Evolution of Local Circuit Neurons in Two Sensory Thalamic Nuclei in Amniotes.

Local circuit neurons are present in the thalamus of all vertebrates where they are considered inhibitory. They play an important role in computation and influence the transmission of information from the thalamus to the telencephalon. In mammals, the percentage of local circuit neurons in the dorsal lateral geniculate nucleus remains relatively constant across a variety of species. In contrast, the numbers of local circuit neurons in the ventral division of the medial geniculate body in mammals vary significantly depending on the species examined. To explain these observations, the numbers of local circuit neurons were investigated by reviewing the literature on this subject in these two nuclei in mammals and their respective homologs in sauropsids and by providing additional data on a crocodilian. Local circuit neurons are present in the dorsal geniculate nucleus of sauropsids just as is the case for this nucleus in mammals. However, sauropsids lack local circuits neurons in the auditory thalamic nuclei homologous to the ventral division of the medial geniculate body. A cladistic analysis of these results suggests that differences in the numbers of local circuit neurons in the dorsal lateral geniculate nucleus of amniotes reflect an elaboration of these local circuit neurons as a result of evolution from a common ancestor. In contrast, the numbers of local circuit neurons in the ventral division of the medial geniculate body changed independently in several mammalian lineages.

Processing symmetry between visual and auditory spatial representations in updating working memory.

Updating spatial representations in visual and auditory working memory relies on common processes, and the modalities should compete for attentional resources. If competition occurs, one type of spatial information is presumably weighted over the other, irrespective of sensory modality. This study used incompatible spatial information conveyed from two different cue modalities to examine relative dominance in memory updating. Participants mentally manoeuvred a designated target in a matrix according to visual or auditory stimuli that were presented simultaneously, to identify a terminal location. Prior to the navigation task, the relative perceptual saliences of the visual cues were manipulated to be equal, superior, or inferior to the auditory cues. The results demonstrate that visual and auditory information competed for attentional resources, such that visual/auditory guidance was impaired by incongruent cues delivered from the other modality. Although visual bias was generally observed in working-memory navigation, stimuli of relatively high salience interfered with or facilitated other stimuli regardless of modality, demonstrating the processing symmetry of spatial updating in visual and auditory spatial working memory. Furthermore, this processing symmetry can be identified during the encoding of sensory inputs into working-memory representations. The results imply that auditory spatial updating is comparable to visual spatial updating in that salient stimuli receive a high priority when selecting inputs and are used when tracking spatial representations.

Auditory timing-tuned neural responses in the human auditory cortices.

Perception of sub-second auditory event timing supports multisensory integration, and speech and music perception and production. Neural populations tuned for the timing (duration and rate) of visual events were recently described in several human extrastriate visual areas. Here we ask whether the brain also contains neural populations tuned for auditory event timing, and whether these are shared with visual timing. Using 7T fMRI, we measured responses to white noise bursts of changing duration and rate. We analyzed these responses using neural response models describing different parametric relationships between event timing and neural response amplitude. This revealed auditory timing-tuned responses in the primary auditory cortex, and auditory association areas of the belt, parabelt and premotor cortex. While these areas also showed tonotopic tuning for auditory pitch, pitch and timing preferences were not consistently correlated. Auditory timing-tuned response functions differed between these areas, though without clear hierarchical integration of responses. The similarity of auditory and visual timing tuned responses, together with the lack of overlap between the areas showing these responses for each modality, suggests modality-specific responses to event timing are computed similarly but from different sensory inputs, and then transformed differently to suit the needs of each modality.

Away Rotation Applications in Emergency Medicine: Medical Student Behaviors, Outcomes, and Stressors.

BACKGROUND: Completing an emergency medicine (EM) away rotation is integral to matching successfully into an EM residency program. The demand for EM away rotations (ARs) drives students to submit numerous applications without evidence-based recommendations to guide stakeholders on the approach or number to submit. OBJECTIVES: We conducted a survey study of EM-bound fourth-year medical students to gain insight into their AR application experiences, outcomes, and perceptions. METHODS: We distributed a 40-item questionnaire to EM applicants in Fall 2018 via e-mail through the Clerkship Directors in Emergency Medicine, Council of Residency Directors in EM, and Emergency Medicine Residents' Association listservs. Responses were evaluated using quantitative and qualitative analysis. Primary outcomes were the number of AR applications submitted and AR offers received by students. Secondary outcomes were students' self-assessment of their competitiveness, differences in AR application numbers by degree type, sources of student advising, and student perceptions of the AR application process. RESULTS: There were 253 respondents, consisting of 192 allopathic (MD) and 61 osteopathic (DO) medical students, who met the inclusion criteria, representing about 10% of the applicant pool. On average, students submitted 13.97 applications (95% confidence interval [CI] 11.59-16.35), received 3.25 offers (95% CI 3.01-3.49), and accepted 2.22 offers (95% CI 2.08-2.36). DO candidates submitted twice as many applications as MD candidates while experiencing a similar rate of offers received. Peer influence (n = 154, 61%), peer online advising networks (n = 83, 33%), and self-assessment (n = 114, 45%) were the most often reported causes of increased applications; cost (n = 104, 41%) and geographic limitations (n = 114, 45%) were the most often reported causes of decreased applications. Open-response analysis revealed frustration with lack of standardization (n = 44, 29.5%), insufficient transparency on available positions (n = 37, 24.8%), limited communication (n = 30, 20.1%), and cost (n = 12, 8.1%). CONCLUSIONS: This study showed that, as a whole, students received one away rotation offer for every four to five applications submitted. It clarified factors contributing to increased EM away rotation application submissions and associated stressors inherent in the application experience. Our findings offer insights to inform advising recommendations. They also suggest that stakeholders consider standardizing the process and improve communication over spot availability and application status.

