Maintenance of Thermal Homeostasis with Special Emphasis on Testicular Thermoregulation

The testicular thermoregulation is maintained by a complex anatomical structures and physiological phenomena including convection, conduction, radiation and evaporation. The testicular temperature variation has a deleterious outcome on the reproductive performance of male reproductive system, though, duration and intensity of the insult is the pivot point. By regulating the altered thermal offense, the reproductive performance can be improved, however, with a consistent increase in the global temperature, there is a need of extensive investigations to undermine the thermoregulatory mechanism, its performance, anatomical and physiological adaptations, in the upcoming hot and humid environment.


Abu-zaid RM, Gad MR, 1999. Goat testicular blood supply: A study by corrosion cast and angiography. Journal of the Egyptian Veterinary Medical Association 59.

Abdullahi IA, Musa HA, Jibril A, 2012. Scrotal circumference and testicular morphometric characteristics of the camel (Camelus Dromedarius) in the Semi-Arid Environment of Northern Nigeria. International Journal of Morphology 30, 1369-1372. [CrossRef]

Amakiri SF, 1974. Sweat gland measurements in some tropical and temperate breeds of cattle in Nigeria. Animal Production 18, 285- 291. [CrossRef]

Amselgruber W, Sinowatz F, 1987. Zur Beziehung zwischen der Arteria testicularis und den Venen des Plexus pampiniformis beim Bullen. Anatomia, Histologia, Embryologia 16, 363-370.

Artyukhin AA, 2007a. Anatomy and microanatomy of the venous system of scrotal organs and spermatic cord. Bulletin of Experimental Biology and Medicine 143, 99-104. [CrossRef]

Artyukhin AA, 2007b. Vascular buffer system of the scrotal organs. Bulletin of Experimental Biology and Medicine 143, 642-647. [CrossRef]

Asala S, Chaudhary SC, Masumbuko-Kahamba N, Bidmos M, 2001. Anatomical variations in the human testicular blood vessels. Annals of Anatomy - Anatomischer Anzeiger 183, 545-549. [CrossRef]

Atkinson S, 1997. Reproductive biology of seals. Reviews of Reproduction 2, 175-194. [CrossRef]

Austin J, Hupp E, Murphee R, 1961. Effect of scrotal insulation on semen of Hereford bulls. Journal of Animal Science 20, 307.

Barth AD, Bowman PA, 1994. The sequential appearance of sperm abnormalities after scrotal insulation or dexamethasone treatment in bulls. The Canadian Veterinary Journal 35, 93-102.

Bartlett JM, Sharpe RM, 1987. Effect of local heating of the rat testis on the levels in interstitial fluid of a putative paracrine regulator of the Leydig cells and its relationship to changes in Sertoli cell secretory function. Journal of Reproduction and Fertility 80, 279-287. [CrossRef]

Bedford JM, 2004. Enigmas of mammalian gamete form and function. Biological Reviews of the Cambridge Philosophical Society 79, 429- 460. [CrossRef]

Blazquez NB, Long SE, Mayhew TM, Perry GC, Prescott NJ, Wathes C, 1994. Rate of discharge and morphology of sweat glands in the perineal, lumbodorsal and scrotal skin of cattle. Research in Veterinary Science 57, 277-284. [CrossRef]

Blazquez NB, Mallard GJ, Wedd SR, 1988. Sweat glands of the scrotum of the bull. Journal of Reproduction and Fertility 83, 673-677. [CrossRef]

Borthakur S, Dhingra LD, 1979. Anatomical studies on the blood vessels of the testis in buffalo (Bubalus bubalis). Indian Journal of Animal Science 49, 266.

