Neurobiology of Speech and Language :: Syllabus

Term: Fall 2016
Instructor: Daniel Meliza (cdm8j)
Class times: T/Th 11:00A-12:15P, Gilmer 240
Office Hours: Gilmer 183, T 1-2P, Th 3-4, or by appointment
Collab site:
Last revised: 8/23/2016

What is this course about?

Spoken language is a complicated, rich, and beautiful behavior. When you speak or listen, billions of neurons in your brain are engaged in translating between tiny variations in air pressure and the concepts, images, and memories they carry.

How does this all come about? How do the circuits involved in speech and language work? How do neurons wire themselves together into circuits and learn how to produce and perceive speech? Did language suddenly happen to our species, or did it evolve out of simpler components that we can study in other animals?

In this course, we will learn about mechanisms of speech and language by discussing a series of experimental animal models that exhibit constituents of language or speech, building towards behaviors that integrate multiple processes and culminating with a look at some of the data from human imaging studies.

Course objectives

Speech and language make us who we are as a species, and understanding them better can be both intrinsically fascinating and deeply practical. By the end of the course, you will be able to answer the following questions:

You will also gain experience reading journal articles, analyzing scientific arguments, and researching topics, enabling you to answer these questions as well:

Meeting the objectives

Reading responses and forum discussion (25%)

To prepare ourselves for discussions that go beyond the basic content of the readings, by Monday morning of each week, we will each post a blog entry to the Collab site that describes an aspect of the readings to discuss further in class. Entries will be between 200-300 words and should:

You’ll each be assigned to small groups who will read each other’s responses before coming to class, so that we can respond to each other’s ideas and reach a better understanding together.

In-class discussion (25%)

Active participation in class discussions is essential to your learning experience in this course. In order to participate, you must

Of course, none of this can happen unless you are present. If unforeseen events illness, religious holidays, or academic/athletic field trips prevent you from attending, I expect you to notify me ahead of time so we can discuss make-up work. Only major, documented emergencies are acceptable excuses after the fact.

Discussion facilitation (30%)

You will have two opportunities to take a more active role in guiding the class meeting, using papers I’ve selected. Facilitation involves:

I will act as a co-facilitator on these exercises, but it will be your responsibility to lead. Your grade will be based on a rubric (that I will share with you) that assesses your understanding of the concepts and background, your clear and accurate presentation of the data, and your ability to engage the class in a good discussion.

Topic selection and discussion (20%)

In the last weeks of class, each of you will have the opportunity to facilitate a discussion on a topic of your choice. You may choose something new to address or revisit an earlier subject, and you’re encouraged to draw on your previous coursework and interests. In addition to the tasks described in the previous section, you will need to select one or more papers as readings and post the references to Collab by Monday of the week before your assigned date. We’ll make initial topic selections early in the semester, but you may change your topic (subject to my approval) at any point up until the readings are posted.

What materials will you need?

All of our assigned readings will be primary research papers and reviews that you can find using PubMed, Google Scholar, or VIRGO. Some papers may only be accessible from on grounds or by using the VPN or web proxy as described here. In some rare cases the readings may not be available online, in which case I will scan them and make them available through the class Collab page.

You will probably need to consult additional resources to help you understand some of the concepts we will discuss. Like working research scientists, you can use handbooks, textbooks, online resources, peer-reviewed articles, and personal communications to learn what you need to know to complete the full story surrounding the questions we’ll be addressing. If you need additional help finding sources, please contact me, your reference librarian, or the Source Dorks.

What is the class schedule?

This schedule will be updated with readings and presentations as the semester progresses. Resources in italics are optional. Check back frequently.

