Morphofunctional characteristics of the оs penis of mammals

Abstract

The study analyzed the morphofunctional organization of the mammalian baculum (os penis). In many mammals, bone tissue caused the penis to harden, which forms the bone of the penis (os penis or baculum). This structure appeared later in evolution than others, and it is present only in representatives of the most evolutionarily young class of vertebrates, i.e., mammals. The baculum is present in the penis of marsupials and placental mammals (rodents, some bats, pinnipeds, cetaceans, individual predators, semi-monkeys, and some monkeys).

The bone tissue is usually located in the anterior region of the penis between the cavernous and spongy bodies. The baculum is an unpaired bone. On the cross-section of representatives of different species, the baculum can be rounded, square, triangular, or flat. Most often, the baculum is a cylindrical rod, expanded in the proximal and distal parts, and it has outgrowths of various shapes. Histogenesis of the bone tissue during os penis formation in embryogenesis occurs by indirect osteogenesis. In the area of the future os penis, a cartilaginous model is formed from the mesenchyme, in place of which the bone tissue then develops. The base of the baculum is formed by lamellar bone tissue, in which osteones are well expressed. Coarse fibrous (reticulofibrous) bone tissue and fibrous cartilage tissue are found on the surface of the baculum. The baculum was found to increase the overall hardness of the penis when bending during copulation.

Keywords

mammals / penis / bone / os penis / baculum

References

[1] Dzerzhinsky FYa, Vasiliev BD, Malakhov VV. Zoology of vertebrates. Moscow: Izdatel’skij centr «Akademiya»; 2013. 408 p. (In Russ).

[2] Дзержинский Ф.Я., Васильев Б. Д., Малахов В. В. Зоология позвоночных. Москва: Издательский центр «Академия», 2013. 408 с.

[3] Shevlyuk NN. Comparative histology of the male reproductive system of vertebrates. Orenburg: Orenburgskij gosudarstvennyj medicinskij universitet; 2017. 168 p. EDN: MIODAN

[4] Шевлюк Н. Н. Сравнительная гистология мужской репродуктивной системы позвоночных. Оренбург: Оренбургский государственный медицинский университет, 2017. 168 с. EDN: MIODAN

[5] Comelis MT, Bueno LM, Góes RM, et al. Morphological and histological characters of penile organization in eleven species of molossid bats. Zoology (Jena). 2018;127:70–83. doi: 10.1016/j.zool.2018.01.006

[6] Comelis M.T., Bueno L. M., Góes R. M., et al. Morphological and histological characters of penile organization in eleven species of molossid bats // Zoology (Jena). 2018. Vol. 127. P. 70–83. doi: 10.1016/j.zool.2018.01.006

[7] Dixson AF. Copulatory and postcopulatory sexual selection in primates. Folia Primatol (Basel). 2018;89(3–4):258–286. doi: 10.1159/000488105

[8] Dixson A. F. Copulatory and postcopulatory sexual selection in primates // Folia Primatol (Basel). 2018. Vol. 89, N 3–4. P. 258–286. doi: 10.1159/000488105

[9] Ferguson SH, Lariviere S. Are long penis bone an adaptation to high latitude snowy environments? Oikos. 2004;105:255–267. doi: 10.1111/j.0030-1299.2004.13173.x

[10] Ferguson S.H., Lariviere S. Are long penis bone an adaptation to high latitude snowy environments? // Oikos. 2004. Vol. 105. P. 255–267. doi: 10.1111/j.0030-1299.2004.13173.x

[11] Rodriguez E Jr, Weiss DA, Yang JH, et al. New insights on the morphology of adult mouse penis. Biol Reprod. 2011;85(6):1216–1221. doi: 10.1095/biolreprod.111.091504

[12] Rodriguez E. Jr, Weiss D. A., Yang J. H., et al. New insights on the morphology of adult mouse penis // Biol Reprod. 2011. Vol. 85, N 6. P. 1216–1221. doi: 10.1095/biolreprod.111.091504

[13] Kalaida ML, Nigmetzyanova MV, Borisova SD. General histology and embryology of fish. 2th ed. Saint Petersburg: Lan’; 2018. 148 p. (In Russ).