Boulant JA, Hardy JD, 1974. The effect of spinal and skin temperatures on the firing rate and thermosensitivity of preoptic neurones. The Journal of Physiology 240, 639-660. [CrossRef]

Bratincsák A, Palkovits M, 2005. Evidence that peripheral rather than intracranial thermal signals induce thermoregulation. Neuroscience 135, 525-532. [CrossRef]

Brito LFC, Barth AD, Wilde RE, Kastelic JP, 2012. Testicular vascular cone development and its association with scrotal temperature, semen quality, and sperm production in beef bulls. Animal Reproduction Science 134, 135-140. [CrossRef]

Brito LFC, Silva AEDF, Barbosa RT, Kastelic JP, 2004. Testicular thermoregulation in Bos indicus, crossbred and Bos taurus bulls: relationship with scrotal, testicular vascular cone and testicular morphology, and effects on semen quality and sperm production. Theriogenology 61, 511-528. [CrossRef]

Chance MRA, 1996. Reason for externalization of the testis of mammals. Journal of Zoology 239, 691-695. [CrossRef]

Cividini FG, 2017. Characterization of the circulatory parameters of the testicular supra artery in young bulls of the aberdeen angus race. Unopar Universidade.

Cook RB, Coulter GH, Kastelic JP, 1994. The testicular vascular cone, scrotal thermoregulation, and their relationship to sperm production and seminal quality in beef bulls. Theriogenology 41, 653-671. [CrossRef]

Coulter G, Kastelic J, 1994. Testicular thermoregulation in bulls In: Proceedings of the 15th Technical Conference on Artificial Insemination Reproduction (pp. 28–34).

Craig AD, 2002. How do you feel? Interoception: the sense of the physiological condition of the body. Nature Reviews Neuroscience 3, 655- 666. [CrossRef]

Dhingra LD, 1979. Angioarchitecture of the buffalo testis. Anatomischer Anzeiger 146, 60-68.

Eheenberg L, Ehrenstein GV, Hedgran A, 1957. Gonad Temperature and Spontaneous Mutation-rate in Man. Nature 180, 1433-1434. [CrossRef]

Einer-Jensen N, Hunter RHF, 2005. Counter-current transfer in reproductive biology. Reproduction 129, 9-18. [CrossRef]

El-Gaafary MA, Aly AE, El-Ayat MA, 1980. Morphological studies on the testicular artery in the Egyptian water buffalo (Bos Bubalis L.). The Egyptain Veterinary Medical Journal 28.

Elayat MA, Khalil KM, Farag FM, Rizk HM, 2014. Gross anatomical studies on the pattern and density of the tunica vasculosa testis in some farm animals (buffalo, ram, camel, donkey and rabbit). Benha Veterinary Medical Journal 26, 1-9.

Fehlings K, Pohlmeyer K, 1978. Die Arteria testicularis und ihre Aufzweigung im Hoden und Nebenhoden des Esels (Equus africanus f. asinus): Korrosionsanatomische und röntgenologische Untersuchungen. Anat Histol Embryol 7, 74-78. [CrossRef]

Freeman S, 1990. The evolution of the scrotum: a new hypothesis. Journal of Theoretical Biology 145, 429-445. [CrossRef]

Gábor G, Sasser RG, Kastelic JP, Coulter GH, Falkay G, Mézes M, Szász F, 1998. Morphologic, endocrine and thermographic measurements of testicles in comparison with semen characteristics in mature Holstein-Friesian breeding bulls. Animal Reproduction Science 51, 215-224. [CrossRef]

Galil KA, Setchell BP, 1988. Effects of local heating of the testis on testicular blood flow and testosterone secretion in the rat. International Journal of Andrology 11, 73-85. [CrossRef]

Garcia AR, 2013. Estratégias para elevar a eficiência reprodutiva de búfalos em ambientes tropicais In: Proceedings of the 10th Brazilian Congress of Buiatrics (pp. 1–11). Belém: Brazilian Buiatrics Association.

