Baculum of a dog's penis; the arrow shows the urethral sulcus, which is the groove in which the urethra lies.

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

A raccoon baculum

Mammals having a penile bone (in males) and a clitoral bone (in females) include various eutherians:

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, 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] 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

Walrus baculum, around 22 inches (56 cm) long

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

References

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  2. ↑ Dollé, P.; Izpistia-Belmonte, J.-C.=; Brown, J.M.; Tickle, C.; Duboule, D. (1991). "HOX-4 genes and the morphogenesis of mammalian genitalia" (PDF). Genes & Development. 5 (10): 1767–1776. doi:10.1101/gad.5.10.1767. PMID 1680771. S2CID 6307427. Archived from the original (PDF) on 20 July 2018.
  3. ↑ 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. Archived from the original on 31 December 2013. Retrieved 13 February 2018.
  4. 1 2 Patterns of Sexual Behavior Clellan S. Ford and Frank A. Beach, published by Harper & Row, New York in 1951. ISBN 0-313-22355-6
  5. ↑ 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. S2CID 216556342.
  6. 1 2 3 4 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. Archived from the original on 21 October 2021. Retrieved 30 October 2020.
  7. ↑ Best, Troy L.; Granai, Nancy J. (2 December 1994). "Tamius merriami" (PDF). Mammalian Species (476): 1–9. doi:10.2307/3504203. JSTOR 3504203. S2CID 253909941. Archived (PDF) from the original on 23 June 2010. Retrieved 15 January 2010.
  8. ↑ Burrows, Harold (1945). Biological Actions of Sex Hormones. Cambridge University Press. p. 264. ISBN 9780521043946. Archived from the original on 21 October 2021. Retrieved 4 August 2012.
  9. 1 2 3 Ewer, R. F. (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. Archived from the original on 28 May 2013. Retrieved 16 December 2012.
  10. ↑ Liddell, Henry George; Scott, Robert. "βάκλον". An Intermediate Greek-English Lexicon. Tufts University. Archived from the original on 5 February 2017. Retrieved 9 April 2017.
  11. ↑ Ramm, Steven A. "Sexual selection and genital evolution in mammals: a phylogenetic analysis of baculum length Archived 30 September 2017 at the Wayback Machine." The American Naturalist 169.3 (2007): 360–369.
  12. ↑ Dixson, A. F. "Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)." Journal of Zoology 235.1 (1995): 67–76. Archived 3 June 2016 at the Wayback Machine
  13. ↑ DIXSON33, Alan, N. YHOL T. Jenna, and Matt Anderson. "A positive relationship between baculum length and prolonged intromission patterns in mammals Archived 6 August 2017 at the Wayback Machine." 动物学报 50.4 (2004): 490–503.
  14. ↑ H Ferguson, Steven, and Serge Lariviere. "Are long penis bones an adaption to high latitude snowy environments? Archived 6 August 2017 at the Wayback Machine." Oikos 105.2 (2004): 255–267.
  15. 1 2 3 4 "Godinotia". Walking With Beasts. ABC – BBC. 2002. pp. Question: How do we know how Godinotia (the primate in program 1) mated?. Archived from the original on 29 April 2014. Retrieved 7 July 2015.
  16. ↑ 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.
  17. ↑ Stockley, Paula (2012). "The baculum". Current Biology. 22 (24): R1032–R1033. doi:10.1016/j.cub.2012.11.001. PMID 23257184.
  18. ↑ 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. PMC 7657853. PMID 33049172.
  19. ↑ Brindle, Matilda, and Christopher Opie. "Postcopulatory sexual selection influences baculum evolution in primates and carnivores Archived 24 April 2022 at the Wayback Machine." Proc. R. Soc. B. Vol. 283. No. 1844. The Royal Society, 2016.
  20. ↑ 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. PMC 4337662. PMID 25655647.
  21. ↑ Ronald M. Nowak; Ernest Pillsbury Walker (28 October 1999). Walker's Primates of the World. JHU Press. ISBN 978-0-8018-6251-9. baculum.
  22. ↑ Harvey, Suzanne. "How Did Man Lose His Penis Bone?". University College London, Researchers in Museums blog, 26 November 2012. Archived from the original on 25 October 2014. Retrieved 20 November 2014.
  23. ↑ 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. Archived from the original on 24 April 2022. Retrieved 27 October 2015.
  24. 1 2 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. Archived from the original on 24 April 2022. Retrieved 13 February 2018.