[14] Калайда М.Л., Нигметзянова М. В., Борисова С. Д. Общая гистология и эмбриология рыб. 2-е издание. Санкт-Петербург: Лань, 2018. 148 с.

[15] Ognev SI. Zoology of vertebrates. 4th ed. Moscow: Sovetskaya nauka; 1945. 520 p. (In Russ).

[16] Огнев С. И. Зоология позвоночных. 4-е издание. Москва: Советская наука, 1945. 520 с.

[17] Ognev SI. Essays on the ecology of mammals. Moscow: Moskovskoe obshhestvo ispytanija prirody; 1951. 253 p. (In Russ).

[18] Огнев С. И. Очерки экологии млекопитающих. Москва: Московское общество испытания природы, 1951. 253 с.

[19] Aire TA. Anatomy of the testis and male reproductive tract. In: Jamieson BGM, editor. Reproductive biology and phylogeny of birds. Vol. 6A of series “Reproductive Biology and Phylogene. Enfield (NH): Science Publishers; 2007. P. 37–114.

[20] Aire T. A. Anatomy of the testis and male reproductive tract. In: Jamieson B. G.M., editor. Reproductive biology and phylogeny of birds. Vol. 6A of series “Reproductive Biology and Phylogene”. Enfield (NH): Science Publishers, 2007. P. 37–114.

[21] Sarma DP, Weilbaecher TG. Human os penis. Urology. 1990;35(4):349–350. doi: 10.1016/0090-4295(90)80163-h

[22] Sarma D.P., Weilbaecher T. G. Human os penis // Urology. 1990. Vol. 35, N 4. P. 349–350. doi: 10.1016/0090-4295(90)80163-h

[23] Jubilato FC, Comelis MT, Bueno LM, et al. Histomorphology of the glans penis in Vespertilionidae and Phyllostomidae (Chiroptera, Mammalia). J Morphol. 2019;280(12):1759–1776. doi: 10.1002/jmor.21062

[24] Jubilato F.C., Comelis M. T., Bueno L. M., et al. Histomorphology of the glans penis in Vespertilionidae and Phyllostomidae (Chiroptera, Mammalia) // J Morphol. 2019. Vol. 280, N 12. P. 1759–1776. doi: 10.1002/jmor.21062

[25] Herdina AN, Hulva P, Horasek I, et al. МicroCT imaging reveals morphometric baculum differences for discriminating the cryptic species Pipistrellus pipistrellus and P. pygmaeus. Acta Ciropterologica. 2014;16(1):157–168. doi: 10.3161/150811014X683372

[26] Herdina A.N., Hulva P., Horasek I., et al. МicroCT imaging reveals morphometric baculum differences for discriminating the cryptic species Pipistrellus pipistrellus and P. pygmaeus // Acta Ciropterologica. 2014. Vol. 16, N 1. P. 157–168. doi: 10.3161/150811014X683372

[27] Herdina AN, Herzig-Straschil B, Hilgers H, et al. Histomorphology of the penis bone (baculum) in the gray long-eared bat Plecotus austriacus (Chiroptera, Vespertilionidae). Anat Rec (Hoboken). 2010;293(7):1248–1258. doi: 10.1002/ar.21148

[28] Herdina A.N., Herzig-Straschil B., Hilgers H., et al. Histomorphology of the penis bone (baculum) in the gray long-eared bat Plecotus austriacus (Chiroptera, Vespertilionidae) // Anat Rec (Hoboken). 2010. Vol. 293, N 7. P. 1248–1258. doi: 10.1002/ar.21148

[29] Herdina AN, Kelly DA, Jahelková H, et al. Testing hypotheses of bat baculum function with 3D models derived from microCT. J Anat. 2015;226(3):229–235. doi: 10.1111/joa.12274

[30] Herdina A.N., Kelly D. A., Jahelková H., et al. Testing hypothesis of bat baculum function with 3D models derived from microCT // J Anat. 2015. Vol. 226, N 3. P. 229–235. doi: 10.1111/joa.12274

[31] Lariviere S, Ferguson SH. On the evolution of the mammalian baculum: vaginal friction, prolonged intromission or induced ovulation? Mammalian Review. 2002;32(4):283–294. doi: 10.1046/j.1365-2907.2002.00112.x