Geerling JC, Loewy AD, 2008. Central regulation of sodium appetite. Experimental Physiology 93, 177-209. [CrossRef]

Glad Sørensen H, Lambrechtsen J, Einer-Jensen N, 1991. Efficiency of the countercurrent transfer of heat and 133Xenon between the pampiniform plexus and testicular artery of the bull under in-vitro conditions. International Journal of Andrology 14, 232-240. [CrossRef]

Gibson A, Akinrinsola A, Patel T, Ray A, Tucker J, McFadzean I, 2002. Pharmacology and thermosensitivity of the dartos muscle isolated from rat scrotum. British Journal of Pharmacology 136, 1194-1200. [CrossRef]

Godynicki SZ, Meyer W, Jackowiak H, Bollwein H, Pfarrer CH, 2013. Microvascularisation of the scrotal integument of Holstein-Friesian bulls. Folia Morphologica (Poland) 72, 132-136. [CrossRef]

Guieu JD, Hardy JD, 1970. Effects of heating and cooling of the spinal cord on preoptic unit activity. Journal of Applied Physiology 29, 675- 683. [CrossRef]

Gunn S, Gould T, 1975. Vasculature of the testes and adnexa In Handbook of Physiology (pp. 117–142). Washington: American Physiological Society.

Gupta BN, Nier K, Hensel H, 1979. Cold-sensitive afferents from the abdomen. Pflugers Archiv: European Journal of Physiology 380, 203- 204. [CrossRef]

Guyton AC, Hall JE, John E, 2006. Textbook of medical physiology. Elsevier Saunders.

Hansen PJ, 2009. Effects of heat stress on mammalian reproduction. Philosophical Transactions of the Royal Society B: Biological Sciences 364, 3341-3350. [CrossRef]

Hees H, Leiser R, Kohler T, Wrobel KH, 1984. Vascular morphology of the bovine spermatic cord and testis. Cell and Tissue Research 237, 31-38. [CrossRef]

Hodges GJ, Kosiba WA, Zhao K, Johnson JM, 2008. The involvement of norepinephrine, neuropeptide Y, and nitric oxide in the cutaneous vasodilator response to local heating in humans. Journal of Applied Physiology 105, 233-240. [CrossRef]

Holowatz LA, Thompson CS, Minson CT, Kenney WL, 2005. Mechanisms of acetylcholine-mediated vasodilatation in young and aged human skin. The Journal of Physiology 563, 965–973. [CrossRef]

Jannes P, Spiessens C, Van der Auwera I, D’Hooghe T, Verhoeven G, Vanderschueren D, 1998. Male subfertility induced by acute scrotal heating affects embryo quality in normal female mice. Human Reproduction (Oxford, England) 13, 372-375. [CrossRef]

Jung A, Eberl M, Schill WB, 2001. Improvement of semen quality by nocturnal scrotal cooling and moderate behavioural change to reduce genital heat stress in men with oligoasthenoteratozoospermia. Reproduction (Cambridge, England) 121, 595-603. [CrossRef]

Kastelic JP, 2014a. Thermoregulation of the Testes In R. M. Hopper (Ed.), Bovine Reproduction (1st ed., pp. 26–29). John Wiley & Sons, Inc. [CrossRef] Kastelic JP, 2014b. Understanding and evaluating bovine testes. Theriogenology 81, 18-23. [CrossRef]

Kastelic JP, Cook RB, Coulter GH, 1996a. Contribution of the scrotum and testes to scrotal and testicular thermoregulation in bulls and rams. Journal of Reproduction and Fertility 108, 81-85. [CrossRef]

Kastelic JP, Cook RB, Coulter GH, 1997a. Contribution of the scrotum, testes, and testicular artery to scrotal/testicular thermoregulation in bulls at two ambient temperatures. Animal Reproduction Science 45, 255-261. [CrossRef]

Kastelic JP, Cook RB, Coulter GH, 1997b. Scrotal/testicular thermoregulation and the effects of increased testicular temperature in the bull. The Veterinary Clinics of North America. Food Animal Practice 13, 271-282. [CrossRef]

Kastelic JP, Cook RB, Coulter GH, 1999. Effects of ambient temperature and scrotal fleece cover on scrotal and testicular temperatures in rams. Canadian Journal of Veterinary Research = Revue Canadienne de Recherche Veterinaire 63, 157-160.

