Confirming an antiphasic bicyclic pattern of forward entrainment in signal detection: A reanalysis of Sun et al. (2021).

Forward entrainment refers to that part of the entrainment process that persists after termination of an entraining stimulus. Hickok et al. (2015) reported forward entrainment in signal detection that lasted for two post-stimulus cycles. In a recent paper, Sun et al. (2021) reported new data which suggested an absence of entrainment effects (Eur. J. Neurosci, 1-18, doi.org/10.1111/ejn.15367). Here we show that when Sun et al.'s data are analysed using unbiased detection-theoretic measures, a clear antiphasic bicyclic pattern of entrainment is observed. We further show that the measure of entrainment strength used by Sun et al., the normalized Fourier transform of performance curves, is not only erroneously calculated but is also unreliable in estimating entrainment strength due to signal-processing artifacts.

A critical analysis of Lin et al.'s (2021) failure to observe forward entrainment in pitch discrimination.

Forward entrainment refers to that part of the entrainment process that outlasts the entraining stimulus. Several studies have demonstrated psychophysical forward entrainment in a pitch-discrimination task. In a recent paper, Lin et al. (2021) challenged these findings by demonstrating that a sequence of 4 entraining pure tones does not affect the ability to determine whether a frequency modulated pulse, presented after termination of the entraining sequence, has swept up or down in frequency. They concluded that rhythmic sequences do not facilitate pitch discrimination. Here, we describe several methodological and stimulus design flaws in Lin et al.'s study that may explain their failure to observe forward entrainment in pitch discrimination.

Detection of dynamic changes in interaural delay by older adults (L).

The ability of older adults (48 to 72) with relatively intact low-frequency hearing to detect the motion of an acoustic source was investigated using dynamically varying interaural delays. Thresholds were measured using a single-interval two-alternative forced-choice task in which listeners determined if the sound source was moving or stationary. Motion thresholds were significantly larger than stationary localization thresholds. No correlation was observed between age and motion-detection ability for the age range tested. An interesting finding was that there were similar thresholds for older and younger adults. Results suggest reliance on dominant low-frequency binaural timing cues unaffected by high-frequency hearing loss in older adults.

The Rhythm of Perception: Entrainment to Acoustic Rhythms Induces Subsequent Perceptual Oscillation.

Acoustic rhythms are pervasive in speech, music, and environmental sounds. Recent evidence for neural codes representing periodic information suggests that they may be a neural basis for the ability to detect rhythm. Further, rhythmic information has been found to modulate auditory-system excitability, which provides a potential mechanism for parsing the acoustic stream. Here, we explored the effects of a rhythmic stimulus on subsequent auditory perception. We found that a low-frequency (3 Hz), amplitude-modulated signal induces a subsequent oscillation of the perceptual detectability of a brief nonperiodic acoustic stimulus (1-kHz tone); the frequency but not the phase of the perceptual oscillation matches the entrained stimulus-driven rhythmic oscillation. This provides evidence that rhythmic contexts have a direct influence on subsequent auditory perception of discrete acoustic events. Rhythm coding is likely a fundamental feature of auditory-system design that predates the development of explicit human enjoyment of rhythm in music or poetry.

Orthogonal acoustic dimensions define auditory field maps in human cortex.

The functional organization of human auditory cortex has not yet been characterized beyond a rudimentary level of detail. Here, we use functional MRI to measure the microstructure of orthogonal tonotopic and periodotopic gradients forming complete auditory field maps (AFMs) in human core and belt auditory cortex. These AFMs show clear homologies to subfields of auditory cortex identified in nonhuman primates and in human cytoarchitectural studies. In addition, we present measurements of the macrostructural organization of these AFMs into "clover leaf" clusters, consistent with the macrostructural organization seen across human visual cortex. As auditory cortex is at the interface between peripheral hearing and central processes, improved understanding of the organization of this system could open the door to a better understanding of the transformation from auditory spectrotemporal signals to higher-order information such as speech categories.

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.

Enhanced optomotor efficiency by expression of the human gene superoxide dismutase primarily in Drosophila motorneurons.

Mutation of the human gene superoxide dismutase (hSOD1) triggers the fatal neurodegenerative motorneuron disorder, familial amyotrophic lateral sclerosis (ALS or Lou Gehrig's 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.

