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Daniel Tollin, PhD


Department of Physiology and Biophysics
University of Colorado School of Medicine

Joint Appointment: Department of Otolaryngology

Physiology and Biophysics
Biomedical Sciences​


Behavioral and physiological mechanisms of binaural and spatial hearing


The goal of the work in my lab is to understand the neural mechanisms of auditory perception with particular emphasis on how sources of sounds are localized.  Because the peripheral receptors of the ear have no mechanism to directly sense sound location on their own (unlike the topographic organization of the retina), location must be computed at more central levels.  This makes sound localization a fascinating neurocomputational problem, particularly from a developmental perspective.  Our experiments seek answers to at least four basic questions:

  1. What are the acoustical cues to sound source location and what are their physical bases?
  2. How are the acoustical cues represented and transformed in the activity patterns of neurons in the various nuclei of the ascending auditory pathway?
  3. How are the neural representations of the cues used by observers to determine location?
  4. How do each of these aspects of hearing develop?

I use a multidisciplinary approach to tackle these questions employing both experimental and theoretical techniques including human and animal psychophysics, extracellular physiology, signal detection and information theory, systems identification techniques, acoustic transfer function measurement and modeling, digital filter design and estimation, acoustic signal design, and physiological systems modeling.

Fig. 1
Illustration of a frontal section through the brainstem showing the ascending pathways through the nuclei of the superior olivary complex that are believed to be responsible for encoding interaural level differences (ILDs).  Neurons of the lateral superior olive (LSO) receive bilateral inputs from both ears.  The input from the ipsilateral ear via the spherical bushy cells (SBCs) is excitatory (open symbols) but the input from the contralateral ear via the globular bushy cells (GBCs) of the contralateral anteroventral cochlear nucleus (AVCN) is inhibitory (filled symbols) due to the additional synapse in the ipsilateral medial nucleus of the trapezoid body (MNTB).  The interplay of the ipsilateral excitation and contralateral inhibition confers on LSO neurons sensitivity to ILDs.  LSO neurons send excitatory projections to the contralateral inferior colliculus (IC) and dorsal nucleus of the lateral lemniscus (DNLL) and inhibitory projections (not shown) to the ipsilateral IC and DNLL.  The color bar and shading indicates the tonotopic organization and shows that the neurons comprising the MNTB and LSO are sensitive to predominantly high frequency sounds.

​Image coming soon
John Peacock, PhD
Research Associate
Linked in headshot Carol.jpg

​​Carol Sammeth
Research Associate​
​​​​Image coming soon Monica Benson
Graduate Student
​​Image coming soon Zoe Owrutsky
Graduate Student


Selected Publications

Brown, AD, Beemer BT, Greene NT, Argo T, Meegan GD and Tollin DJ (2015)  Effects of active and passive hearing protection devices on sound source localization, speech recognition, and tone detection, PLoS One Aug 27;10(8):e0136568. doi: 10.1371/journal.pone.0136568
Jones HG, Brown AD, Koka K, Thornton JL and Tollin DJ (2015) Sound frequency-invariant neural coding of a frequency-dependent cue to sound source location. J Neurophysiol 114:531-539.
Brown AD, Stecker GC and Tollin DJ (2015) The precedence effect in sound localization, J. Assoc. Res. Otolaryngol (JARO), 16:1-28.
Greene NT, Mattingly JK, Jenkins HA, Tollin DJ, Easter JR and Cass SP (2015) Cochlear Implant Electrode Effect on Sound Energy Transfer within the Cochlea during Acoustic Stimulation, Otology & Neurotology 36:1554-1561.
Mattingly JK, Greene NT, Jenkins HA, Tollin DJ, Easter JR and Cass SP (2015) Effects of Skin Thickness on Cochlear Input Signal using Transcutaneous Bone Conduction Implants, Otology & Neurotology 36:1403-1411.
Beutelmann R, Laumen G, Tollin DJ and Klump GM (2015) Amplitude and phase equalization of stimuli for click evoked auditory brainstem responses, J. Acoust. Soc. Am. 137: 71-77.
Koka K and Tollin DJ (2014). Linear coding of complex sound spectra by discharge rate by neurons of the medial nucleus of the trapezoid body (MNTB) and it inputs. Front. Neural Circuits 8:144. doi:10.3389/fncir.2014.00144
Greene NT, Anbuhl KL, Williams W and Tollin DJ (2014) The acoustical cues to sound location in the Guinea pig (cavia porcellus), Hearing Research 316:1-15.
Tringali S, Koka K, Jenkins HA and Tollin DJ (2014) Sound Location Modulation of Electrocochleographic Responses in Chinchilla with Single Sided Deafness and Fitted with an Osseointegrated Bone-conducting Hearing Prosthesis, Otol & Neurotol 36:678-686.
Bierman HS, Thornton JL, Jones HG, Koka K, Young BA, Carr CE and Tollin DJ (2014) Biophysics of directional hearing in the American Alligator (Alligator mississippiensis), J Exp Biol 217:1094-1107.  (Top 5 most read article in the Journal of Experimental Biology in 2014)