Kastelic JP, Cook RB, Coulter GH, Saacke RG, 1996b. Insulating the scrotal neck affects semen quality and scrotal/testicular temperatures in the bull. Theriogenology 45, 935-942. [CrossRef]

Kastelic JP, Cook RB, Pierson RA, Coulter GH, 2001. Relationships among scrotal and testicular characteristics, sperm production, and seminal quality in 129 beef bulls. Canadian Journal of Veterinary Research = Revue Canadienne de Recherche Veterinaire 65, 111-115.

Kastelic JP, Coulter GH, Cook RB, 1995. Scrotal surface, subcutaneous, intratesticular, and intraepididymal temperatures in bulls. Theriogenology 44, 147-152. [CrossRef]

Kellogg DL, Hodges GJ, Orozco CR, Phillips TM, Zhao JL, Johnson JM, 2007. Cholinergic mechanisms of cutaneous active vasodilation during heat stress in cystic fibrosis. Journal of Applied Physiology 103, 963-968. [CrossRef]

Khalifa EF, Noor NA, 2017. Anatomical study on the testicular artery of bull (Bos Taurus). International Journal of Advanced Research in Biological Sciences 4, 80-89.

Khalil KMHA, 2013. Comparative anatomical studies on the vascular architecture of the blood vessels of the testis of some farm animals (Thesis). Faculty of Veterinary Medicine - Department of Anatomy and Embryology, Cairo University.

Lagerlof N, 1938. Infertility in male domestic animals. Veterinary Medicine 33, 550.

Langley JN, Anderson HK, 1895. The Innervation of the Pelvic and adjoining Viscera: Part II. The Bladder. Part III. The External Generative Organs. Part IV. The Internal Generative Organs. Part V. Position of the Nerve Cells on the Course of the Efferent Nerve Fibres. The Journal of Physiology 19, 71-139. [CrossRef]

Li Q, Thornhill J, 1993. Neuronal activity changes of ventromedial hypothalamic neurons and associated temperature responses in rats following scrotal thermal stimulation. Canadian Journal of Physiology and Pharmacology 71, 604-610. [CrossRef]

Li Q, Thornhill J, 1996. A functional medial preoptic nucleus (MPO) is required for scrotal thermal stimuli to alter the neuronal activity of thermoresponsive ventromedial hypothalamic (VMH) neurons. Brain Research 716, 134-140. [CrossRef]

Li Q, Thornhill J, 1998. Thermoresponsiveness of posterior hypothalamic (PH) neurons of rats to scrotal and abdominal thermal stimulation. Brain Research 794, 80-87. [CrossRef]

Liedtke WB, 2017. Deconstructing mammalian thermoregulation. Proceedings of the National Academy of Sciences 114, 1765-1767. [CrossRef]

Lomax P, Malveaux E, Smith RE, 1964. Brain temperatures in the rat during exposure to low environmental temperatures. American Journal of Physiology-Legacy Content 207, 736-739. [CrossRef]

Lue YH, Sinha Hikim AP, Swerdloff RS, Im P, Taing KS, Bui T, Wang C, 1999. Single Exposure to Heat Induces Stage-Specific Germ Cell Apoptosis in Rats: Role of Intratesticular Testosterone on Stage Specificity. Endocrinology 140, 1709-1717. [CrossRef]

Lunstra DD, Coulter GH, 1997. Relationship between scrotal infrared temperature patterns and natural-mating fertility in beef bulls. Journal of Animal Science 75, 767-774. [CrossRef]

Lyne AG, Hollis DE, 1968. The skin of the sheep: a comparison of body regions. Australian Journal of Biological Sciences 21, 499-527. [CrossRef]

Main SJ, Waites GM, 1977. The blood-testis barrier and temperature damage to the testis of the rat. Journal of Reproduction and Fertility 51, 439-450. [CrossRef]

Maloney SK, Bonomelli JM, DeSouza J, 2003. Scrotal heating stimulates panting and reduces body temperature similarly in febrile and non-febrile rams (Ovis aries). Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 135, 565-573. [CrossRef]

Marai IFM, Darawany AAE, Fadiel A, Abdel-Hafez MAM, 2008. Reproductive performance traits as affected by heat stress and its alleviation in sheep. Tropical and Subtropical Agroecosystems 8, 209- 234.

