Bruce Young, PhD


Work: 660.626.2491
Fax: 660.626.2346
Email Address:

Research Interests: Dr. Young's current research area is the neurobiology and biophysics of sensory systems, and particularly audition, in reptiles. Audition in this group is remarkably diverse, including: pressure-coupled tympani in some taxa; tympanic and non-tympanic ears; elaborate stapedial dampening systems; separate low-frequency and high-frequency auditory pathways; as well as integration with pressure and/or vibration detection systems. A second area of research is the neurobiology of venom secretion, storage, and activation. Snake venom is extremely diverse in terms of its biochemical and toxicological properties; perhaps the only common attribute is the proteolytic nature of these secretions. Though the system is poorly known, it appears that snakes secrete and store an inactive precursor to venom, then rapidly "activate" the venom by mixing it with other compounds (especially mineral ions). I am interested in the underlying neural control of this process, including such topics as: the neural pathway for venom activation, the relationship to salivation, and the pharmacological properties of the inactive precursors.


PhD - Biological Sciences (Anatomy), University of Calgary (1989)
Summer Course - Functional Morphology of Vertebrates at Leiden University, The Netherlands
MS - Biology, Northern Arizona University (1986)
BS - Zoology, Washington State University (1984)

Scholarly Activity

Research Publications:

2016 | Vedurmudi AP, Young BA, van Hemmen JL. Internally coupled ears: mathematical structures and mechanisms underlying ICE. Biol Cybern. 2016;110(4-5):359-382.

2016 | Nasoori A, Shahbazzadeh D, Tsubota T, Young BA. The defensive behaviour of Naja oxiana,with comments on the visual displays of cobras. Herpetological Bulletin. 2016

2016 | Jackson TNW, Young B, Underwood G, McCarthy CJ, Kochva E, Vidal N, van der Weerd L, Nabuurs R, Dobson J, Whitehead D, Vonk FJ, Hendrikx I, Hay C, Fry BG. (2016) Endless forms most beautiful: the evolution of ophidian oral glands, including the venom system, and the use of appropriate terminology for homologous structures. Zoomorphology, 1-24; DOI 10.1007/s00435-016-0332-9.

2016 | Young BA. Anatomical influences on internally coupled ears in reptiles. Biol Cybern. 2016;110(4-5):255-261

2016 | Young BA, Dumais J, John N, Lyons B, Macduff A, Most M, Reiser NA, Reiser PJ. Functional segregation within the muscles of aquatic propulsion in the Asiatic Water Monitor Lizard (Varanus salvator).Front Physiol. 2016;7:380

2016 | Han D, Young BA. Anatomical Basis of Dynamic Modulation of Tympanic Tension in the Water Monitor Lizard, Varanus salvator. Anat Rec (Hoboken). 2016 Jun 17;299(9):1270-1280. doi: 10.1002/ar.23382

2016 | Vedurmudi AP, Goulet J, Christensen-Dalsgaard J, Young BA, Williams R, van Hemmen JL. How Internally Coupled Ears Generate Temporal and Amplitude Cues for Sound Localization. Phys Rev Lett. 2016 Jan 15;116(2):028101

2014 | Bierman HS, Thornton JL, Jones HG, Koka K, Young BA, Brandt C, Christensen-Dalsgaard J, Carr CE, Tollin DJ. Biophysics of directional hearing in the American alligator (Alligator mississippiensis). J Exp Biol. 2014 Apr 1;217(Pt 7):1094-107. doi: 10.1242/jeb.092866

2014 | Most MG, Grace KM, Young BA. Terrestrial locomotion in the marine filesnake, Acrochordus granulatus (Schneider, 1799). Hamadryad 2014 36(2):142-8

2012 | Kohl T, Colayori SE, Westhoff G, Bakken GS, Young BA. Directional sensitivity in the thermal response of the facial pit in western diamondback rattlesnakes (Crotalus atrox). J Exp Biol. 2012 Aug 1;215(Pt 15):2630-6. doi:10.1242/jeb.065896

