Journal of Medical Genetics 101 (3): 284–85. doi:10.1002/ajmg.1387. PMID 11424148.
- ↑ Joanne O'Sullivan (1 March 2010). Book of Superstitious Stuff: Weird Happenings, Wacky Rites, Frightening Fears, Mysterious Myths & Other Bizarre Beliefs. Charlesbridge Publishing. p. 87. ISBN 978-1-60734-367-7. https://books.google.com/books?id=kQAUTJS5frAC&pg=PA87. Retrieved 27 October 2015. "In the hoodoo (folk magic) tradition of the American South, a raccoon penis bone (scientifically known as the baculum) is a lucky charm used to attract love. In some areas, it's boiled to remove any trace of the animal, and then tied to a red ribbon and worn as a necklace. In other areas, the bones were traditionally given to girls and young women by suitors, and in still other places, the charms are worn by men. Earrings made from cast raccoon penis bones became a fad in 2004, and celebrities such as Sarah Jessica Parker and Vanessa Williams were photographed wearing them. New Orleans gamblers are said to use the bones (also called coon dogs and Texas toothpicks) for luck."
- ↑ "Walrus penis sells for $8,000 at Beverly Hills action". AP. http://sfgate.com/cgi-bin/article.cgi?f=/n/a/2007/08/26/state/n154935D40.DTL.
- ↑ "A brief history of Rep. Don Young's incendiary remarks. (All right, it's a long history.)". Washington Post. https://www.washingtonpost.com/news/the-fix/wp/2018/02/28/a-brief-history-of-alaska-rep-don-youngs-incendiary-remarks-okay-its-a-long-history/.
Further reading
- Gilbert SF, Zevit Z (July 2001). "Congenital human baculum deficiency: the generative bone of Genesis 2:21–23". Am. J. Med. Genet. 101 (3): 284–5. doi:10.1002/ajmg.1387. PMID 11424148.
- Clellan S., Frank A. Beach (1951). Patterns of Sexual Behavior. New York: Harper, and Paul B. Hoeber, Inc. Medical Books. ISBN 978-0-313-22355-6.
External links
- Beresford WA, Burkart S (December 1977). "The penile bone and anterior process of the rat in scanning electron microscopy". J. Anat. 124 (3): 589–97. PMID 604330.
- The San Diego Zoo's Conservation and research for endangered species projects. 'What is the significance of the baculum in animals?'
- On the evolution of the mammalian baculum: vaginal friction, prolonged intromission or induced ovulation?
- The structure of the penis with the associated baculum in the male greater cane rat (Thryonomys swinderianus)
- Panciroli, Elsa (24 January 2018). "How do you sex a fossil? | Elsa Panciroli" (in en). The Guardian. https://www.theguardian.com/science/2018/jan/24/cloacae-sex-arms-and-penis-bones-the-tricky-art-of-fossil-sexing.
Original source: https://en.wikipedia.org/wiki/Baculum.
Read more |