Maurya VP, Sejian V, Kumar D, Naqvi SMK, 2016. Impact of heat stress, nutritional restriction and combined stresses (heat and nutritional) on growth and reproductive performance of Malpura rams under semi-arid tropical environment. Journal of Animal Physiology and Animal Nutrition 100, 938-946. [CrossRef]

McKemy DD, Neuhausser WM, Julius D, 2002. Identification of a cold receptor reveals a general role for TRP channels in thermosensation. Nature 416, 52-58. [CrossRef]

McLaren TT, Foster PM, Sharpe RM, 1994. Identification of stage-specific changes in protein secretion by isolated seminiferous tubules from rats following exposure to short-term local testicular heating. Journal of Reproduction and Fertility 102, 293-300. [CrossRef]

Mieusset R, Sowerbutts SF, Zupp JL, Setchell BP, 1992. Increased flow of testicular blood plasma during local heating of the testes of rams. Journal of Reproduction and Fertility 94, 345-352. [CrossRef]

Moore CR, Quick WJ, 1924. The scrotum as a temperature regulator for the testes. American Journal of Physiology-Legacy Content 68, 70-79. [CrossRef]

Morrison SF, Nakamura K, 2011. Central neural pathways for thermoregulation. Frontiers in Bioscience (Landmark Edition) 16, 74-104. [CrossRef]

Mostafa T, Labib I, El-Khayat Y, El-Rahman El-Shahat A, Gadallah A, 2008. Human testicular arterial supply: gross anatomy, corrosion cast, and radiologic study. Fertility and Sterility 90, 2226-2230. [CrossRef]

Nakamura K, Morrison SF, 2007. Central efferent pathways mediating skin cooling-evoked sympathetic thermogenesis in brown adipose tissue. American Journal of Physiology. Regulatory, Integrative and Comparative Physiology 292, 127-36. [CrossRef]

Nakamura K, Morrison SF, 2008 A thermosensory pathway that controls body temperature. Nature-Neuroscience 11, 62-71. [CrossRef]

Nakayama T, Eisenman JS, Hardy JD, 1961. Single unit activity of anterior hypothalamus during local heating. Science (New York, N.Y.) 134, 560-561. [CrossRef]

Nakayama T, Hammel HT, Hardy JD, Eisenman JS, 1963. Thermal stimulation of electrical activity of single units of the preoptic region. American Journal of Physiology-Legacy Content 204, 1122-1126. [CrossRef]

Nunes AS, Filho MFC, Júnior AANM, Abreu-Silva AL, Júnior AMC, Souza JAT, Carvalho MAM, 2010. Scrotum histological description in native goats from Piaui State, according to scrotal bipartition level. Ciência Rural 40, 1808-1813. [CrossRef]

Osman DI, Tingari MD, Moniem KA, 1979. Vascular supply of the testis of the camel (Camelus dromedarius). Acta Anatomica 104, 16-22. [CrossRef]

Pasqualotto FF, Lucon AM, Hallak J, Goes PM, Saldanha LB, Arap S, 2003. Induction of spermatogenesis in azoospermic men after varicocele repair. Human Reproduction 18, 108-112. [CrossRef]

Pasha RH, Qureshi AS, Lodhi LA, Jamil H, 2011. Biometric and Ultrasonographic evaluation of the testis of one-humped camel (Camelus dromedarius). Pakistan Veterinary Journal 31, 129-133.