2012 | Fry BG, Casewell NR, Wüster W, Vidal N, Young B, Jackson TN. The structural and functional diversification of the Toxicofera reptile venom system. Toxicon. 2012 Sep 15;60(4):434-48. doi: 10.1016/j.toxicon.2012.02.013. Epub 2012 Mar 14

2012 | Burnell A, Collins S, Young BA. On the postpulmonary septum of Varanus and it’s implications for Mosasaurine ventilation and physiology. 2012 Bulletin of the Geological Society of France. 183: 159-169

2012 | Burnell A, Collins S, Young BA. Vertebral morphometrics in Varanus. Bulletin of the Geological Society of France. 2012 183: 151-158

2011 | Young BA, Herzog F, Friedel P, Rammensee S, Bausch A, van Hemmen JL. Tears of venom: hydrodynamics of reptilian envenomation. Phys Rev Lett. 2011 May 13;106(19):198103. Epub 2011 May 12

2010 | Lindgren J, Polcyn M, Young BA. Landlubbers to leviathans: modular evolution and swimming in mosasaurine mosasaurs. Paleobiology 2010 37: 445-469

2010 | Westhoff G, Boetig M, Bleckmann H, Young BA. Target tracking during venom 'spitting' by cobras. J Exp Biol. 2010 Jun 1;213(11):1797-802. doi:10.1242/jeb.037135

2010 | Young BA, Kardong KV. The functional morphology of hooding in cobras. J Exp Biol. 2010 May;213(Pt 9):1521-8. doi: 10.1242/jeb.034447

2010 | . Young BA. How a heavy-bodied snake strikes quickly: high-power axial musculature in the puff adder (Bitis arietans). J Exp Zool A Ecol Genet Physiol. 2010 Feb 1;313(2):114-21. doi: 10.1002/jez.579

2009 | Young,BA, Boetig M, Westhoff G. Spitting behaviour of hatchling red spitting cobras (Naja pallida). Herpetological Journal 2009 19: 185-191

2009 | Young BA, Boetig M, Westhoff G. Functional bases of the spatial dispersal of venom during cobra spitting. Physiol Biochem Zool. 2009 Jan-Feb;82(1):80-9.doi: 10.1086/595589

2008 | Young BA, Boetig M, Fahey A, Lawrence A. The diversity of aquatic locomotion in extant varanoid lizards. Proc. 2008 Fort Hayes St. Univ. 3: 159-167

2008 | Friedel P, Young BA, van Hemmen JL. Auditory localization of ground-borne vibrations in snakes. Phys Rev Lett. 2008 Feb 1;100(4):048701. Epub 2008 Jan 28

2008 | Fry BG, Scheib H, van der Weerd L, Young B, McNaughtan J, Ramjan SF, Vidal N, Poelmann RE, Norman JA. Evolution of an arsenal: structural and functional diversification of the venom system in the advanced snakes (Caenophidia). Mol Cell Proteomics. 2008 Feb;7(2):215-46. Epub 2007 Sep 12

2008 | Parker MR, Young BA, Kardong KV. The forked tongue and edge detection in snakes (Crotalus oreganus): an experimental test. J Comp Psychol. 2008 Feb;122(1):35-40. doi: 10.1037/0735-7036.122.1.35

2008 | Young BA, Aguiar A, Lillywhite H. Foraging cues used by Cottonmouth snakes (Agkistrodon piscivorus conanti) on Sea Horse Key, Florida. South Amer. J. Herpetol. 2008 3(2): 135-144

2007 | Young BA, Kardong KV. Mechanisms controlling venom expulsion in the western diamondback rattlesnake, Crotalus atrox. J Exp Zool A Ecol Genet Physiol. 2007 Jan 1;307(1):18-27

2006 | Young BA, Harris J. Auditory response of the Northern Pacific rattlesnake (Crotalus viridis oreganus): Does behavior match physiology? Northwest Sci. 2006 80:218-233

2005 | Young BA. Snake sounds. Nature Austr. 2005 28(4): 34-41

2005 | Young BA, O’Shea M. Analyses of venom spitting in African cobras (Elapidae: Serpentes). Afr. Zool. 2005 40(1): 71-76.