[32] Lariviere S., Ferguson S. H. On the evolution of the mammalian baculum: vaginal friction, prolonged intromission or induced ovulation? // Mammalian Review. 2002. Vol. 32, N 4. P. 283–294. doi: 10.1046/j.1365-2907.2002.00112.x

[33] Orr TJ, Brennan PL. All features great and small-the potential roles of the baculum and penile spines in mammals. Integr Comp Biol. 2016;56(4):635–643. doi: 10.1093/icb/icw057

[34] Orr T.J., Brennan P. L. All features great and small — the potential roles of the baculum and penile spines in mammals // Integr Comp Biol. 2016. Vol. 56, N 4. P. 635–643. doi: 10.1093/icb/icw057

[35] Schultz NG, Lough-Stevens M, Abreu E, et al. The baculum was gained and lost multiple times during mammalian evolution. Integr Comp Biol. 2016;56(4):644–656. doi: 10.1093/icb/icw034

[36] Schultz N.G., Lough-Stevens M., Abreu E., et al. The baculum was gained and lost multiple times during mammalian evolution // Integr Comp Biol. 2016. Vol. 56, N 4. P. 644–656. doi: 10.1093/icb/icw034

[37] Simmnons LW, Firman RC. Experimental evidence for the evolution of the Mammalian baculum by sexual selection. Evolution. 2014;68(1):276–283. doi: 10.1111/evo.12229

[38] Simmnons L.W., Firman R. C. Experimental evidence for the evolution of the mammalian baculum by sexual selection // Evolution. 2014. Vol. 68, N 1. P. 276–283. doi: 10.1111/evo.12229

[39] Stockley P. The baculum. Curr Biol. 2012;22(24): R1032–R1033. doi: 10.1016/j.cub.2012.11.001

[40] Stockley P. The baculum // Curr Biol. 2012. Vol. 22, N 24. P. R1032–R1033. doi: 10.1016/j.cub.2012.11.001

[41] Vilmann A, Vilmann H. Os penis of the rat. IV. The proximal growth cartilage. Acta Anat (Basel). 1983;117(2):136–144. doi: 10.1159/000145779

[42] Vilmann A., Vilmann H. Os penis of the rat. IV. The proximal growth cartilage // Acta Anat (Basel). 1983. Vol. 117, N 2. P. 136–144. doi: 10.1159/000145779

[43] Ozen AHS. Morphological vatiability of the baculum in Martes foina (Carnivora: Mustelidae) from Turkey. Turkish Journal of Zoology. 2018;42:666–672. doi: 10.3906/zoo-1802-39

[44] Ozen A.H.S. Morphological vatiability of the baculum in Martes foina (Carnivora: Mustelidae) from Turkey // Turkish Journal of Zoology. 2018. Vol. 42. P. 666–672. doi: 10.3906/zoo-1802-39

[45] Smirnov DG, Tsytsulina K. The ontogeny of the baculum in Ntctalus noctula and Vespertilio murinus (Chiroptera: Vespertilionidae). Acta Chiropterologica. 2003;5(1):117–123. doi: 10.3161/001.005.0111

[46] Smirnov D.G., Tsytsulina K. The ontogeny of the baculum in Ntctalus noctula and Vespertilio murinus (Chiroptera: Vespertilionidae) // Acta Chiropterologica. 2003. Vol. 5, N 1. P. 117–123. doi: 10.3161/001.005.0111

[47] Murakami R. Immunohistochemical and immunoblot analyses of collagens in the developing fibrocartilage in the glans penis of the rat. Acta Morphol Neerl Scand. 1987;25(4):279–288.

[48] Murakami R. Immunohistochemical and immunoblot analyses of collagens in the developing fibrocartilage in the glans penis of the rat // Acta Morphol Neerl Scand. 1987. Vol. 25, N 4. P. 279-288.