Paul C, Murray AA, Spears N, Saunders PTK, 2008. A single, mild, transient scrotal heat stress causes DNA damage, subfertility and impairs formation of blastocysts in mice. Reproduction 136, 73-84. [CrossRef]

Paul C, Teng S, Saunders PTK, 2009. A single, mild, transient scrotal heat stress causes hypoxia and oxidative stress in mouse testes, which induces germ cell death. Biology of Reproduction 80, 913-919. [CrossRef]

Pérez-Crespo M, Pintado B, Gutiérrez-Adán A, 2008. Scrotal heat stress effects on sperm viability, sperm DNA integrity, and the offspring sex ratio in mice. Molecular Reproduction and Development 75, 40-47. [CrossRef]

Portmann A, 1952. Animal forms and patterns : a study of the appearance of animals. London. Raman JD, Goldstein M, 2004. Intraoperative characterization of arterial vasculature in spermatic cord. Urology 64, 561-564. [CrossRef]

Robertshaw D, Vercoe J, 1980a. Scrotal thermoregulation of the bull (Bos sp.) Australian Journal of Agricultural Research 31, 401. [CrossRef]

Robertshaw D, Vercoe J, 1980b. Scrotal thermoregulation of the bull (Bos sp.) Australian Journal of Agricultural Research 31, 401-407. [CrossRef]

Qureshi AS, Masood A, Ali, MZ, Shahid, RU, 2015. Effects of natural environment conditions on reproductive histo-morphometric dynamics of male dromedary camel, in: Qureshi, M.S., Khan, S., Qureshi, I.U.H. (Eds.), 3rd International Workshop Dairy Science Park. Dairy Science Park, Peshawar, p. 13.

Saab Abi S, Fawwak TS, Najib K, Walid YD, Pauline YA, 2011. Effect of adaptation and heat stress on reproductive performances of fattail awassi rams in eastern mediterranean. Lebanese Science Journal 12, 31-44.

Sailer BL, Sarkar LJ, Bjordahl JA, Jost LK, Evenson DP, 1997. Effects of heat stress on mouse testicular cells and sperm chromatin structure. Journal of Andrology 18, 294-301.

Schäfer SS, Schäfer S, 1973. The role of the primary afference in the generation of a cold shivering tremor. Experimental Brain Research 17, 381-393. [CrossRef]

Schwarz GM, Hirtler L, 2017. The cremasteric reflex and its muscle - a paragon of ongoing scientific discussion: A systematic review. Clinical Anatomy 30, 498-507. [CrossRef]

Sealfon AI, Zorgniotti AW, 1991. A theoretical model for testis thermoregulation. Advances in Experimental Medicine and Biology 286, 123-135. [CrossRef]

Setchell BP, 1978. The Scrotum and Thermoregulation In The Mammalian Testis (pp. 90–108). New York: Cornell University Press.

Setchell BP, 1998. The Parkes Lecture. Heat and the testis. Journal of Reproduction and Fertility 114, 179-194. [CrossRef]

Setchell BP, 2006. The effects of heat on the testes of mammals Anim. Reprod. International Symposium on Animal Biology of Reproduction 3, 81-91.

Setchell BP, Bergh A, Widmark A, Damber JE, 1995. Effect of testicular temperature on vasomotion and blood flow. International Journal of Andrology 18, 120-126. [CrossRef]

Setchell BP, Locatelli A, Perreau C, Pisselet C, Fontaine I, Kuntz C, Hochereau-de Reviers MT, 1991. The form and function of the Leydig cells in hypophysectomized rams treated with pituitary extract when spermatogenesis is disrupted by heating the testes. The Journal of Endocrinology 131, 101-112. [CrossRef]

Setchell BP, Tao L, Zupp JL, 1996. The penetration of chromium-EDTA from blood plasma into various compartments of rat testes as an indicator of function of the blood-testis barrier after exposure of the testes to heat. Journal of Reproduction and Fertility 106, 125-133. [CrossRef]

Silva LKX, Sousa JS, Silva AOA, Lourenço Junior JB, Faturi C, Martorano LG, Garcia AR, 2017. Testicular thermoregulation, scrotal surface temperature patterns and semen quality of water buffalo bulls reared in a tropical climate. Andrologia e12836. [CrossRef]

Skinner JD, Louw GN, 1966. Heat stress and spermatogenesis in Bos indicus and Bos taurus cattle. Journal of Applied Physiology 21, 1784-1790. [CrossRef]

Souto PLG, McManus C, Zago FC, Martins E, Fonteque JH, Egito AA, Ramos AF, 2017. Reproductive characteristics of Crioulo Lageano breed bulls (Bos taurus) at puberty. Animal Reproduction 14, 1034- 1042. [CrossRef]

Stanfield C, Germann W, 2008. Principles of human physiology (Third). San Francisco, Calif: Benjamin Cummings.