Forward entrainment: Psychophysics, neural correlates, and function.

We define forward entrainment as that part of behavioral or neural entrainment that outlasts the entraining stimulus. In this review, we examine conditions under which one may optimally observe forward entrainment. In Part 1, we review and evaluate studies that have observed forward entrainment using a variety of psychophysical methods (detection, discrimination, and reaction times), different target stimuli (tones, noise, and gaps), different entraining sequences (sinusoidal, rectangular, or sawtooth waveforms), a variety of physiological measures (MEG, EEG, ECoG, CSD), in different modalities (auditory and visual), across modalities (audiovisual and auditory-motor), and in different species. In Part 2, we describe those experimental conditions that place constraints on the magnitude of forward entrainment, including an evaluation of the effects of signal uncertainty and attention, temporal envelope complexity, signal-to-noise ratio (SNR), rhythmic rate, prior experience, and intersubject variability. In Part 3 we theorize on potential mechanisms and propose that forward entrainment may instantiate a dynamic auditory afterimage that lasts a fraction of a second to minimize prediction error in signal processing.

Informational Masking in Aging and Brain-lesioned Individuals.

Auditory stream segregation and informational masking were investigated in brain-lesioned individuals, age-matched controls with no neurological disease, and young college-age students. A psychophysical paradigm known as rhythmic masking release (RMR) was used to examine the ability of participants to identify a change in the rhythmic sequence of 20-ms Gaussian noise bursts presented through headphones and filtered through generalized head-related transfer functions to produce the percept of an externalized auditory image (i.e., a 3D virtual reality sound). The target rhythm was temporally interleaved with a masker sequence comprising similar noise bursts in a manner that resulted in a uniform sequence with no information remaining about the target rhythm when the target and masker were presented from the same location (an impossible task). Spatially separating the target and masker sequences allowed participants to determine if there was a change in the target rhythm midway during its presentation. RMR thresholds were defined as the minimum spatial separation between target and masker sequences that resulted in 70.7% correct-performance level in a single-interval 2-alternative forced-choice adaptive tracking procedure. The main findings were (1) significantly higher RMR thresholds for individuals with brain lesions (especially those with damage to parietal areas) and (2) a left-right spatial asymmetry in performance for lesion (but not control) participants. These findings contribute to a better understanding of spatiotemporal relations in informational masking and the neural bases of auditory scene analysis.

Functional anatomy of language and music perception: temporal and structural factors investigated using functional magnetic resonance imaging.

Language and music exhibit similar acoustic and structural properties, and both appear to be uniquely human. Several recent studies suggest that speech and music perception recruit shared computational systems, and a common substrate in Broca's area for hierarchical processing has recently been proposed. However, this claim has not been tested by directly comparing the spatial distribution of activations to speech and music processing within subjects. In the present study, participants listened to sentences, scrambled sentences, and novel melodies. As expected, large swaths of activation for both sentences and melodies were found bilaterally in the superior temporal lobe, overlapping in portions of auditory cortex. However, substantial nonoverlap was also found: sentences elicited more ventrolateral activation, whereas the melodies elicited a more dorsomedial pattern, extending into the parietal lobe. Multivariate pattern classification analyses indicate that even within the regions of blood oxygenation level-dependent response overlap, speech and music elicit distinguishable patterns of activation. Regions involved in processing hierarchical aspects of sentence perception were identified by contrasting sentences with scrambled sentences, revealing a bilateral temporal lobe network. Music perception showed no overlap whatsoever with this network. Broca's area was not robustly activated by any stimulus type. Overall, these findings suggest that basic hierarchical processing for music and speech recruits distinct cortical networks, neither of which involves Broca's area. We suggest that previous claims are based on data from tasks that tap higher-order cognitive processes, such as working memory and/or cognitive control, which can operate in both speech and music domains.

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.

Functionally distinct regions for spatial processing and sensory motor integration in the planum temporale.