Lupo JE, Koka K, Jenkins HA, and Tollin DJ (2014) Vibromechanical Assessment of Active Middle Ear Implant Stimulation in Simulated Middle Ear Effusion: A Temporal Bone Study, Otology and Neurotology, 35:470-475
Gai Y, Ruhland JL, Yin TCT and Tollin DJ (2013) Behavioral and Modeling Studies of Sound Localization in Cats: Effects of Stimulus Level and Duration, J. Neurophysiol., 110:607-620. PMC3742992
Ruhland JL,Yin TCT and Tollin DJ (2013) Gaze shifts of the cat to auditory and visual stimuli, J. Assoc. Res. Otolaryngol (JARO), 14:731-755.  PMC3767881
Thornton JL, Chevallier KM, Koka K and Tollin DJ (2013) Conductive hearing loss induced by experimental middle ear effusion in a chinchilla model reveals impaired TM-coupled ossicular chain movement, J. Assoc. Res. Otolaryngol (JARO), 14:451-464.  PMC3705087
Tollin DJ, Ruhland JL and Yin TCT (2013) The role of spectral composition of sounds on the localization of sound sources by cats, J. Neurophysiol. 109:1658-1668.  PMC3602938
Dondzillo, A., Thornton, J.L., Tollin, D.J., Klug, A. (2013) Manufacturing and Using Piggy-back Multibarrel Electrodes for In vivo Recordings. J Vis Exp (71) e4358 DOI:10.3791/4358. PMC3582659

Deveze A, Koka K, Tringali S, Jenkins HA, and Tollin DJ (2012). Techniques to improve the performance of a middle ear implant: Effect of different methods of coupling to the ossicular chain, Otology & Neurotology, 34:158-166.

Thornton JL, Chevallier KM, Koka K, Lupo JE and Tollin DJ (2012) The conductive hearing loss due to an experimentally-induced middle ear effusion alters the interaural level and time difference cues to sound location, J Assoc Res Otolaryngol, 13:641-654.  (Journal cover figure).

Lupo JE, Koka K, Hyde BJ, Jenkins HA, and Tollin DJ (2012) Third window vibroplasty with an active middle ear implant: Assessment of physiological responses in a model of stapes fixation in Chinchilla Lanigera, Otology and Neurotology, 33:425-431.

Jones HG, Koka K and Tollin DJ (2011) Postnatal development of cochlear microphonic and compound action potentials in a precocious species, chinchilla lanigera, J Acoust Soc Am 130:38-43.

Lupo JE, Koka K, Hyde BJ, Jenkins HA, and Tollin DJ (2011) Physiologic assessment of active middle ear implant coupling to the round window in Chinchilla Lanigera, Otolaryngology - Head and Neck Surgery, 145:641-647

Jones H, Koka K, Thornton J and Tollin DJ (2011). Concurrent development of the head and pinnae and the acoustical cues to sound location in a precocious species, the Chinchilla (Chinchilla laniger), J Assoc Res Otolaryngol., 12:127-140.

Koka K, Jones H, Thornton JL and Tollin DJ (2011) The acoustical cues to sound location in the adult Chinchilla: measurements of Directional Transfer Functions (DTFs), Hear Res, 272:135-147.

Lupo JE, Koka K, Thornton JL and Tollin DJ (2011) The effects of experimentally induced conductive hearing loss on the spectral and temporal aspects of sound transmission through the ear, Hear Res 272:30-41.

Tringali S, Koka K, Deveze A, Ferber AT, Jenkins HA, and Tollin DJ (2011). Intraoperative adjustments to optimize active middle ear implant performance, Acta-Otolaryngologica, 131:27-35.

Tollin DJ (2010). The development of sound localization mechanisms, Oxford Handbook of Developmental Behavioral Neuroscience, MS Blumberg, JH Freeman, and SR Robinson (Eds), Oxford University Press, p. 262-282. [PDF]

Deveze A, Koka K, Tringali S, Jenkins HA, and Tollin DJ (2010). Active middle ear implant application in case of stapes fixation: A temporal bone study, Otology & Neurotology, 31:1027-1034.

Oliva AM, Salcedo E, Hellier J, Koka K, Tollin DJ, Restrepo D (2010). Toward a mouse neuroethology in the laboratory environment, PLoS ONE 5(6): e11359. doi:10.1371/journal.pone.0011359.

Tsai JJ, Koka K and Tollin DJ (2010). Roving overall sound intensity to the two ears impacts interaural level difference discrimination thresholds by single neurons in the lateral superior olive, J. Neurophysiol. 103:875-886.

​Tollin DJ, McClaine L, and Yin TCT (2010). Short-latency, goal-directed movements of the pinnae to sounds that produce auditory spatial illusions, J Neurophysiol 103:446-457.