[49] Williams-Ashman HG, Reddi AH. Differentiation of mesenchymal tissues during phallic morphogenesis with emphasis on the os penis: roles of androgens and other regulatory agents. J Steroid Biochem Mol Biol. 1991;39(6):873–881. doi: 10.1016/0960-0760(91)90344-5

[50] Williams-Ashman H.G., Redid A. H. Differentiation of mesenchymal tissues during phallic morphogenesis with emphasis on the os penis: roles of androgens and other regulatory agents // J Steroid Biochem Mol Biol. 1991. Vol. 39, N 6. P. 873–881. doi: 10.1016/0960-0760(91)90344-5

[51] Pole SB. Subspecific and interpopulation variability of the gray marmot baculum. Holarctic marmots as a factor of biodiversity. In: Proceedings of the III International Conference on marmots. Cheboksary, 1997 Aug 25–30. Moscow; 1997. P. 76–77. (In Russ).

[52] Поле С. Б. Подвидовая и межпопуляционная изменчивость бакулюма серого сурка. Сурки Голарктики как фактор биоразнообразия. В кн.: Сборник тезисов III международной конференции по суркам. Чебоксары, 25–30 августа 1997 г. Москва, 1997. С. 76–77.

[53] Chervyakova VP. On the structure of the baculum of marmots. Zoologicheskiy zhurnal. 1966;45(11):1712–1719.

[54] Червякова В.П. О строении бакулюма сурков // Зоологический журнал. 1966. Т. 45, № 11. С. 1712–1719.

[55] Shevlyuk NN, Rudi VN, Stadnikov AA. Biology of reproduction of terrestrial rodents from the squirrel family (morphological, physiological and ecological aspects). Ekaterinburg: Izdatel’stvo UrO RAN; 1999. 146 p. EDN: TLSMJP

[56] Шевлюк Н.Н., Руди В. Н., Стадников А. А. Биология размножения наземных грызунов из семейства беличьих (морфологические, физиологические и экологические аспекты). Екатеринбург: УрО РАН, 1999. 146 с. EDN: TLSMJP

[57] Kelly DA. Anatomy of the baculum-corpus cavernosum interface in the norway rat (Rattus norvegicus), and implications for force transfer during copulation. J Morphol. 2000;244(1):69–77. doi: 10.1002/(SICI)1097–4687(200004)244:1<69::AID-JMOR7>3.0.CO;2-#

[58] Kelly D. A. Anatomy of the baculum — corpus cavernosum interface in the Norway rat (Rattus norvegicus), and implications for force transfer during copulation // J Morphol. 2000. Vol. 244, N 1. P. 69–77. doi: 10.1002/(SICI)1097-4687(200004)244:1<69::AID-JMOR7>3.0.CO;2-#

[59] Leon-Alvarado OD, Ramirez-Chaves HE. Morphological drscription of the glans penis and baculum of Coendou quichua (Rodentia: Erethizontidae). Therya. 2017;8(4):283–294. doi: 10.12933/therya-17-495

[60] Leon-Alvarado O.D., Ramirez-Chaves H. E. Morphological drscription of the glans penis and baculum of Coendou quichua (Rodentia: Erethizontidae) // Therya. 2017. Vol. 8, N 4. P. 283–294. doi: 10.12933/therya-17-495

[61] Ramm SA, Khoo L, Stockley P. Sexual selection and the rodent baculum: an intraspecific study in the house mouse (Mus musculus domesticus). Genetica. 2010;138(1):129–137. doi: 10.1007/s10709-009-9385-8

[62] Ramm S.A., Khoo L., Stockley P. Sexual selection and the rodent baculum: an intraspecific study in the house mouse (Mus musculus domesticus) // Genetica. 2010. Vol. 138, N 1. P. 129–137. doi: 10.1007/s10709-009-9385-8

[63] Reserford WA, Burkart S. The penile bone and anterior process of the rat in scanning electron microscopy. J Anat. 1977;124(Pt 3):589–597.

[64] Reserford W.A., Burkart S. The penile bone and anterior process of the rat in scanning electron microscopy // J Anat. 1977. Vol. 124, N 3. P. 589–597.

[65] Schultz NG, Ingels J, Hillhouse A, et al. The genetic basis of baculum size and shape variation in mice. G3 (Bethesda). 2016;6(5):1141–1151. doi: 10.1534/g3.116.027888

[66] Schultz N.G., Ingels J., Hillhouse A., et al. The genetic basis of baculum size and shape variation in mice // G3 (Bethesda). 2016. Vol. 6, N 5. P. 1141–1151. doi: 10.1534/g3.116.027888

[67] Vilmann H, Vilmann A. Os penis of the rat. II. Morphology of the mature bone. Anat Anz. 1979;146(5):483–493.

