Our Research

For a complete list publications out of the Schofield lab, please take a look at our NCBI bibliography. You can also read about our techniques, our funding, and our collaborators.

Journal Covers

  • Cover of The Journal of Comparative Neurology, Volume 375, Number 3, yellow background with a large microscope image of large blue cells surrounded by smaller red label
  • Cover of The Journal of Comparative Neurology, Volume 453, Number 3, yellow background with a large microscope image of red and green labeled neurons
  • Cover for Hearing Research, Volume 191, Issues 1 and 2. Published May 2004. Purple background and showing digital photomicrographs of labeled cells in the IC.
  • Cover of Journal of the Association for Reserach in Otolaryngology, 7:1, March 2006, black background with closeup microscope image of a red-labeled neuron and its dendrites
  • Cover of Hearing Research, Volume 388, March 2020, featuring a microscope picture of magenta-labeled neurons in the LSO region
  • Cover of Hearing Research, Volume 411, November 2021. Green dots, squares, and lines represent cholinergic boutons, soma, and axons throughout various regions of two illustrated brain slices.
  • Cover of Volume 131, Sept 2023, Journal of CHEMICAL NEUROANATOMY, yellow background with a figure panel of neurons labeled red "Retro from CN," neurons labeled blue "Ach," and the merged image showing overlap
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Schofield, B.R. and Cant, N.B. (1996), Origins and targets of commissural connections between the cochlear nuclei in guinea pigs. J. Comp. Neurol., 375: 128-146.
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Schofield, B.R. (2002), Ascending and descending projections from the superior olivary complex in guinea pigs: Different cells project to the cochlear nucleus and the inferior colliculus. J. Comp. Neurol., 453: 217-225.
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Diana L. Coomes, Brett R. Schofield,
Separate projections from the inferior colliculus to the cochlear nucleus and thalamus in guinea pigs, Hearing Research, Volume 191, Issues 1–2, 2004, Pages 67-78,
ISSN 0378-5955
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Schofield, B.R., Coomes, D.L. & Schofield, R.M. Cells in Auditory Cortex that Project to the Cochlear Nucleus in Guinea Pigs. JARO 7, 95–109 (2006).
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Beebe, et al. (2020) Generation of a ChATCre mouse line without the early onset hearing loss typical of the C57BL/6J strain. Hear Res 388:107896
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Noftz, et al. (2021) Dense cholinergic projections to auditory and multisensory nuclei of the intercollicular midbrain. Hear Res, 411: 108352
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Beebe, et al. (2023) Characterization of three cholinergic inputs to the cochlear nucleus. J Chem Neuroanat, 131: 102284

Current Projects

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Above: A retrograde viral vector labels cells in the auditory cortex that make descending projections to the inferior colliculus

Descending Auditory Pathways: A major advance over the past 25 years has been the discovery of extensive pathways from auditory cortex to many lower brainstem auditory regions. Many of our studies have focused on identifying the auditory circuits that are contacted directly by the descending projections from auditory cortex. Another important focus of this work is the interaction between descending auditory pathways and neuromodulatory pathways.

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Above: Cholinergic axons in the inferior colliculus are labeled with a chemically-selective viral tract tracing technique.

Cholinergic Circuits: Acetylcholine is a neurotransmitter that is important in normal hearing, development, aging, learning, and after damage to the nervous system. Understanding these functions is critical for developing approaches to help people with age-related hearing loss, difficulty understanding speech in noisy environments, learning to hear with cochlear implants or coping with hearing dysfunctions in disorders such as autism or schizophrenia. Our work will answer critical questions about the circuitry that underlies cholinergic function in the brainstem.

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Above: A GABAergic cell (magenta) in the rostral pole of the inferior colliculus is surrounded by glutamatergic boutons (cyan) and a perineuronal net (green)

Types of Inhibitory Cells: Inhibitory neurons that use GABA or glycine as a neurotransmitter play key roles throughout the auditory system. Another area of emphasis in our lab focuses on several questions about these inhibitory cells: 1) are there specific molecular markers that distinguish subtypes of inhibitory cells that serve different functions in hearing? 2) do top-down circuits have direct connections with the inhibitory cells? 3) do cholinergic circuits have direct connections with the inhibitory cells?