Week Topic/Readings Assignments
8/23 What is language and where did it come from?
Feher O et al (2009) De novo establishment of wild-type song culture in the zebra finch. Nature, doi:10.1038/nature07994
Hauser MD et al (2002) The Faculty of Language: What Is It, Who Has It, and How Did It Evolve? Science, doi:10.1126/science.298.5598.1569
Deacon TW (2010) A role for relaxed selection in the evolution of the language capacity. PNAS, doi:10.1073/pnas.0914624107
Optional: Pinker S and Jackendoff R (2005). The faculty of language: what’s special about it? Cognition, doi:10.1016/j.cognition.2004.08.004
8/30 Communication, information, and signal detection
Maynard Smith J and Harper DGC (1995) Animal Signals: Models and Terminology. J Theor Biol, doi:10.1006/jtbi.1995.0248
Bee MA and Gerhardt HC (2002) Individual voice recognition in a territorial frog (Rana catesbeiana). Proc R Soc Lond B, doi:10.1098/rspb.2002.2041
Wiley RH (2006) Signal Detection and Animal Communication, in Advances in the Study of Behavior, Vol. 36
Iverson P and Kuhl PK (1995) Mapping the perceptual magnet effect for speech using signal detection theory and multidimensional scaling. J Acoust Soc Amer, 97(1):553-62
9/6 Bioacoustics & biophysics of sound generation Choose dates for facilitation I
Johnson, KS et al (2011) Acoustic and Auditory Phonetics. Hoboken: Wiley-Blackwell. Chs 1-2 ONLY. 9/7: last day to drop
Riede T and Goller F (2010) Peripheral mechanisms for vocal production in birds – differences and similarities to human speech and singing. Brain Lang, doi:10.1016/j.bandl.2009.11.003
Fitch WT et al (2002) Calls out of chaos: the adaptive significance of nonlinear phenomena in mammalian vocal production. Animal Behaviour, doi:10.1006/anbe.2001.1912
9/13 The auditory system and vocal perception
Sharpee TO et al (2011) Hierarchical representations in the auditory cortex. Curr Opin Neurobiol, doi:10.1016/j.conb.2011.05.027
de Charms RC et al (1998) Optimizing sound features for cortical neurons. Science, doi:10.1126/science.280.5368.1439
Prather JF (2009) Neural correlates of categorical perception in learned vocal communication. Nat Neurosci, doi:10.1038/nn.2246
Romanski LM et al (2004) Neural representation of vocalizations in the primate ventrolateral prefrontal cortex. J Neurophys, doi:10.1152/jn.00675.2004
9/20 Bird calls, songs & the evolution of learned and unlearned signals
Pepperberg IM (2013) Evolution of vocal communication: an avian model. Ch 26 in Birdsong, speech, and language : exploring the evolution of mind and brain, eds JJ Bolhuis and M Everaert. Cambridge, Mass: MIT Press.
Marler P (2004) Bird calls: their potential for behavioral neurobiology. Ann N Y Acad Sci, doi:10.1196/annals.1298.034
Marler P and Tamura M (1964) Culturally transmitted patterns of vocal behavior in sparrows. Science 146:1483–86
Kroodsma DE and Konishi M (1991) A suboscine bird (eastern phoebe, Sayornis phoebe) develops normal song without auditory feedback. Anim Behav 42:477-487
9/27 Sensorimotor learning in the song system Student facilitation
Kelsey: Lalazar H, Vaadia E (2008). Neural basis of sensorimotor learning: modifying internal models. Curr Opin Neurobiol, doi:10.1016/j.conb.2008.11.003 Choose dates for facilitation II
Anthony: Brainard MS and Doupe AJ (2000) Interruption of a basal ganglia–forebrain circuit prevents plasticity of learned vocalizations. Nature 404:762-766
Owen: Olveczky BP et al (2005) Vocal Experimentation in the Juvenile Songbird Requires a Basal Ganglia Circuit. Plos Biol doi:10.1371/journal.pbio.0030153
Grace: Hadipour-Niktarash A et al (2007) Impairment of retention but not acquisition of a visuomotor skill through time-dependent disruption of primary motor cortex. J Neurosci 27:13413-13419.
Optional: Tin C and Poon C-S (2005) Internal Models in Sensorimotor Integration: Perspectives from Adaptive Control Theory. J Neur Eng doi:10.1088/1741-2560/2/3/S01
10/4 Interactions between production and perception No class Tuesday
Michael: Liberman and Mattingly (1985) The motor theory of speech perception revised. Cognition 21, 1-36
Rachel: Skipper JI et al (2007) Hearing Lips and Seeing Voices: How Cortical Areas Supporting Speech Production Mediate Audiovisual Speech Perception Cereb. Cortex doi:10.1093/cercor/bhl147
10/11 Phonetics and perception
Johnson, KS et al (2011) Acoustic and Auditory Phonetics. Hoboken: Wiley-Blackwell. Chs 6-7 ONLY.
Abby: Bouchard, KE et al (2013). Functional organization of human sensorimotor cortex for speech articulation. Nature, doi:0.1038/nature11911
Sravya: Maye and Werker (2008). Infant sensitivity to distributional information can affect phonetic discrimination. Cognition, doi:10.1016/S0010-0277(01)00157-3
10/18 Development of human speech perception
Kuhl PK (2000) A new view of language acquisition. PNAS doi:10.1073/pnas.97.22.11850 10/18 last day to withdraw
Sravya: Dehaene-Lambertz G et al (2002) Functional Neuroimaging of Speech Perception in Infants. Science doi:10.1126/science.1077066
Rachel: Bortfield H et al (2009) Identifying cortical lateralization of speech processing in infants using near-infrared spectroscopy. Dev Neuropsyc 34(1) 52-65
Owen: Bruderer AG et al (2015) Sensorimotor influences on speech perception in infancy. Proc Natl Acad Sci doi:10.1073/pnas.1508631112
10/25 Development of human speech production
Michael: Perani D et al (2011) Neural language networks at birth. Proc Natl Acad Sci doi:10.1073/pnas.1102991108
Grace: Golstein MH et al (2003) Social interaction shapes babbling: Testing parallels between birdsong and speech. Proc Natl Acad Sci doi:10.1073/pnas.1332441100
Abby: Nip ISB et al (2009) Early speech motor development: Cognitive and linguistic considerations. J Comm Disord doi:10.1016/j.jcomdis.2009.03.008
Kelsey: Vallentin D et al (2016) Inhibition protects acquired song segments during vocal learning in zebra finches. Science doi:10.1126/science.aad3023
11/1 Grammar and temporal patterns
Anthony:Steinhauer K et al (1999) Brain potentials indicate immediate use of prosodic cues in natural speech processing. Nature Neurosci doi:10.1038/5757
Saffran J et al (2008) Grammatical Pattern Learning by Human Infants and Cotton-Top Tamarin Monkeys. Cognition doi:10.1016/j.cognition.2007.10.010
11/3 Student topic: Owen - Handedness and Lateralization
11/8 Student topic: Rachel - Autism, Lateralization, and Speech
11/10 Student topic: Grace - Bilingualism
11/17 Student topic: Sravya - Aphasia No class Tuesday
11/22 Student topic: Abby - Speech Pathologies No class Thursday
11/29 Student topic: Michael - Aphasia in Bilinguals
12/1 Student topic: Kelsey - Language without Speech
12/6 Student topic: Anthony - Music and Language