[68] Vilmann H., Vilmann A. Os penis of the rat. II. Morphology of mature bone // Anat Anz. 1979. Vol. 146, N 5. P. 483–493.

[69] Abramov AV. Variation of the baculum structure of the Palearctic badger (Carnivora, Mustelidae, Meles). Russian J Ther. 2002;(1):57–60. doi: 10.15298/rusjtheriol.01.1.04

[70] Abramov A. V. Variation of the baculum structure of the Palearctic badger (Carnivora, Mustelidae, Meles) // Russian J Ther. 2002. N 1. P. 57–60. doi: 10.15298/rusjtheriol.01.1.04

[71] Baryshnikov GF, Bininda-Emonds OR, Abramov AV. Morphological variability and evolution of the baculum (os penis) in Mustelidae (Carnivora). Journal Mammal. 2003;84:673–690. doi: 10.1644/1545-1542(2003)084<0673:MVAEOT>2.0.CO;2

[72] Baryshnikov G.F., Bininda-Emonds O.R., Abramov A. V. Morphological variability and evolution of the baculum (os penis) in Mustelidae (Carnivora) // Journal Mammal. 2003. Vol. 84. P. 673–690. doi: 10.1644/1545–1542(2003)084<0673: MVAEOT>2.0.CO;2

[73] Canady A. Variability of the baculum in the red fox (Vulpes vulpes) from Slovakia. Zool Ecol. 2013;23:165–170. doi: 10.1080/21658005.2013.832848

[74] Canady A. Variability of the baculum in the red fox (Vulpes vulpes) from Slovakia // Zool Ecol. 2013. Vol. 23. P. 165–170. doi: 10.1080/21658005.2013.832848

[75] Canady A, Comor L. Contribution to knowledge of the variability of the penis bone (baculum) in the Eurasian wolf (Canis lupus) from Slovakia. Lynx. 2013;44:5–12.

[76] Canady A., Comor L. Contribution to knowledge of the variability of the penis bone (baculum) in the Eurasian wolf (Canis lupus) from Slovakia // Lynx. 2013. Vol. 44. P. 5–12.

[77] Canady A, Comor L. Allometry of the baculum in the wolf (Canis lupus, Canidae) as an indicator viability and quality in males. Zool Ecol. 2015;25:192–198. doi: 10.1080/21658005.2015.1044164

[78] Canady A., Comor L. Allometry of the baculum in the wolf (Canis lupus, Canidae) as an indicator viability and quality in males // Zool Ecol. 2015. Vol. 25. P. 192–198. doi: 10.1080/21658005.2015.1044164

[79] Canady A, Onderkova A. Are size, variability and allometry of the baculum in relation to body length signals of a good condition in male weasels Mustela nivalis. Zool Anz. 2016;264:29–33. doi: 10.1016/j.jcz.2016.07.003

[80] Canady A., Onderkova A. Are size, variability and allometry of the baculum in relation to body length signals of a good condition in male weasels Mustela nivalis // Zool Anz. 2016. Vol. 264. P. 29–33. doi: 10.1016/j.jcz.2016.07.003

[81] Dixson AF. Baculum length and copulatort behavior in carnivores and ginnipeds (grand order Ferae). Journal of Zoology. 1995;235(1):67–76. doi: 10.1111/j.1469-7998.1995.tb05128.x

[82] Dixson A. F. Baculum length and copulatort behavior in carnivores and ginnipeds (grand order Ferae) // Journal of Zoology. 1995. Vol. 235, N 1. P. 67–76. doi: 10.1111/j.1469–7998.1995.tb05128.x

[83] Schulte-Hostedde A, Bowman J, Middel KR. Allometry of the baculum and sexual size dimorphism in American martens and fishers (Mammalia: Mustelidae). Biol J Linn Soc. 2011;104:955–963. doi: 10.1111/j.1095-8312.2011.01775.x