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Above: A cell expressing neuropeptide Y (green) also projects to the medial geniculate nucleus (magenta)

Circuitry of Subclasses of Inferior Colliculus Cells: Recent collaborative work with the Roberts lab has shown that two markers, vasoactive intestinal peptide, or VIP, and neuropeptide Y, or NPY, can be used to identify distinct subclasses of cells in the inferior colliculus, an auditory hub. We are working to understand what brain regions send inputs to these cell types, and to find out what brain regions these cell types send projections to.

Schofield Lab Publications

Below are images from our recent publications. Click an image to see the full paper.

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Beebe, Silveira, et al. (2022) Neurotransmitter phenotype and axonal projection patterns of VIP-expressing neurons in the inferior colliculus. J Chem Neuroanat, 126:102189
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Beebe, et al. (2021) Cholinergic boutons are closely associated with excitatory and four subtypes of inhibitory cells in the inferior colliculus. J Chem Neuroanat, 116:101998
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Beebe, Zhang, et al. (2021) Multiple sources of cholinergic input to the superior olivary complex. Front Neural Circuits 15:715369
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Noftz, et al. (2020) Cholinergic projections from the pedunculopontine tegmental nucleus contact excitatory and inhibitory neurons in the inferior colliculus. Front Neural Circuits 14:43
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Beebe, et al. (2018) Inhibitory projections from the inferior colliculus to the medial geniculate body originate from four subtypes of GABAergic cells. eNeuro 5:ENEURO.0406-18.2018
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Crish and Schofield (2018) Anterograde tract tracing for assaying axonopathy and transport deficits in glaucoma. Methods Mol Biol 1695:171
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Mellott, et al. (2014) Excitatory and inhibitory projections in parallel from the inferior colliculus to the auditory thalamus. Front Neuroanat 8:124
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Schofield, et al. (2014) Subcollicular projections to the auditory thalamus and collateral projections to the inferior colliculus. Front Neuroanat 8:70
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Foster, et al. (2014) Perineuronal nets and GABAergic cells in the inferior colliculus of guinea pigs. Front Neuroanat 7:53
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Anair, et al. (2022) Inhibitory NPY neurons provide a large and heterotopic commissural projection in the inferior colliculus. Front Neural Circuits 16: 8711924
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Kristaponyte, et al. (2021) Group II metabotropic glutamate receptor modulate sound evoked and spontaneous activity in the mouse inferior colliculus. eNeuro 8 (1) ENEURO.0328-20.2020
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Silveira, et al. (2020) Neuropeptide Y expression defines a novel class of GABAergic projection neuron in the inferior colliculus. J Neurosci 40:4685
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Schofield and Beebe (2019) Subtypes of GABAergic cells in the inferior colliculus. Hear Res 376:1
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Mellott, et al. (2018) GABAergic and nonGABAergic projections to the superior colliculus from the auditory brainstem. Brain Struct Funct 223:1923
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Smith, et al. (2016) Persistence of intact retinal ganglion cell terminals after axonal transport loss in the DBA/2J mouse model of glaucoma. 524:3503
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Mellott, et al. (2014) Descending projections from auditory cortex to excitatory and inhibitory cells in the nucleus of the brachium of the inferior colliculus. Front Syst Neurosci 8:188
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Mellott, et al. (2014) Distribution of GABAergic cells in the inferior colliculus that project to the thalamus. Front Neuroanat 8:17
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Zhang, et al. (2021) Endogenous cholinergic signaling modulates sound-evoked responses of medial nucleus of trapezoid body. J Neurosci, 41:674
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Beebe, et al. (2020) Perineuronal nets and subtypes of GABAergic cells differentiate auditory and multisensory nuclei in the intercollicular area of the midbrain. J Comp Neurol, 528:2695
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Goyer, et al. (2019) A novel class of inferior colliculus principal neurons labeled in vasoactive intestinal peptide-cre mice. Elife 2019;8:e43770
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Mellott, et al. (2019) Bilateral projections to the thalamus from individual neurons in the inferior colliculus. J Comp Neurol 527:1118
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Beebe, et al. (2018) Perineuronal nets in subcortical auditory nuclei of four rodent species with differing hearing ranges. J Comp Neurol 526:972
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Beebe, et al. (2016) Extracellular molecular markers and soma size of inhibitory neurons: evidence for four subtypes of GABAergic cells in the inferior colliculus. J Neurosci 36:3988
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Nakamoto, et al. (2014) Ultrastructural characterization of GABAergic and excitatory synapses in the inferior colliculus. Front Neuroanat 8:108
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Schofield, et al. (2014) Projections from the dorsal and ventral cochlear nuclei to the medial geniculate body. Front Neuroanat 8:10