Tringali S, Koka K, Deveze A, Holland NH, Jenkins HA, and Tollin DJ (2010). Round window membrane implantation with an active middle ear implant: Effects on performance of round window exposure and transducer tip diameter, Audiology & Neurotology 15:291-302.

Tollin DJ and Koka K (2009).  Postnatal development of sound pressure transformations by the head and pinnae of the cat: binaural characteristics, J. Acoust. Soc. Am. 126:3125-3136.

Lupo JE, Koka K, Holland NJ, Jenkins HA, and Tollin DJ (2009) Prospective Electrophysiologic Findings of Round Window Stimulation in a Model of Experimentally-induced Stapes Fixation, Otology & Neurotology, 30:1215-1224.

Koka K, Holland NJ, Lupo JE, Jenkins HA, and Tollin DJ (2009). Electrocochleographic and mechanical assessment of round window stimulation with an active middle ear prosthesis, Hearing Research. doi:10.1016/j.heares.2009.08.009.

Al-Sheikh Hussein B, Matin M, and Tollin DJ (2009). All-pole and all-zero models of human and cat head related transfer functions, edited by Mark S. Schmalz, Gerhard X. Ritter, Junior Barrera, Jaakko T. Astola, Franklin T. Luk, Proc. of SPIE Vol. 7444, 74440X1-11, SPIE doi: 10.1117/12.829872.

Tollin DJ and Koka K (2009).  Postnatal development of sound pressure transformations by the head and pinnae of the cat: monaural characteristics, J. Acoust. Soc. Am. 125:980-994.

Tollin DJ, Ruhland JL, and Yin TCT (2009).  The vestibulo-auricular reflex, J. Neurophysiol. 101: 1258-1266.

Dent ML, Tollin DJ, and Yin TCT (2009).  The influence of sound source location on the behavior and physiology of the precedence effect, J. Neurophysiol, 102:724-734.

Tollin DJ, Koka K, and Tsai JJ (2008). Interaural level difference discrimination thresholds for single neurons in the lateral superior olive, J. Neurosci. 28:4848-4860.

Koka K, Read HE, and Tollin DJ (2008).  The acoustical cues to sound location in the rat: measurements of Directional Transfer Functions, J. Acoust. Soc. Am. 123:4297-4309.

Moore J, Tollin DJ, and Yin TCT (2008).  Can measures of sound localization acuity be related to the precision of absolute location estimates? Hear. Res. 238:94-109.

Tollin DJ and Yin TCT (2005). Interaural phase and level difference sensitivity in low-frequency neurons in the lateral superior olive, J. Neurosci. 25:10648-10657.

Tollin DJ (2003).  The lateral superior olive: A functional role in sound source localization, Neuroscientist, 9(2): 127-143. [PDF] 

Tollin DJ (1998). Computational model of the lateralization of clicks and their echoes, in S. Greenberg and M. Slaney (Eds.), Proceedings of the NATO Advanced Study Institute on Computational Hearing, pp. 77-82. [PDF]​

Greene NT, Jenkins HA, Tollin DJ, and Easter JR (2017) Stapes displacement and intracochlear pressure in response to very high level, low frequency sounds, Hear Res 348:16-30.

Brown, AD and Tollin DJ (2016) Slow temporal integration enables robust neural coding and perception of a cue to sound source location, Journal of Neuroscience 36:9908-9921.


Brown, AD and Tollin DJ (2016) Time-varying distortions of binaural information by bilateral hearing aids: Effects of nonlinear frequency compression, Trends in Hearing 20:1-15.


Ashida G, Kretzberg J, Tollin DJ (2016) Roles for coincidence detection in coding amplitude-modulated sounds. PLoS Comput Biol 12 (6): e1004997. doi:10.1371/journal.pcbi.1004997


Banakis-Hartl RM, Mattingly JK, Greene NT, Jenkins HA, Cass SP and Tollin DJ (2016) A preliminary investigation of the air-bone gap changes in intracochlear sound pressure with air- and bone-conducted stimuli after cochlear implantation, Otology & Neurotology 37:1291-1299.


Greene NT, Mattingly JK, Banakis-Hartl RM, Tollin DJ, and Cass SP (2016)Intracochlear pressure transients during cochlear implant electrode insertion, Otology & Neurotology 37:1541-1548.


Laumen G, Ferber AT, Klump GM and Tollin DJ (2016).  The physiological basis and clinical use of the binaural interaction component of the auditory brainstem response, Ear & Hearing 37:276–290


Ferber AT, Benichoux, V and Tollin DJ (2016).  Test-retest reliability of the binaural interaction component of the auditory brainstem response, Ear & Hearing 37:291–301


Laumen G, Tollin DJ, Beutelmann R and Klump GM (2016).  Aging effects on the binaural interaction component of the auditory brainstem response in the mongolian gerbil: effects of interaural time and level differences, Hear Res 337:46-58.​

PubMed search (Tollin D)