2004 | Young BA, Dunlap K, Koenig K, Singer M. The buccal buckle: the functional morphology of venom spitting in cobras. J Exp Biol. 2004 Sep;207(Pt 20):3483-94

2004 | Grande T, Young BA. The ontogeny of the Weberian apparatus in the zebrafish Danio rerio (Ostariophysi: Cypriniformes) with comments on the homology of otophysan Weberian ossicles. Zool. J. Linn. Soc. 2004 140: 241-254

2003 | Hartdegen RW, Russell MJ, Young BA, Reams RD. Krokotiiliskinkin Tribolonotus gracilis (DE ROOY, 1909) ääntely ja todisteita poikasten hoidosta. Herpetomania 2003 12(4): 5-9

2003 | Young BA. Snake bioacoustics: toward a richer understanding of the behavioral ecology of snakes. Q Rev Biol. 2003 Sep;78(3):303-25. Review

2003 | oung BA, Morain M. The mechanics of vertical burrowing in the Saharan sand vipers (Cerastes). Copeia 2003: 132-138

2003 | Young, BA, Phelan M, Morain M, Ommundsen M, Kurt R. Venom injection in rattlesnakes (Crotalus): Peripheral resistance and the pressure-balance hypothesis. Can. J. Zool. 2003 81: 313-320

2002 | Young BA, Aguiar A. Response of western diamondback rattlesnakes Crotalus atrox to airborne sounds. J Exp Biol. 2002 Oct;205(Pt 19):3087-92

2002 | Young BA, Lee CE, Daley KM. Do snakes meter venom? Biosciences 2002 52:1121-1126

2002 | Young BA, Lee CE, Daley KM. On a flap and a foot: Aerial locomotion in the Flying gecko, Ptychozoon kuhli. J. Herpetol. 2002 36(3): 412-418

2002 | Young BA, Morain M. The use of ground-borne vibrations for prey localization in the Saharan sand vipers (Cerastes). J Exp Biol. 2002 Mar;205(Pt 5):661-5

2001 | Berkhoudt H, Wilson P, Young BA. Taste buds in the palatal mucosa of snakes. Afr. Zool. 2001 36(2): 185-188

2001 | Hartdegen RW, Russell MJ, Young BA, Reams RD. Vocalization of the Crocodile skink, Tribolonotus gracilis, and evidence of parental care. Cont. Herpetol. 2001(on-line publication)

2001 | Blair M, Zahn K, Marvin J. Mechanics of venom expulsion in Crotalus,with special reference to the role of the fang sheath. Anat Rec. 2001 Dec 1;264(4):415-26

2001 | Young BA, Jaggers J, Nejman N, Kley N. Defensive buccal expansion in Bitis arietans. Copeia 2001(1): 270-273

2001 | Young BA, Phelan M, Jaggers J, Nejman N. Kinematic modulation of the strike of the western diamondback rattlesnakes (Crotalus atrox). Hamadryad 2001 26(2): 316-349

2001 | Young BA, Zahn K. Venom flow in rattlesnakes: mechanics and metering. J Exp Biol. 2001 Dec;204(Pt 24):4345-51

2000 | Young BA. The comparative morphology of the ophidian larynx. Acta Zool. 2000 81:177-193

2000 | Young BA, Marvin J, Marosi K. The potential significance of groundborne vibration to predator-prey relationships in snakes. Hamadryad 2000 25(2): 164-174

2000 | Young BA, Nejman N, Jaggers J. Snakes that inflate: The functional morphology of a defensive behavior. Zoology 2000 103: 25-30

2000 | Young BA, Zahn K, Blair M, Lalor J. Functional subdivision of the venom gland musculature and the regulation of venom expulsion in rattlesnakes. J Morphol. 2000 Dec;246(3):249-59