  25. 1 2 3 Schultz, Nicholas G.; Lough-Stevens, Michael; Abreu, Eric; Orr, Teri; Dean, Matthew D. (1 June 2016). "The Baculum was Gained and Lost Multiple Times during Mammalian Evolution". Integrative and Comparative Biology. 56 (4): 644–56. doi:10.1093/icb/icw034. ISSN 1540-7063. PMC 6080509. PMID 27252214.
  26. ↑ 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. CiteSeerX 10.1.1.464.4517. doi:10.1017/S0952836904005606. Archived from the original on 31 October 2014. Retrieved 15 January 2010.
  27. 1 2 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. Archived from the original on 24 April 2022. Retrieved 27 October 2015.
  28. ↑ 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.
  29. ↑ Hosken, D., et al. "Is the bat os penis sexually selected? Archived 1 August 2017 at the Wayback Machine." Behavioral Ecology and Sociobiology 50.5 (2001): 450–460.
  30. ↑ Lüpold, S., A. G. McElligott, and D. J. Hosken. "Bat genitalia: allometry, variation and good genes Archived 22 August 2017 at the Wayback Machine." Biological Journal of the Linnean Society 83.4 (2004): 497–507.
  31. ↑ Elizabeth G. Crichton; Philip H. Krutzsch (12 June 2000). Reproductive Biology of Bats. Academic Press. pp. 103–. ISBN 978-0-08-054053-5. Archived from the original on 24 April 2022. Retrieved 30 October 2020.
  32. ↑ Ronald M. Nowak (7 April 1999). Walker's Mammals of the World. JHU Press. pp. 1007–. ISBN 978-0-8018-5789-8. Archived from the original on 24 April 2022. Retrieved 30 October 2020.
  33. ↑ 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. S2CID 536033. Archived from the original on 24 April 2022. Retrieved 3 June 2020.
  34. 1 2 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. Archived from the original on 24 April 2022. Retrieved 30 October 2020.
  35. ↑ 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. Retrieved 12 December 2012. boneless.
  36. ↑ Abramov, Alexei V. "Variation of the baculum structure of the Palaearctic badger (Carnivora, Mustelidae, Meles) Archived 30 January 2019 at the Wayback Machine." Russian Journal of Theriology 1.1 (2002): 57–60.
  37. ↑ Ahnlund, H. "Age determination in the European badger, Meles meles L. Archived 30 January 2019 at the Wayback Machine" Zeitschrift für Säugetierkunde 41.1 (1976): 119–125.
  38. ↑ 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. S2CID 44416632.
  39. ↑ Friderun Ankel-Simons (27 July 2010). Primate Anatomy: An Introduction. Elsevier. ISBN 978-0-08-046911-9.
  40. ↑ Sarma, Deba; Thomas Weilbaecher (1990). "Human os penis". Urology. 35 (4): 349–350. doi:10.1016/0090-4295(90)80163-H. PMID 2108520.
  41. ↑ 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.
  42. ↑ Carrion, Hernan, et al. "A history of the penile implant to 1974." Sexual medicine reviews 4.3 (2016): 285–293.
  43. ↑ Dawkins R (2006) The Selfish Gene. Oxford University Press, Oxford, UK.
  44. ↑ Cormier LA, Jones SR (2015) The Domesticated Penis: How Womanhood Has Shaped Manhood. University of Alabama Press, Tuscaloosa, USA.
  45. ↑ 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
  46. ↑ London, University College. "Study sheds light on the function of the penis bone in male competition". phys.org. Retrieved 15 July 2023.
  47. ↑ Ilyich, Iryna. "Dr". Flo.health. Retrieved 1 June 2023.
  48. ↑ "Scientists have answered one of the biggest questions people have about their penis". The Independent. 14 December 2016. Archived from the original on 19 August 2017. Retrieved 15 December 2016.
  49. ↑ "ABC - Science - Beasts - Evidence - Programme 1 - Godinotia". Australian Broadcasting Corporation. 29 April 2014. Archived from the original on 29 April 2014. Retrieved 23 November 2020.
  50. ↑ 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
  51. ↑ 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'"
  52. ↑ 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.
  53. ↑ 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. Archived from the original on 29 December 2015. 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.
  54. ↑ "Walrus penis sells for $8,000 at Beverly Hills action". AP. Archived from the original on 6 November 2007. Retrieved 30 August 2007.
  55. ↑ "A brief history of Rep. Don Young's incendiary remarks. (All right, it's a long history.)". Washington Post. Archived from the original on 5 December 2020. Retrieved 18 March 2021.

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.
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