Turner TT, D’Addario DA, Forrest JB, Howards SS, 1982. The Effects of Experimental Cryptorchidism on the Entry of [ 3 H]-Inulin and [ 3 H]-Horseradish Peroxidase into the lumen of the rat seminiferous tubules. Journal of Andrology 3, 178-183. [CrossRef]

Van Zelst SJ, Zupp JL, Hayman DL, Setchell BP, 1995. X-Y chromosome dissociation in mice and rats exposed to increased testicular or environmental temperatures. Reproduction, Fertility, and Development 7, 1117-1121. [CrossRef]

Vash PD, Engels TM, Kandeel FR, Greenway F, 2002. Scrotal cooling increases rectal temperature in man. Experimental Biology and Medicine (Maywood, N.J.) 227, 105-107. [CrossRef]

Vogler CJ, Bame JH, DeJarnette JM, McGilliard ML, Saacke RG, 1993. Effects of elevated testicular temperature on morphology characteristics of ejaculated spermatozoa in the bovine. Theriogenology 40, 1207-1219. [CrossRef]

Waites GMH, Voglmayr JK, 1962. Apocrine sweat glands of the scrotum of the ram. Nature 196, 965-967. [CrossRef]

Waites GMH, Voglmayr JK, 1963. The functional activity and control of the apocrine sweat glands of the scrotum of the ram. Australian Journal of Agricultural Research 14, 839. [CrossRef]

Wallage AL, Johnston SD, Lisle AT, Beard L, Lees AM, Collins CW, Gaughan JB, 2017. Thermoregulation of the bovine scrotum 1: measurements of free-range animals in a paddock and pen. International Journal of Biometeorology 61, 1381-1387. [CrossRef]

Wallin BG, Charkoudian N, 2007. Sympathetic neural control of integrated cardiovascular function: Insights from measurement of human sympathetic nerve activity. Muscle & Nerve 36, 595-614. [CrossRef]

Weiner JS, Hellmann K, 1960. The sweat glands. Biological Reviews 35, 141-186. [CrossRef]

Werdelin L, Nilsonne Å, 1999. The Evolution of the scrotum and testicular descent in mammals: a phylogenetic view. Journal of Theoretical Biology 196, 61-72. [CrossRef]

Whyte DG, Johnson AK, 2005. Lesions of the anteroventral third ventricle region (AV3V) disrupt cardiovascular responses to an elevation in core temperature. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology 288, 1783-1790. [CrossRef]

Wildeus S, Entwistle KW, 1983. Spermiogram and sperm reserves in hybrid Bos indicus X Bos taurus bulls after scrotal insulation. Journal of Reproduction and Fertility 69, 711-716. [CrossRef]

Wildeus S, Entwistle KW, 1986. Effects of scrotal insulation and unilateral vasoligation on ejaculate characteristics and sperm reserves in the bull. Animal Reproduction Science 10, 11-21. [CrossRef]

Yasui T, Fukui K, Nara T, Habata I, Meyer W, Tsukise A, 2007. Immunocytochemical localization of lysozyme and β-defensin in the apocrine glands of the equine scrotum. Archives of Dermatological Research 299, 393-397. [CrossRef]

Zhang W, Zhou Q, Chen W, Ni X, Zhao Z, Yang WZ, Zhou Q, 2017. A hypothalamic circuit that controls body temperature. Proceedings of the National Academy of Sciences 114, 1755. [CrossRef]

Kaynak Göster