1999 | Cascardi J, Young BA, Husic HD, Sherma J. Protein variation in the venom spat by the red spitting cobra, Naja pallida (Reptilia: Serpentes). Toxicon. 1999 Sep;37(9):1271-9

1999 | Young BA. The comparative morphology of the mandibular midline raphe in snakes (Reptilia: Squamata). Zool. Anz. 1999 237: 217-241

1999 | Young BA, Lalor J, Solomon J. The comparative biomechanics of an ophidian defensive behavior: Head triangulation in hognose snake (Heterodon) and egg-eating snake (Dasypeltis). J. Zool. Lond. 1999 248: 169-177

1999 | Young BA, Marsit C, Meltzer K. Comparative morphology of the cloacal scent gland in snakes (Serpentes: Reptilia). Anat Rec. 1999 Oct 1;256(2):127-38

1999 | Young BA, Meltzer K, Marsit C. Scratching the surface of mimicry: Sound production through scale abrasion in snakes. Hamadryad 1999 24: 29-38

1999 | Young BA, Meltzer K, Marsit C, Abishahin G. Cloacal popping in snakes. J. Herpetol. 1999 33: 557-566

1999 | Young BA, Nejman N, Meltzer K, Marvin J. The mechanics of sound production in the puff adder bitis arietans (Serpentes: viperidae) and the information content of the snake hiss. J Exp Biol. 1999 Sep;202 (Pt 17):2281-9

1999 | Young BA, Saunders M. Direct visualization of blood flow through the interaortic foramen of the eastern diamondback rattlesnake, Crotalus adamanteus, using echocardiography and color Doppler imaging. J Exp Zool. 1999 Dec 1;284(7):742-745

1999 | Young BA, Solomon J, Abishahin G. How many ways can a snake growl? The morphology of sound production in Ptyas mucosus and its potential mimicry of Ophiophagus. Herpetol. J. 1999 9(3): 89-94

1998 | Kinney C, Abishahin G, Young BA. Hissing in rattlesnakes: Redundant signaling or inflationary epiphenomenon? J. Exp. Zool. 1998 280: 107-113

1998 | Young BA. How to hiss loudly: Narial flaring and acoustic amplification in Russell’s viper (Daboia russelii). Hamadryad 1998 23(2): 103-110

1998 | Young BA. On the acoustic biology of snakes. [Invited paper] Lit. Serpent. 1998 18(4): 84-91

1998 | Young BA. The comparative morphology of the intermandibular connective tissue in snakes (Reptilia: Squamata). Zool. Anz. 1998 237: 59-84

1998 | Young BA, Abishahin G, Bruther M, Kinney C, Sgroi J. Acoustic analysis of the defensive sounds of Varanus salvator with notes on sound production in other varanid species. Hamadryad 1998 23: 1-14

1998 | Young B, Frazer B, Fried B, Lee M, Lalor J, Sherma J. HPTLC determination of cloacal scent gland lipids from two sympatric snakes: The eastern diamondback rattlesnake (Crotalus adamanteus) and the Florida cottonmouth (Agkistrodon piscivorus conanti). J.P. C. 1998 12: 196-201

1998 | Young B, Lalor J. Sound production in the eastern hognose snake, Heterodon platyrhinos (Serpentes: Colubridae): Does it snore? Amph.-Rept. 1998 19:407-418

1998 | Young BA, Wallach V. Description of a papillate tactile organ in the Typhlopidae. S. Afr. J. Zool. 1998 33: 249-253

1997 | Grande T, Young B. Morphological development of the opercular apparatus in Kneria wittei (Ostariophysi: Gonorynchiformes) with comments on its possible function. Acta Zool. 1997 78: 145-162

1997 | Young BA. A review of sound production and hearing in snakes, with a discussion of intraspecific acoustic communication in snakes. J. Penn. Acad. Sci. 1997 71: 39-46

1997 | Young BA. On the absence of taste buds in monitor lizards (Varanus) and snakes. J. Herpetol. 1997 31: 130-137

