I-Hui Hsieh

Hierarchical Organization of Human Auditory Cortex: Evidence from Acoustic Invariance in the Response to Intelligible Speech

Hierarchical organization of human auditory cortex has been inferred from functional imaging observations that core regions respond to simple stimuli (tones) whereas downstream regions are selectively responsive to more complex stimuli (band-pass noise, speech). It is assumed that core regions code low-level features, which are combined at higher levels in the auditory system to yield more abstract neural codes. However, this hypothesis has not been critically evaluated in the auditory domain. We assessed sensitivity to acoustic variation within intelligible versus unintelligible speech using functional magnetic resonance imaging and a multivariate pattern analysis. Core auditory regions on the dorsal plane of the superior temporal gyrus exhibited high levels of sensitivity to acoustic features, whereas downstream auditory regions in both anterior superior temporal sulcus and posterior superior temporal sulcus (pSTS) bilaterally showed greater sensitivity to whether speech was intelligible or not and less sensitivity to acoustic variation (acoustic invariance). Acoustic invariance was most pronounced in more pSTS regions of both hemispheres, which we argue support phonological level representations. This finding provides direct evidence for a hierarchical organization of human auditory cortex and clarifies the cortical pathways supporting the processing of intelligible speech.

Fm-selective networks in human auditory cortex revealed using fmri and multivariate pattern classification

Frequency modulation (FM) is an acoustic feature of nearly all complex sounds. Directional FM sweeps are especially pervasive in speech, music, animal vocalizations, and other natural sounds. Although the existence of FM-selective cells in the auditory cortex of animals has been documented, evidence in humans remains equivocal. Here we used multivariate pattern analysis to identify cortical selectivity for direction of a multitone FM sweep. This method distinguishes one pattern of neural activity from another within the same ROI, even when overall level of activity is similar, allowing for direct identification of FM-specialized networks. Standard contrast analysis showed that despite robust activity in auditory cortex, no clusters of activity were associated with up versus down sweeps. Multivariate pattern analysis classification, however, identified two brain regions as selective for FM direction, the right primary auditory cortex on the supratemporal plane and the left anterior region of the superior temporal gyrus. These findings are the first to directly demonstrate existence of FM direction selectivity in the human auditory cortex.

Detection of sinusoidal amplitude modulation in logarithmic frequency sweeps across wide regions of the spectrum.

Many natural sounds such as speech contain concurrent amplitude and frequency modulation (AM and FM), with the FM components often in the form of directional frequency sweeps or glides. Most studies of modulation coding, however, have employed one modulation type in stationary carriers, and in cases where mixed-modulation sounds have been used, the FM component has typically been confined to an extremely narrow range within a critical band. The current study examined the ability to detect AM signals carried by broad logarithmic frequency sweeps using a 2-alternative forced-choice adaptive psychophysical design. AM-detection thresholds were measured as a function of signal modulation rate and carrier sweep frequency region. Thresholds for detection of AM in a sweep carrier ranged from -8 dB for an AM rate of 8 Hz to -30 dB at 128 Hz. Compared to thresholds obtained for stationary carriers (pure tones and filtered Gaussian noise), detection of AM carried by frequency sweeps substantially declined at low (12 dB at 8 Hz) but not high modulation rates. Several trends in the data, including sweep- versus stationary-carrier threshold patterns and effects of frequency region were predicted from a modulation filterbank model with an envelope-correlation decision statistic.

Detection of spatial cues in linear and logarithmic frequency-modulated sweeps

Frequency- and amplitude-modulated (FM and AM) sounds are the building blocks of complex sounds. In the present study, we investigated the ability of human observers to process spatial information in an important class of FM sounds: broadband directional sweeps common in natural communication signals such as speech. The stimuli consisted of linear or logarithmic unidirectional FM pulses that swept either up or down in frequency at various rates. Spatial localization thresholds monotonically improved as sweep duration decreased and as sweep rate increased, but no difference in performance was observed between logarithmic and linear or between upand down-frequency sweeps. Counterintuitive reversals in localization were observed which suggested that the localization of high-frequency sweeps may be strongly dominated by amplitude information even in situations in which one might consider timing cues to be critical. Implications of these findings for the localization of complex sounds are discussed.

Enhanced Optomotor Efficiency by Expression of the Human Gene Superoxide Dismutase Primarily in Drosophila Motorneurons

Abstract Mutation of the human gene superoxide dismutase (hSOD1) triggers the fatal neurodegenerative motorneuron disorder, familial amyotrophic lateral sclerosis (ALS or Lou Gehrigs disease). Broad expression of this gene in Drosophila has no effect on longevity or functional senescence. We show here that restricting expression of human SOD1 primarily to motorneurons of Drosophila has significant effects on optomotor efficiency during in-flight tracking of rapidly moving visual targets. Under high-stress workloads with a recursive visual-motion stimulus cycle, young isogenic controls failed to track rapidly changing visual cues, whereas their same-aged hSOD1-activated progeny maintained coordinated in-flight tracking of the target by phase locking to the dynamic visual movement patterns. Several explanations are considered for the observed effects, including antioxidant intervention in motorneurons, changes in signal transduction pathways that regulate patterns of gene expression in other cell types, and expression of hSOD1 in a small set of neurons in the central brain. That hSOD1 overexpression improves sensorimotor coordination in young organisms may suggest possible therapeutic strategies for early-onset ALS in humans.

