Evan Balaban

The National Science Foundation · Neural Systems

Mammalian TRPC5 channels are predominantly expressed in the brain, where they increase intracellular Ca²⁺ and induce depolarization. Because they augment presynaptic vesicle release, cause persistent neural activity, and show constitutive activity, TRPC5s could play a functional role in late developmental brain events. We used immunohistochemistry...

Neuroimaging studies show that the brain articulatory motor system is activated during speech perception. These results, however, cannot determine whether the motor system is engaged in motor simulation (e.g., lip movement) or in non-motor computations. To gauge the causal role of the articulatory motor system, here we examine the effect of mechani...

The present study was undertaken because no previous developmental studies exist on MCH neurons in any avian species. After validating a commercially-available antibody for use in chickens, immunohistochemical examinations first detected MCH neurons around embryonic day (E) 8 in the posterior hypothalamus. This population increased thereafter, reac...

The CUBIC tissue clearing protocol has been optimized to produce translucent immunostained whole chicken embryos and embryo brains. When combined with multispectral light sheet microscopy, the validated protocol presented here provides a rapid, inexpensive and reliable method for acquiring accurate histological images that preserve three-dimensiona...

Coordinated activity in different sets of widely-projecting neurochemical systems characterize waking (W) and sleep (S). How and when this coordination is achieved during development is not known. We used embryos and newborns of a precocial bird species (chickens) to assess developmental activation in different neurochemical systems using cFos expr...

Individuals with dyslexia exhibit subtle impairments in speech processing. Informed by these findings, a large literature has attributed dyslexia to a phonological deficit. Phonology, however, is only one of many systems engaged in speech processing. Accordingly, the speech perception deficit is consistent with any of multiple loci, including both...

A major unanswered question in the field of sleep studies concerns the developmental emergence of cycles of sleep and waking during embryonic life. One means for gaining valuable information about the development of this aspect of global brain function is to relate brain electrical activity recorded with electroencephalography (EEG) signals to brai...

The mechanisms underlying perceptual adaptation to severely spectrally-distorted speech were studied by training participants to comprehend spectrally-rotated speech, which is obtained by inverting the speech spectrum. Spectral-rotation produces severe distortion confined to the spectral domain while preserving temporal trajectories. During five 1-...

cFos expression (indicating a particular kind of neuronal activation) was examined in embryonic day (E) 18 chick embryos after exposure to 4 hours of either normoxia (21% O2 ), modest hypoxia (15% O2 ), or medium hypoxia (10% O2 ). Eight regions of the brainstem and hypothalamus were surveyed, including seven previously shown to respond to hypoxia...

Categorical perception occurs when a perceiver's stimulus classifications affect their ability to make fine perceptual discriminations and is the most intensively studied form of category learning. On the basis of categorical perception studies, it has been proposed that category learning proceeds by the deformation of an initially homogeneous perc...

Terrestrial vertebrate embryos face a risk of low oxygen availability (hypoxia) that is especially great during their transition to air-breathing. To better understand how fetal brains respond to hypoxia, we examined the effects of low oxygen availability on brain activity in late-stage chick embryos (day 18 out of a 21 day incubation period). Usin...

Pitch and timbre perception are both based on the frequency content of sound, but previous perceptual experiments have disagreed about whether these two dimensions are processed independently from each other. We tested the interaction of pitch and timbre variations using sequential comparisons of sound pairs. Listeners judged whether two sequential...

The developmental origins of sleep and brain activity rhythms in higher vertebrate animals (birds and mammals) are currently unknown. In order to create an experimental system in which these could be better elucidated, we designed, built and tested a system for recording EEG and EMG signals in-ovo from chicken embryos incubated for 16-21 days. This...

The human cerebral cortex appears to shrink during adolescence. To delineate the dynamic morphological changes involved in this process, 52 healthy male and female adolescents (11-17 years old) were neuroimaged twice using magnetic resonance imaging, approximately 2 years apart. Using a novel morphometric analysis procedure combining the FreeSurfer...

Dyslexia is commonly attributed to a phonological deficit, but whether it effectively compromises the phonological grammar or lower level systems is rarely explored. To address this question, we gauge the sensitivity of dyslexics to grammatical phonological restrictions on spoken onset clusters (e.g., bl in block). Across languages, certain onsets...

Dyslexia is associated with numerous deficits to speech processing. Accordingly, a large literature asserts that dyslexics manifest a phonological deficit. Few studies, however, have assessed the phonological grammar of dyslexics, and none has distinguished a phonological deficit from a phonetic impairment. Here, we show that these two sources can...

