Neurobiology of Speech and Language :: Syllabus

Term: Fall 2018
Instructor: Daniel Meliza (cdm8j)
Class times: T/Th 11:00A-12:15P, Shannon House 111
Course site:
Collab site:
Piazza site:
Office Hours: Gilmer 481, Tu 1-3 or by appointment
Last revised: 8/31/2018

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 the biology of language-learning disabilities.

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
8/28 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
9/4 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
9/11 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.
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
9/18 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
9/25 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
10/2 Sensorimotor learning Student facilitation begins
Background: 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
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
Tyler: Brainard MS and Doupe AJ (2000) Interruption of a basal ganglia–forebrain circuit prevents plasticity of learned vocalizations. Nature 404:762-766
Madison: Olveczky BP et al (2005) Vocal Experimentation in the Juvenile Songbird Requires a Basal Ganglia Circuit. Plos Biol doi:10.1371/journal.pbio.0030153
Andrew: Valletin D et al (2016) Inhibition protects acquired song segments during vocal learning in zebra finches. Science doi:10.1126/science.aad3023
Matt: Giret N et al (2014) Evidence for a causal inverse model in an avian cortico-basal ganglia circuit. PNAS doi:10.1073/pnas.1317087111
10/16 Phonetics, production and perception
Background: Johnson, KS et al (2011) Acoustic and Auditory Phonetics. Hoboken: Wiley-Blackwell. Chs 6-7 ONLY.
Dan: Liberman and Mattingly (1985) The motor theory of speech perception revised. Cognition 21, 1-36
Wangqi: Maye and Werker (2008). Infant sensitivity to distributional information can affect phonetic discrimination. Cognition, doi:10.1016/S0010-0277(01)00157-3
Cat: Bouchard, KE et al (2013). Functional organization of human sensorimotor cortex for speech articulation. Nature, doi:0.1038/nature11911
Sophia: 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
10/23 Development of human speech perception
Kenna: Bosseler AN et al (2013) Theta brain rhythms index perceptual narrowing in infant speech perception. Front Psychol doi:10.3389/fpsyg.2013.00690
Yafi: Partanen E et al (2015) Learning-induced neural plasticity of speech processing before birth. PNAS doi:10.1073/pnas.1302159110
Flo: Bruderer AG et al (2015) Sensorimotor influences on speech perception in infancy. Proc Natl Acad Sci doi:10.1073/pnas.1508631112
Adina: Aslin et al (1998) Computation of Conditional Probability Statistics by 8-Month-Old Infants. Psychol Sci doi:10.1111/1467-9280.00063
10/30 Developmental language disorders
Background: Pennington and Bishop (2009) Relations among speech, language, and reading disorders. Annu Rev Psychol doi:10.1146/annurev.psych.60.110707.163548
Julia: van Zuijen et al (2013) Infant ERPs separate children at risk of dyslexia who become good readers from those who become poor readers. doi:10.1111/desc.12049
Peri: Halliday et al (2017) Auditory processing deficits are sometimes necessary and sometimes sufficient for language difficulties in children: Evidence from mild to moderate sensorineural hearing loss. Cognition doi:10.1016/j.cognition.2017.04.014
Miriam: Pinel et al (2012) Genetic variants of FOXP2 and KIAA0319/TTRAP/THEM2 locus are associated with altered brain activation in distinct language-related regions. J Neurosci doi:10.1523/JNEUROSCI.5996-10.2012
Helina: Centanni et al (2014) Knockdown of the dyslexia-associated gene Kiaa0319 impairs temporal responses to speech stimuli in rat primary auditory cortex. Cereb Cortex doi:10.1093/cercor/bht028
11/6 Grammar and temporal patterns
Xiaomu: Steinhauer K et al (1999) Brain potentials indicate immediate use of prosodic cues in natural speech processing. Nature Neurosci doi:10.1038/5757
Dan: 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
11/8 Andrew: Seery AM et al (2013) Atypical lateralization of ERP response to native and non-native speech in infants at risk for autism spectrum disorder. Dev Cog Neurosci doi:10.1016/j.dcn.2012.11.007
Madison: Young BK et al (2017) Adult zebra finches rehearse highly variable song patterns during sleep. PeerJ doi:10.7717/peerj.4052
11/13 Helina: TBD
Julia: Mock JR et al (2015) Speech preparation in adults with persistent developmental stuttering. Brain & Lang doi:10.1016/j.bandl.2015.05.009
11/15 Cat: Haebig E et al (2017) Statistical word learning in children with autism spectrum disorder and specific language impairment. J Child Psychol Psych doi:10.1111/jcpp.12734
Peri: Berko J (1958) The Child’s Learning of English Morphology. Word doi:10.1080/00437956.1958.11659661
11/20 (class canceled by instructor)
11/22 No class (Thanksgiving)
11/27 Tyler: Doelling KB et al (2014) Acoustic landmarks drive delta-theta oscillations to enable speech comprehension by facilitating perceptual parsing. Neuroimage doi:10.1016/j.neuroimage.2013.06.035
Miriam: Brookshire G et al (2017) Visual cortex entrains to sign language. PNAS doi:10.1073/pnas.1620350114
11/29 Kenna: Byers-Heinlein K et al (2017) Bilingual infants control their languages as they listen. PNAS doi:10.1073/pnas.1703220114
Yafi: Lytle SR et al (2017) Two are better than one: Infant language learning from video improves in the presence of peers. PNAS. doi:10.1073/pnas.1611621115
12/4 Sophia: Moeller MP (2000) Early Intervention and Language Development in Children Who Are Deaf and Hard of Hearing. Pediatrics 106(3):e43
Matt: Dionne G et al (2011) Associations Between Sleep-Wake Consolidation and Language Development in Early Childhood: A Longitudinal Twin Study. Sleep doi:10.5665/SLEEP.1148
Flo: Boutonnet B et al (2013) Seeing Objects through the Language Glass. J Cog Neurosci doi:10.1162/jocn_a_00415
12/6 Adina: Newman AJ et al (2002) A critical period for right hemisphere recruitment in American Sign Language processing. Nature Neuroscience doi:10.1038/nn775 Last Day of Class
Wanqi: Zinzser BD et al (2015) Second language experience modulates neural specialization for first language lexical tones. J Neurolinguistics doi:10.1016/j.jneuroling.2014.09.005
Xiaomu: Grey S and Van Hell JG (2017) Foreign-accented Speaker Identity Affects Neural Correlates of Language Comprehension. J Neuroling doi:10.1016/j.jneuroling.2016.12.001

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.