1997 | Young BA, Wassersug RJ, Pinder A. Gravitational gradients and blood flow patterns in specialized arboreal (Ahaetulla nasuta) and terrestrial (Crotalus adamanteus) snakes. J Comp Physiol B. 1997;167:481-93

1996 | Young BA, Kardong KV. Dentitional surface features in snakes (Lepidosauria: Serpentes). Amph.-Rept. 1996 17: 261-276

1995 | Rasmussen S, Young BA, and Krimm H. On the spitting behavior in cobras. J. Zool. Lond. 1995 237(1): 27-36

1995 | Young BA, Brown IP. The physical basis of the rattling sound in the rattlesnake, Crotalus viridis oreganus. J. Herpetol. 1995 29(1): 80-85

1995 | Young BA, Sheft S, Yost W. Sound production in Pituophis melanoleucus (Serpentes: Colubridae) with the first description of a vocal cord in snakes. J Exp Zool. 1995 Dec 15;273(6):472-81

1994 | Young BA. Cartilago cordis in serpents. Anat Rec. 1994 Oct;240(2):243-7

1994 | Young BA, Street SL, Wassersug RJ. Anatomical and gravitational influences on cardiac displacement in snakes (Lepidosauria, Serpentes). Zoomorphology.1994;114:169-75

1993 | Young BA, Brown IP. On the acoustic profile of the rattlesnake rattle. Amph.-Rept. 1993 14: 373-380

1993 | Young BA, Lillywhite HB, Wassersug RJ. On the structure of the aortic valves in snakes (Reptilia: Serpentes). J. Morphol. 1993 216(2): 141-160

1992 | Young BA. On the necessity of an archetypal concept in morphology: With special reference to the concepts of structure and homology. Biol. and Phil. 1992 8:225-248

1992 | Young BA. Tracheal diverticula in snakes: Possible functions and evolutionary origins. J. Zool. Lond. 1992 227: 567-583

1991 | Young BA. Evaluating hypotheses for the transfer of stimulus particles to Jacobson’s organ in snakes. [invited paper] Brain Behav. and Evol. 1991 41: 203-209

1991 | Young BA. Morphological basis of growling in the king cobra, Ophiophagus hannah. J Exp Zool. 1991 Dec;260(3):275-87

1990 | Young BA. Is there a direct link between the ophidian tongue and Jacobson’s organ? Amph.-Rept. 1990 11: 263-276

1990 | Young BA. The influences of the aquatic medium on the prey capture system of snakes. J. Nat. Hist. 1990 25: 519-531

1990 | Young BA, Magon DK, Goslow GE. Length-tension and histochemical properties of selected shoulder muscles of the savannah monitor lizard (Varanus exanthematicus): Implications for function and evolution. J. Exp. Zool. 1990 256: 63-74

1989 | Young BA. Ontogenetic changes in the feeding system of the red-sided garter snake, Thamnophis sirtalis parietalis. I. Allometric analysis. J. Zool. Lond. 1989 218(3): 365-382

1989 | Young BA. The anatomy of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Part I - Osteology. Zool. Jb. Anat. 1989 118: 251-272

1989 | Young BA. The anatomy of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Part II - Myology. Zool. Jb. Anat. 1989 118: 325-354

1989 | Young BA. Ashley MA. A mathematical model of relative muscle force with a perturbation analysis of selected muscle parameters. J. Theor. Biol. 1989 138:213-233

1988 | Young BA. A model for the evolution of functional-morphological systems. Evol. Theory. 1988 8(3): 125-150

1988 | Young BA. The arthrology of the head of the red-sided garter snake, Thamnophis sirtalis parietalis. Neth. J. Zool. 1988 38(2-4): 166-205

1988 | Young BA. The cephalic vascular anatomy of three species of sea snakes. J Morphol. 1988 May;196(2):195-204

1988 | Young BA. The subclavian loop of Varanus salvator. Copeia. 1988(4): 1029-1034

1988 | Young BA. The cephalic vascular anatomy of three species of sea snakes. J. Morphol. 1988 196:195-204

1987 | Young BA. The comparative cephalic osteology and myology of three species of sea snakes. Zool. Jb. Anat. 1987 116: 421-452