The effect of reading skill on the discrimination of words from nonwords (in Experiment 1). Note: Box plots mark one SE above and below the mean. Each whisker bar marks 2 SD. Individual data plots are indicated by triangles. (PDF)

The effect of reading skill on the discrimination of speech from nonspeech (in Experiment 2). Note: Box plots mark one SE above and below the mean. Each whisker bar marks two SD. Individual data plots are indicated by triangles. (PDF)

Response of type time to speech and nonspeech as a function of reading skill and stem type (in Experiment 2). Note: Box plots mark one SE above and below the mean. Each whisker bar marks two SD. Individual data plots are indicated by triangles. (PDF)

Identification of the ba-pa continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s response data across step (treated as a continuous variable) using logistic regression. (PDF)

Identification of the a-e continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s response data across step (treated as a continuous variable) using logistic regression. (PDF)

Discrimination in the da-ta continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s accuracy data across step using logistic regression with a natural cubic spline (df = 2). (PDF)

Discrimination in the o-u continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s accuracy data across step using logistic regression with a natural cubic spline (df = 2). (PDF)

Discrimination in the a-e continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s accuracy data across step using logistic regression with a natural cubic spline (df = 2). (PDF)

Response time and response accuracy to nonwords as a function of reading skill and stem type (in Experiment 1). Note: Box plots mark two SE above and below the mean. Each whisker bar marks two SD. (PDF)

Identification of the da-ta continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s response data across step (treated as a continuous variable) using logistic regression. (PDF)

Identification of the o-u continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s response data across step (treated as a continuous variable) using logistic regression. (PDF)

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Discrimination in the ba-pa continuum by dyslexic and skilled readers. Regression lines were fit to each individual’s accuracy data across step using logistic regression with a natural cubic spline (df = 2). (PDF)

Certain ill-formed phonological structures are systematically under-represented across languages and misidentified by human listeners. It is currently unclear whether this results from grammatical phonological knowledge that actively recodes ill-formed structures, or from difficulty with their phonetic encoding. To examine this question, we gauge t...

Like humans, birds that exhibit vocal learning have relatively delayed telencephalon maturation, resulting in a disproportionately smaller brain prenatally but enlarged telencephalon in adulthood relative to vocal non-learning birds. To determine if this size difference results from evolutionary changes in cell-autonomous or cell-interdependent dev...

A frontal view of dissected brain in chimeric head at ED16. Optic nerves of zebra finch eyes innervate the quail optic tecta in the chimeric head. (TIF)

Because acoustic landscapes are complex and rapidly changing, auditory systems have evolved mechanisms that permit rapid detection of novel sounds, sound source segregation and perceptual restoration of sounds obscured by noise. Perceptual restoration is particularly important in noisy environments, because it allows organisms to track sounds over...

The brain network underlying speech comprehension is usually described as encompassing fronto-temporal-parietal regions while neuroimaging studies of speech intelligibility have focused on a more spatially restricted network dominated by the superior temporal cortex. Here we use functional magnetic resonance imaging with a novel whole-brain multiva...

Experience-dependent plastic changes in the brain underlying complex forms of learning are generally initiated when organisms are awake, and this may limit the earliest developmental time at which learning about external events can take place. It is not known whether waking-like brain function is present prenatally in higher vertebrate (bird or mam...

Learners can segment potential lexical units from syllable streams when statistically variable transitional probabilities between adjacent syllables are the only cues to word boundaries. Here we examine the nature of the representations that result from statistical learning by assessing learners’ ability to generalize across acoustically different...

It has long been known that the identification of aural stimuli as speech is context-dependent (Remez et al., 1981). Here, we demonstrate that the discrimination of speech stimuli from their non-speech transforms is further modulated by their linguistic structure. We gauge the effect of phonological structure on discrimination across different mani...

Example of Discontinuity illusion stimulus: Continuous tone with spectrally remote noise. The tone sounds discontinuous to some listeners. (WAV)

Example of Continuity illusion stimulus (noise duration 50 ms). (WAV)

Preliminary Data Streaming and illusory discontinuity are related. (DOC)

No relationship between noise detection thresholds and continuity/discontinuity perception. (DOC)

Important sounds can be easily missed or misidentified in the presence of extraneous noise. We describe an auditory illusion in which a continuous ongoing tone becomes inaudible during a brief, non-masking noise burst more than one octave away, which is unexpected given the frequency resolution of human hearing. Participants strongly susceptible to...

The study was carried out to assess the role that five hearing history variables (chronological age, age at onset of deafness, age of first cochlear implant [CI] activation, duration of CI use, and duration of known deafness) play in the ability of CI users to identify speaker gender. Forty-one juvenile CI users participated in two voice gender ide...