How will you succeed in this course?

Participate. You are expected to participate actively in the course based on your own learning goals. You all come from different backgrounds and science experiences, and your peers are valuable resources for learning. Don’t shortchange them and yourself by coming to class without preparing or by sitting quietly during class discussion.

Indulge your curiosity. Unlike some of the other topics you may have studied, the neurobiology of speech and language is in many ways still in its infancy. We will cover some foundational knowledge, but our ultimate goals are to identify open problems in the field and draw connections between different systems and species. You will likely come away with more questions than when you started, but with a greater ability to understand the questions and where they might lead. To help you think of new questions and ways to research them, I’ll share some of the ways I approach the literature and use it to stimulate further reading, research, and experiment.

Write frequently. One of the interesting features of language is that putting your ideas into words is often the best way of clarifying them. Striving for clarity and conciseness will magnify this effect. In this course, we will write weekly responses to the readings and submit them to a small group of peers for comment and clarification. Although these exercises will be short, they will help you the most if you craft them carefully.

Professional and academic integrity

As practicing professionals, scientists trust each other to maintain the highest standards of ethics, integrity, and personal responsibility. Since you have joined this community of trust to prepare for your future career, I expect you to fully comply with all of the provisions of the UVa Honor System. In addition to pledging that you have neither received nor given aid on an assignment, your signature also affirms that you have not knowingly represented as your own any opinions or ideas that are attributable to another author in published or unpublished notes, study outlines, abstracts, articles, textbooks, or web pages. In other words, I expect that all assignments and reports are your original work and that references are cited appropriately. Breaking this trust agreement not only will result in zero credit for the assignment in question and referral to the Honor Committee but also will jeopardize your future as a professional scientist or in any field. Don’t let yourself down.

What accomodations are available for students with disabilities?

Students with disabilities are entitled to reasonable accommodations. The Student Disability Access Center (434-243-5180), located in the Elson Student Health Center, can arrange diagnostic testing and make recommendations for specific accommodations. Your Association Dean can also respond to requests for information and assistance.