Our Research

You can check out a complete list of Schofield lab publications here. You can also read about our techniques, our funding, and our collaborators.

Current Projects

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.

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.

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?

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.

Beebe, Silveira, et al. (2022) Neurotransmitter phenotype and axonal projection patterns of VIP-expressing neurons in the inferior colliculus. J Chem Neuroanat, 126:102189
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
Beebe, Zhang, et al. (2021) Multiple sources of cholinergic input to the superior olivary complex. Front Neural Circuits 15:715369
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
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
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
Crish and Schofield (2018) Anterograde tract tracing for assaying axonopathy and transport deficits in glaucoma. Methods Mol Biol 1695:171
Mellott, et al. (2014) Excitatory and inhibitory projections in parallel from the inferior colliculus to the auditory thalamus. Front Neuroanat 8:124
Schofield, et al. (2014) Subcollicular projections to the auditory thalamus and collateral projections to the inferior colliculus. Front Neuroanat 8:70
Foster, et al. (2014) Perineuronal nets and GABAergic cells in the inferior colliculus of guinea pigs. Front Neuroanat 7:53
Anair, et al. (2022) Inhibitory NPY neurons provide a large and heterotopic commissural projection in the inferior colliculus. Front Neural Circuits 16: 8711924
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
Silveira, et al. (2020) Neuropeptide Y expression defines a novel class of GABAergic projection neuron in the inferior colliculus. J Neurosci 40:4685
Schofield and Beebe (2019) Subtypes of GABAergic cells in the inferior colliculus. Hear Res 376:1
Mellott, et al. (2018) GABAergic and nonGABAergic projections to the superior colliculus from the auditory brainstem. Brain Struct Funct 223:1923
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
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
Mellott, et al. (2014) Distribution of GABAergic cells in the inferior colliculus that project to the thalamus. Front Neuroanat 8:17
Beebe, et al. (2023) Characterization of three cholinergic inputs to the cochlear nucleus. J Chem Neuroanat, 131: 102284
Noftz, et al. (2021) Dense cholinergic projections to auditory and multisensory nuclei of the intercollicular midbrain. Hear Res, 411: 108352
Zhang, et al. (2021) Endogenous cholinergic signaling modulates sound-evoked responses of medial nucleus of trapezoid body. J Neurosci, 41:674
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
Goyer, et al. (2019) A novel class of inferior colliculus principal neurons labeled in vasoactive intestinal peptide-cre mice. Elife 2019;8:e43770
Mellott, et al. (2019) Bilateral projections to the thalamus from individual neurons in the inferior colliculus. J Comp Neurol 527:1118
Beebe, et al. (2018) Perineuronal nets in subcortical auditory nuclei of four rodent species with differing hearing ranges. J Comp Neurol 526:972
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
Nakamoto, et al. (2014) Ultrastructural characterization of GABAergic and excitatory synapses in the inferior colliculus. Front Neuroanat 8:108
Schofield, et al. (2014) Projections from the dorsal and ventral cochlear nuclei to the medial geniculate body. Front Neuroanat 8:10

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