Music and speech are complex sound streams with hierarchical rules of temporal organization that become elaborated over time. Here, we use functional magnetic resonance imaging to measure brain activity patterns in 20 right-handed nonmusicians as they listened to natural and temporally reordered musical and speech stimuli matched for familiarity, e...

Domain-specific systems are hypothetically specialized with respect to the outputs they compute and the inputs they allow (Fodor, 1983). Here, we examine whether these 2 conditions for specialization are dissociable. An initial experiment suggests that English speakers could extend a putatively universal phonological restriction to inputs identifie...

Comparative biology can offer important insights into the evolution of dynamic coordination in the brain. This chapter explores the neural machinery and computations shared by all nervous systems across the animal kingdom, taking into account the fact the complex relationships in coordination between nervous systems and the behavior they produce. I...

An examination of how widely distributed and specialized activities of the brain are flexibly and effectively coordinated. A fundamental shift is occurring in neuroscience and related disciplines. In the past, researchers focused on functional specialization of the brain, discovering complex processing strategies based on convergence and divergence...

A numerical thought experiment was conducted to assess whether stimulus-specific, short-term changes in auditory neural responsiveness could explain the formation of auditory objects underlying the auditory continuity illusion. A tonotopic, two-layer feedforward network model with one time constant for synaptic weight augmentation based on firing r...

Reading impairments have previously been associated with auditory processing differences. We examined auditory stream biasing, a global aspect of auditory temporal processing. Children with reading impairments, control children and adults heard a 10 s long stream-bias-inducing sound sequence (a repeating 1000 Hz tone) and a test sequence (eight rep...

Gender identification of human voices was studied in a juvenile population of cochlear implant (CI) users exposed to naturalistic speech stimuli from 20 male and 20 female speakers using two different voice gender perception tasks. Stimulus output patterns were recorded from each individual CI for each stimulus, and features related to voice fundam...

Psychosis is associated with volumetric decreases of cortical structures. Whether these volumetric decreases imply abnormalities in cortical thickness, surface, or cortical folding is not clear. Due to differences in cytoarchitecture, cortical gyri and sulci might be differentially affected by psychosis. Therefore, we examined differences in gyral...

It is known that unpleasant emotions can modulate the speed of involuntary movements, yet the effects of aversive stimulation on voluntary motor acts have not been systematically investigated. The effects of aversive stimulation on subsequent movement-related cortical activity were examined using a task invalving compatible and incompatible movemen...

Relative percentage of rotated consonants with the indicated feature (rows) identified as a vowel (columns), and the relative mutual information between rotated consonant features and vowel identification (rMI). (0.04 MB DOC)

Percentages of identification of rotated consonants (rows) as unrotated vowels (columns) different from the following vowel in the syllable (0.04 MB DOC)

Percentages of identification of PLACE feature properties after rotation. (0.03 MB DOC)

Percentages of identification of VOICING feature properties after rotation. (0.03 MB DOC)

Percentages of identification of MANNER feature properties after rotation. (0.03 MB DOC)

Comparison of consonant identification performance with the Blesser study [s1]. (0.03 MB DOC)

Normal Syllables. The syllabic sequence \ba\-\be\-\bo\-\bu\ pronounced by a male speaker. (0.22 MB WAV)

Spectrally-rotated Syllables. The spectrally-rotated syllabic sequence \ba\-\be\-bo\-\bu\ pronounced by a male speaker (rotation frequency = 1.5 kHz). (0.22 MB WAV)

Sex Differences in the Brain . From Genes to Behavior. Jill B. Becker, Karen J. Berkley, Nori Geary, Elizabeth Hampson, James P. Herman, and Elizabeth A. Young, Eds. . Oxford University Press, New York, 2008. 504 pp. $98.50, £54. ISBN 9780195311587. Offering a comprehensive introduction for researchers, students, and clinicians, the contributors c...

Neuroimaging studies of speech processing increasingly rely on artificial speech-like sounds whose perceptual status as speech or non-speech is assigned by simple subjective judgments; brain activation patterns are interpreted according to these status assignments. The naïve perceptual status of one such stimulus, spectrally-rotated speech (not con...

During neural development, neurons from downstream, presynaptic regions of the nervous system (such as the retina) send spatially patterned axonal projections to upstream, target regions (the tectum or superior colliculus). A servomechanism model has been proposed to explain the pattern and time-course of axonal growth between these two regions [Ho...

The effects of acoustic features and native phonology on the naïve perception of unfamiliar spectrally‐rotated speech syllables were evaluated. Spectrally‐rotated syllables were created by rotating the spectrum of naturally produced Italian CV syllables around 1500 Hz resulting in changes in the energy distribution of harmonics while preserving the...