1987 | Young BA. The cranial nerves of three species of sea snakes. Can J. Zool. 1987 65(9):2236-2240

2016 | Han D, Young BA. Biophysics of the Varanid Tympanic Membrane. Poster presented at the 8th Annual Interdisciplinary Biomedical Research Conference; Kirksville, MO; November 5, 2016

2016 | Nasoori A, Young BA, Tsubata T, Shahbazzadeh D. The defensive behavior of Naja oxiana with comments on the visual displays of cobras. Poster presented at the 3rd International Veterinary Congress, London, August 18-20, 2016.

2015 | Young BA. Active Tuning Mechanisms in the Reptile Ear. Invited oral presentation at the Symposium on Biomimetics of Sensory System and Locomotion; The Second Sino-German Symposium on Biomimetics: From Animal Sensory Systems to Locomotion; Garching; Germany; June 29, 2015

2014 | Young BA, Most M, Dumais J, John N, Lyons B, MaCduff A, Reiser P. Multiple Perspectives of the Functional Divisions within Swimming Muscles of the Asiatic Water Monitor (Varanus salvator). Oral presentation at the Society for Integrative and Comparative Biology; Austin, TX; Jan. 3-7, 2014

Books, Chapters, and Reviews:

2015 | Young BA. 2015. The evolution of the snake ear and its restricted frequency response range. In: All animals are interesting, A Festschrift in honour of Anthony P. Russell. (O.R.P. Bininda-Edwards, G. Powell, H. Jamniczky, and A. Bauer, eds.) Universitat Oldensburg Press. pp. 423-438

2015 | Fry B, Sunagar K, Casewell N, Kochva E, Scheib H, Wüster W, Vidal N, Young B, Burbrink F, Pyron A, Vonk F, Jackson, T. 2015. The origin and evolution of the Toxicofera reptile venom system.In B. G. Fry (Ed.). Venomous Reptiles and Their Toxins. Evolution, Pathophysiology and Biodiscovery. Oxford University Press , 2015. p. 1-576

2014 | Young BA, Mathevon N, Tang Y. 2014. Reptile auditory neuroethology: What do reptiles do with their hearing. In: Insights from Comparative Hearing Research, Springer Handbook for Auditory Research (R. Fay, C. Koppel, G. Manley, A. Popper, eds.) Springer, New York. 49: 323 – 346

2010 | Young BA. 2010. Vibration detection in snakes. Pp. 85 – 98 In: The Use of Vibrations in Communication: properties, mechanisms and function across taxa. (O’Connell-Rodwell C.E., ed.) Research Signpost

2008 | Young BA. 2008. Perspectives on the regulation of venom expulsion in snakes. Pp. 181 – 190 In: The Biology of Rattlesnakes, W. Hayes, M. Cardwell, K. Beaman, and S. Bush, eds., Loma Linda University Press

2008 | Young BA, Jackson K. 2008. Functional specializations of the extrinsic venom gland musculature in crotaline snakes: What is the role of the M. Pterygoideus Glandulae?;Pp. 47 – 54 In: The Biology of Rattlesnakes, W. Hayes, M.Cardwell, K. Beaman, and S. Bush, eds., Loma Linda University Press

2007 | Young BA. 2007. Response of the yellow anaconda (Eunectes notaeus) to underwater pressure waves. Pp. 199 – 206 In: Biology of Boas and Pythons, R. Henderson,Ed., Eagle Mountain Press

2006 | Young BA. 2006. Auditory atavism and integrated pathways for hearing in snakes. Pp. 229 – 232 in: Herpetological Bonnensis II (M. Vences, J. Köhler, T. Ziegler, W. Böhme, eds.) European Society of Herpetology, Bonn

1997 | Young BA. 1997. Hearing, taste, tactile reception, and olfaction. Pp. 195 – 223 In: The Biology, Husbandry, and Health Care of Reptiles, Vol. 1. (Ackerman, L., ed.), T.F.H. Publications, New Jersey