There has been much debate recently over the functional role played by the planum temporale (PT) within the context of the dorsal auditory processing stream. Some studies indicate that regions in the PT support spatial hearing and other auditory functions, whereas others demonstrate sensory-motor response properties. This multifunctionality has led to the claim that the PT is performing a common computational pattern matching operation, then routing the signals (spatial, object, sensory-motor) into an appropriate processing stream. An alternative possibility is that the PT is functionally subdivided with separate regions supporting various functions. We assess this possibility using a within subject fMRI block design. DTI data were also collected to examine connectivity. There were four auditory conditions: stationary noise, moving noise, listening to pseudowords, and shadowing pseudowords (covert repetition). Contrasting the shadow and listen conditions should activate regions specific to sensory-motor processes, while contrasting the stationary and moving noise conditions should activate regions involved in spatial hearing. Subjects (N = 16) showed greater activation for shadowing in left posterior PT, area Spt, when the shadow and listen conditions were contrasted. The motion vs. stationary noise contrast revealed greater activation in a more medial and anterior portion of left PT. Seeds from these two contrasts were then used to guide the DTI analysis in an examination of connectivity via streamline tractography, which revealed different patterns of connectivity. Findings support a heterogeneous model of the PT, with functionally distinct regions for sensory-motor integration and processes involved in auditory spatial perception.

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.

Auditory spatial and object processing in the human planum temporale: no evidence for selectivity.

Although it is generally acknowledged that at least two processing streams exist in the primate cortical auditory system, the function of the posterior dorsal stream is a topic of much debate. Recent studies have reported selective activation to auditory spatial change in portions of the human planum temporale (PT) relative to nonspatial stimuli such as pitch changes or complex acoustic patterns. However, previous work has suggested that the PT may be sensitive to another kind of nonspatial variable, namely, the number of auditory objects simultaneously presented in the acoustic signal. The goal of the present fMRI experiment was to assess whether any portion of the PT showed spatial selectivity relative to manipulations of the number of auditory objects presented. Spatially sensitive regions in the PT were defined by comparing activity associated with listening to an auditory object (speech from a single talker) that changed location with one that remained stationary. Activity within these regions was then examined during a nonspatial manipulation: increasing the number of objects (talkers) from one to three. The nonspatial manipulation modulated activity within the "spatial" PT regions. No region within the PT was found to be selective for spatial or object processing. We suggest that previously documented spatial sensitivity in the PT reflects auditory source separation using spatial cues rather than spatial processing per se.

The rhythm of attention: Perceptual modulation via rhythmic entrainment is lowpass and attention mediated.

Modulation patterns are known to carry critical predictive cues to signal detection in complex acoustic environments. The current study investigated the persistence of masker modulation effects on postmodulation detection of probe signals. Hickok, Farahbod, and Saberi (Psychological Science, 26, 1006-1013, 2015) demonstrated that thresholds for a tone pulse in stationary noise follow a predictable periodic pattern when preceded by a 3-Hz amplitude modulated masker. They found entrainment of detection patterns to the modulation envelope lasting for approximately two cycles after termination of modulation. The current study extends these results to a wide range of modulation rates by mapping the temporal modulation transfer function for persistent modulatory effects. We found significant entrainment to modulation rates of 2 and 3 Hz, a weaker effect at 5 Hz, and no entrainment at higher rates (8 to 32 Hz). The effect seems critically dependent on attentional mechanisms, requiring temporal and level uncertainty of the probe signal. Our findings suggest that the persistence of modulatory effects on signal detection is lowpass in nature and attention based.

The Effect of Topical Vitamin A and E on Ischemic Random Skin Flap Survival.

BACKGROUND: Ischemia of skin flaps is an important complication in reconstructive surgery. This study evaluated the efficacy of topical vitamins A and E on improving flap survival. METHODS: Twenty-four white-albino male rats were randomly divided into two groups of treatment and control. Standard rectangular, distally based dorsal random pattern skin flap was elevated. Intra-peritoneal cephazoline was administered to prevent any unexpected infection. No pharmaceutical agent was administered for the control group, but pure vaseline ointment. In treatment group, vaseline plus vitamins A and E were administrated daily after surgery for 10 days. The rats were evaluated on the 10th day after surgery for viable and necrotic portions of the flaps. RESULTS: The mean values of necrosis in the flaps were 625±189.56 and 920.00±247.31 in the treatment and control groups, respectively. Vaseline plus vitamins increased flap survival significantly. CONCLUSION: Topical vitamins A and E may be effective pharmaceutical agents to increase viability of random skin flaps in rats. They can be added to vasoactive topical agents to reach better results.