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Mean evoked potential responses to stimuli. (a) The five different tone complexes used are shown color-coded above their constituent components on an amplitude - frequency plot (bottom), where the components of each tone are overlaid on a common set of axes. (b,c) Mean relative amplitude spectra (22 participants) for the 0° (in b) and 180° (in c) r...

Current theories of auditory pitch perception propose that cochlear place (spectral) and activity timing pattern (temporal) information are somehow combined within the brain to produce holistic pitch percepts, yet the neural mechanisms for integrating these two kinds of information remain obscure. To examine this process in more detail, stimuli mad...

Biological contributions to cognitive development continue to be conceived predominantly along deterministic lines, with proponents of different positions arguing about the preponderance of gene-based versus experience-based influences that organize brain circuits irreversibly during prenatal or early postnatal life, and evolutionary influences act...

Working together at Nogent, Marie-Aimee Teillet, Nicole Le Douarin and the author successfully developed an extension of the quail-chick transplant technique for relating species brain cell differences to behavioral differences. This article reviews the application of the technique to species differences in motor behavior (crowing) and auditory per...

Innervation of the visual midbrain by axons from the retina can be described as a stochastic mapping process that maintains topography and polarity between the two regions. Previous work has identified a number of mechanisms that insure proper guidance of the axons. In the current report, we combine three of these mechanisms, servomechanical guidan...

The tendency for animals to form social bonds after sexual activity varies greatly from species to species. Work with voles illuminates a molecular pathway in the brain that influences such differences.

During space flight astronauts show vestibular-related changes in balance, eye movements, and spontaneous and reflex control of cardiovascular, respiratory and gastrointestinal function, sometimes associated with space motion sickness. These symptoms undergo compensation over time. Here we used changes in the expression of two immediate-early gene...

We recorded human auditory cortical activity during the perception of long, changing acoustic signals and analyzed information provided by dynamic neural population measures over a large range of time intervals (approximately 24 ms-5 s). Participants listened to musical scales that were amplitude modulated at a rate of 41.5 Hz, generating an ongoin...

Changes in gene expression were examined in precerebellar structures during and after space flight. These structures included the inferior olive (IO), the source of climbing fibers, and the lateral reticular nucleus (LRt) and basilar pontine nuclei (PN), sources of mossy fibers. We examined two immediate early gene products with two different time...

The nucleus paragigantocellularis lateralis (LPGi) exerts a prominent excitatory influence over locus coeruleus (LC) neurons, which respond to gravity signals. We investigated whether adult albino rats exposed to different gravitational fields during the NASA Neurolab Mission (STS-90) showed changes in Fos and Fos-related antigen (FRA) protein expr...

Auditory-induced expression of the immediate-early gene ZENK was examined in 18 brain areas of domestic chicken and Japanese quail subjects with no previous exposure to parental vocalizations. After one 30-min exposure (approximately 120 calls) within 24 h of hatching to either the chicken or quail maternal call, paired subjects from each species s...

Changes in immediate-early gene (IEG) expression during and after space flight were studied in the rat locus coeruleus (LC) during the NASA Neurolab mission. The LC sends widespread projections throughout the brain and releases the neuromodulator norepinephrine. LC neurons respond to natural vestibular stimulation; their firing rate also increases...

Space flight produces profound changes of neuronal activity in the mammalian vestibular and reticular systems, affecting postural and motor functions. These changes are compensated over time by plastic alterations in the brain. Immediate early genes (IEGs) are useful indicators of both activity changes and neuronal plasticity. We studied the expres...

To assess effects of tonic gravity changes on efferent vestibular neurons, immediate early gene (IEG) protein expression was compared in the efferent vestibular nucleus (EVe) of adult rats exposed to microgravity during the NASA Neurolab Mission (STS-90), and matching control animals on the ground. Rats sacrificed 1 day and 12 days after launch (ex...

'Pitch' refers to a sound's subjective highness or lowness, as distinct from 'frequency,' which refers to a sound's physical structure. In speech, music and other natural contexts, complex tones are often perceived with a single pitch. Using whole-head magnetoencephalography (MEG) and stimuli that dissociate pitch from frequency, we studied cortica...

Inborn species' perceptual preferences are thought to serve as important guides for neonatal learning in most species of higher vertebrates. Although much work has been carried out on experiential contributions to the expression of such preferences, their neural and developmental correlates remain largely unexplored. Here we use embryonic neural tr...

Memory is one of the most fundamental mental processes. Neuroscientists study this process by using extremely diverse strategies. Two different approaches aimed at understanding learning and memory were introduced in this symposium. The first focuses on the roles played by synaptic plasticity, especially in long-term depression in the cerebellum in...