[84] Schulte-Hostedde A., Bowman J., Middel K. R. Allometry of the baculum and sexual size dimorphism in American martens and fishers (Mammalia: Mustelidae) // Biol J Linn Soc. 2011. Vol. 104. P. 955–963. doi: 10.1111/j.1095-8312.2011.01775.x

[85] Sharir A, Israeli D, Milgram J, et al. The canine baculum: the structure and mechanical properties of an unusual bone. J Struct Biol. 2011;175(3):451–456. doi: 10.1016/j.jsb.2011.06.006

[86] Sharir A., Israeli D., Milgram J., et al. The canine baculum: the structure and mechanical properties of an unusual bone // J Struct Biol. 2011. Vol. 175, N 3. P. 451–456. doi: 10.1016/j.jsb.2011.06.006

[87] Vercillo F, Ragni B. Morphometric discrimination between Martes martes and Martyes foina in Italy: the use of rhe baculum. Hystrix. 2011;22:325–331.

[88] Vercillo F., Ragni B. Morphometric discrimination between Martes martes and Martyes foina in Italy: the use of rhe baculum // Hystrix. 2011. Vol. 22. P. 325–331.

[89] Dixson AF. Baculum length and copulatory behavior in primates. Am J Primatol. 1987;13(1):51–60. doi: 10.1002/ajp.1350130107

[90] Dixson A. F. Baculum length and copulatory behavior in primates // Am J Primatol. 1987. Vol. 13, N 1. P. 51–60. doi: 10.1002/ajp.1350130107

[91] Perkin A. Comparative penile morphology of east African galagos of the genus Galagoides (family Galagoidae): implications for taxonomy. Am J Primatol. 2007;69(1):16–26. doi: 10.1002/ajp.20323

[92] Perkin A. Comparative penile morphology of east African galagos of the genus Galagoides (family Galagoidae): implications for taxonomy // Am J Primatol. 2007. Vol. 69, N 1. P. 16–26. doi: 10.1002/ajp.20323

[93] Gilbert T. The Os priapi of suckling animals. Gegenbaurs Morphologisches Jahrbuch. 1892;18:805–808. (In Germany).

[94] Gilbert T. Das Os priapi der Saugethiere // Gegenbaurs Morphologisches Jahrbuch. 1892. Vol. 18. P. 805–808.

[95] Gilbert SF, Zevit Z. Congenital human baculum deficiency: the generative bone of Genesis 2:21–23. Am J Med Genet. 2001;101(3):284–285. doi: 10.1002/ajmg.1387

[96] Gilbert S.F., Zevit Z. Congenital human baculum deficiency: the generative bone of Genesis 2:21–23 // Am J Med Genet. 2001. Vol. 101, N 3. P. 284–285. doi: 10.1002/ajmg.1387

[97] Høeg OM. Human penile ossification. A case report. Scand J Urol Nephrol. 1986;20(3):231–232. doi: 10.3109/00365598609024501

[98] Høeg O. M. Human penile ossification. A case report // Scand J Urol Nephrol. 1986. Vol. 20, N 3. P. 231–232. doi: 10.3109/00365598609024501

[99] Vahlensiesk WK Jr, Schaefer HE, Westenfelder M. Penile ossification and acquired penile deviation. Eur Urol. 1995;27(3):252–256. doi: 10.1159/000475172

[100] Vahlensiesk W. K. Jr, Schaefer H. E., Westenfelder M. Penille ossification and acquired penile deviation // Eur Urol. 1995. Vol. 27, N 3. P. 252–256. doi: 10.1159/000475172

[101] Smirnov DG. Structural features of the baubellum of bats (Chiroptera, Vespertilionidae) of the Middle Volga region. In: Orlov VN, editor. Theriofauna of Russia and adjacent territories. Proceedings of the International Meeting VII Congress of the All-Russian Theriological Society. 2003 Feb 6–7. Moscow: IPEE RAS; 2003. P. 325–327.

[102] Смирнов Д. Г. Особенности строения баубеллюм рукокрылых (Chiroptera, Vespertilionidae) Среднего Поволжья. В кн.: Териофауна России и сопредельных территорий. VII съезд Всероссийского териологического общества. Материалы Международного совещания / под ред. В. Н. Орлова. 6–7 февраля 2003 г. Москва: ИПЭЭ РАН, 2003. С. 325–327.

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