Dissociation of procedural and semantic memory in absolute-pitch processing.

We describe two memory-retrieval systems in absolute-pitch (AP) processing and propose existence of a universal internal pitch template to which subpopulations of musicians selectively gain access through the two systems. In Experiment I, AP and control musicians adjusted the frequency of a pure tone to match the pitch of a visually displayed randomly selected musical note. In Experiment II the same subjects vocally produced within 2s the pitch associated with a randomly selected musical note label. AP musicians, but not controls, were highly accurate in frequency matching. Surprisingly, both AP and non-AP groups were extremely accurate in voicing the target pitch as determined from an FFT of the recorded voiced notes (i.e., sigma=0.97, 0.90 semitones, respectively). Spectrogram analysis showed that notes voiced by non-AP musicians are accurate from onset of voicing suggesting that pitch accuracy does not result from an auditory-motor feedback loop. Findings support existence of two memory-retrieval systems for musical pitch: a semantic associative form of memory used by AP musicians, and a more widespread form of procedural memory which allows precise access to internal pitch representations through the vocal-motor system.

An event-related fMRI study of auditory motion perception: no evidence for a specialized cortical system.

The existence of a specialized human cortical area for the processing of auditory motion is still a matter of debate. Initial functional imaging studies identified the planum temporale as being motion selective. Recent data contrasting spatially varying stationary stimuli with moving stimuli found no difference in the amount of activation between the two types of stimuli in the planum temporale. The present study re-examines this issue using an event-related paradigm. Ten subjects were scanned while listening to pairs of stimuli that were either both moving or both stationary. Consistent with the aforementioned study, we found no difference in the activation levels in the planum temporale when comparing motion and stationary conditions.

Temporal integration in absolute identification of musical pitch.

The effect of stimulus duration on absolute identification of musical pitch was measured in a single-interval 12-alternative forced-choice task. Stimuli consisted of pure tones selected randomly on each trial from a set of 60 logarithmically spaced musical note frequencies from 65.4 to 1975.5Hz (C2-B6). Stimulus durations were 5, 10, 25, 50, 100, and 1000ms. Six absolute-pitch musicians identified the pitch of pure tones without feedback, reference sounds, or practice trials. Results showed that a 5ms stimulus is sufficient for producing statistically significant above chance performance. Performance monotonically increased up to the longest duration tested (1000ms). Higher octave stimuli produced better performance, though the rate of improvement declined with increasing octave number. Normalization by the number of waveform cycles showed that 4cycles are sufficient for absolute-pitch identification. Restricting stimuli to a fixed-cycle waveform instead of a fixed-duration still produced monotonic improvements in performance as a function of stimulus octave, demonstrating that better performance at higher frequencies does not exclusively result from a larger number of waveform cycles. Several trends in the data were well predicted by an autocorrelation model of pitch extraction, though the model outperformed observed performance at short durations suggesting an inability to make optimal use of available periodicity information in very brief tones.

Robustness of speech intelligibility at moderate levels of spectral degradation.

The current study investigated how amplitude and phase information differentially contribute to speech intelligibility. Listeners performed a word-identification task after hearing spectrally degraded sentences. Each stimulus was degraded by first dividing it into segments, then the amplitude and phase components of each segment were decorrelated independently to various degrees relative to those of the original segment. Segments were then concatenated into their original sequence to present to the listener. We used three segment lengths: 30 ms (phoneme length), 250 ms (syllable length), and full sentence (non-segmented). We found that for intermediate spectral correlation values, segment length is generally inconsequential to intelligibility. Overall, intelligibility was more adversely affected by phase-spectrum decorrelation than by amplitude-spectrum decorrelation. If the phase information was left intact, decorrelating the amplitude spectrum to intermediate values had no effect on intelligibility. If the amplitude information was left intact, decorrelating the phase spectrum to intermediate values significantly degraded intelligibility. Some exceptions to this rule are described. These results delineate the range of amplitude- and phase-spectrum correlations necessary for speech processing and its dependency on the temporal window of analysis (phoneme or syllable length). Results further point to the robustness of speech information in environments that acoustically degrade cues to intelligibility (e.g., reverberant or noisy environments).