Imperfect pitch: Gabor’s uncertainty principle and the pitch of extremely brief sounds

How brief must a sound be before its pitch is no longer perceived? The uncertainty tradeoff between temporal and spectral resolution (Gabor’s principle) limits the minimum duration required for accurate pitch identification or discrimination. Prior studies have reported that pitch can be extracted from sinusoidal pulses as brief as half a cycle. This finding has been used in a number of classic papers to develop models of pitch encoding. We have found that phase randomization, which eliminates timbre confounds, degrades this ability to chance, raising serious concerns over the foundation on which classic pitch models have been built. The current study investigated whether subthreshold pitch cues may still exist in partial-cycle pulses revealed through statistical integration in a time series containing multiple pulses. To this end, we measured frequency-discrimination thresholds in a two-interval forced-choice task for trains of partial-cycle random-phase tone pulses. We found that residual pitch cues exist in these pulses but discriminating them requires an order of magnitude (ten times) larger frequency difference than that reported previously, necessitating a re-evaluation of pitch models built on earlier findings. We also found that as pulse duration is decreased to less than two cycles its pitch becomes biased toward higher frequencies, consistent with predictions of an auto-correlation model of pitch extraction.

Velocity Selective Networks in Human Cortex Reveal Two Functionally Distinct Auditory Motion Systems

The auditory system encounters motion cues through an acoustic object’s movement or rotation of the listener’s head in a stationary sound field, generating a wide range of naturally occurring velocities from a few to several hundred degrees per second. The angular velocity of moving acoustic objects relative to a listener is typically slow and does not exceed tens of degrees per second, whereas head rotations in a stationary acoustic field may generate fast-changing spatial cues in the order of several hundred degrees per second. We hypothesized that these two types of systems (i.e., encoding slow movements of an object or fast head rotations) may engage functionally distinct substrates in processing spatially dynamic auditory cues, with the latter potentially involved in maintaining perceptual constancy in a stationary field during head rotations and therefore possibly involving corollary-discharge mechanisms in premotor cortex. Using fMRI, we examined cortical response patterns to sound sources moving at a wide range of velocities in 3D virtual auditory space. We found a significant categorical difference between fast and slow moving sounds, with stronger activations in response to higher velocities in the posterior superior temporal regions, the planum temporale, and notably the premotor ventral-rostral (PMVr) area implicated in planning neck and head motor functions.

A novel signal processing approach to auditory phantom perception

The ear and brain interact in an orchestrated manner to create sensations of phantom tones that are audible to listeners despite lacking physical presence in original sounds. The relative contribution of peripheral sensory cell activity and cortical mechanisms to phantom hearing remains elusive. The current study addressed the question of whether non-linear components of a complex signal exist that are not captured by the linear combination of cosines in a series. To this end, we investigated the source and spectro-temporal dynamics of non-linear components within two-tone complexes related to phantom acoustic perception. The empirical mode decomposition, a method for non-linear and non-stationary processes, was applied to extract the extra-aural existence of an oscillatory component within the original signal associated with the phantom sound. This travelling wave (phantom) has never before been observed in the sound’s linear spectrum. We showed that the wave travels at a velocity that accurately maps onto the perceived phantom tone frequency. Phase coherence of oscillatory mode dynamics predicted discrimination sensitivity to phantom sounds by listeners. Perceived incidences of phantom tones correlated with magnitude of the Hilbert power spectra of the extra-aural component. Findings suggest a possible origin of phantom sounds that exists within the original signal, with potential implications for current models of non-linear cochlear mechanics and cortical dynamics in generating phantom percepts.

Enhanced tethered-flight duration and locomotor activity by overexpression of the human gene SOD1 in Drosophila motorneurons

Mutation of the human gene superoxide dismutase (hSOD1) is associated with the fatal neurodegenerative disease familial amyotrophic lateral sclerosis (Lou Gehrig’s disease). Selective overexpression of hSOD1 in Drosophila motorneurons increases lifespan to 140% of normal. The current study was designed to determine resistance to lifespan decline and failure of sensorimotor functions by overexpressing hSOD1 in Drosophila‘s motorneurons. First, we measured the ability to maintain continuous flight and wingbeat frequency (WBF) as a function of age (5 to 50 days). Flies overexpressing hSOD1 under the D42-GAL4 activator were able to sustain flight significantly longer than controls, with the largest effect observed in the middle stages of life. The hSOD1-expressed line also had, on average, slower wingbeat frequencies in late, but not early life relative to age-matched controls. Second, we examined locomotor (exploratory walking) behavior in late life when flies had lost the ability to fly (age ≥ 60 d). hSOD1-expressed flies showed significantly more robust walking activity relative to controls. Findings show patterns of functional decline dissimilar to those reported for other life-extended lines, and suggest that the hSOD1 gene not only delays death but enhances sensorimotor abilities critical to survival even in late life.

Psychophysical weights estimated for interaural cues in envelope slopes of amplitude-modulated waveforms

Current binaural theory contends that the auditory system encodes interaural delays in highpass-filtered complex sounds by phase locking to their slowly modulating envelopes. Spectrotemporal analysis of interaurally time-delayed highpass waveforms reveals the presence of a concomitant interaural level cue which could contribute to lateralization judgments. The current study systematically investigated the relative contribution of time and concomitant level cues carried by positive and negative envelope slopes of modified sinusoidally amplitude-modulated (SAM) high-frequency carriers. Psychophysical thresholds and observer decision weights were measured independently for the positive and negative modulation slopes of the acoustic signal. Decision weights were also measured to determine whether or not interaural delays are uniformly weighted at different temporal cycles of a SAM waveform. We found that lateralization of interaurally delayed SAM waveforms is influenced equally by ITDs in the rise and decay e...