The role of response set overlap for flexibility and cognitive control in auditory multitasking.

We developed a new variant of auditory task-switching in order to systematically investigate shifting and cognitive control in auditory task-switching and their relation to motor response overlap in a comprehensive way. In two experiments, participants classified either pitch or loudness of a simple tone as either low or high, hence, both tasks were constructed around a common underlying dimension ranging from low to high. In Experiment 1, response sets overlapped in both category and motor modality (both manual), whereas each task was related to a specific response category and motor response modality (manual vs. vocal) in Experiment 2. The data revealed reliable switch costs that were, contrary to our expectations, not reduced with reduced response set overlap. In addition, we found reliable congruency effects and their sequential modulation in both experiments with manual as well as vocal responses, and in the absence of competing motor activation (i.e., without motor response overlap). Congruency effects after auditory task switches were smaller when response sets did not overlap. Our data thus provides an important empirical generalization of known effects to auditory stimuli as well as with both manual and vocal responses. In addition, we demonstrated that reduced congruency effects after switches for non-overlapping response sets were due to the extent of overlap between different response sets in task-switching.

Audiovisual integration in macaque face patch neurons.

Primate social communication depends on the perceptual integration of visual and auditory cues, reflected in the multimodal mixing of sensory signals in certain cortical areas. The macaque cortical face patch network, identified through visual, face-selective responses measured with fMRI, is assumed to contribute to visual social interactions. However, whether face patch neurons are also influenced by acoustic information, such as the auditory component of a natural vocalization, remains unknown. Here, we recorded single-unit activity in the anterior fundus (AF) face patch, in the superior temporal sulcus, and anterior medial (AM) face patch, on the undersurface of the temporal lobe, in macaques presented with audiovisual, visual-only, and auditory-only renditions of natural movies of macaques vocalizing. The results revealed that 76% of neurons in face patch AF were significantly influenced by the auditory component of the movie, most often through enhancement of visual responses but sometimes in response to the auditory stimulus alone. By contrast, few neurons in face patch AM exhibited significant auditory responses or modulation. Control experiments in AF used an animated macaque avatar to demonstrate, first, that the structural elements of the face were often essential for audiovisual modulation and, second, that the temporal modulation of the acoustic stimulus was more important than its frequency spectrum. Together, these results identify a striking contrast between two face patches and specifically identify AF as playing a potential role in the integration of audiovisual cues during natural modes of social communication.

Modulation of early auditory processing by visual information: Prediction or bimodal integration?

What happens if a visual cue misleads auditory expectations? Previous studies revealed an early visuo-auditory incongruency effect, so-called incongruency response (IR) of the auditory event-related brain potential (ERP), occurring 100 ms after onset of the sound being incongruent to the preceding visual cue. So far, this effect has been ascribed to reflect the mismatch between auditory sensory expectation activated by visual predictive information and the actual sensory input. Thus, an IR should be confined to an asynchronous presentation of visual cue and sound. Alternatively, one could argue that frequently presented congruent visual-cue-sound combinations are integrated into a bimodal representation whereby violation of the visual-auditory relationship results in a bimodal feature mismatch (the IR should be obtained with asynchronous and with synchronous presentation). In an asynchronous condition, an either high-pitched or low-pitched sound was preceded by a visual note symbol presented above or below a fixation cross (90% congruent; 10% incongruent), while in a synchronous condition, both were presented simultaneously. High-pitched and low-pitched sounds were presented with different probabilities (83% vs. 17%) to form a strong association between bimodal stimuli. In both conditions, tones with pitch incongruent with the location of the note symbols elicited incongruency effects in the N2 and P3 ERPs; however, the IR was only elicited in the asynchronous condition. This finding supports the sensorial prediction error hypothesis stating that the amplitude of the auditory ERP 100 ms after sound onset is enhanced in response to unexpected compared with expected but otherwise identical sounds.

Cochlea to categories: The spatiotemporal dynamics of semantic auditory representations.

How does the auditory system categorize natural sounds? Here we apply multimodal neuroimaging to illustrate the progression from acoustic to semantically dominated representations. Combining magnetoencephalographic (MEG) and functional magnetic resonance imaging (fMRI) scans of observers listening to naturalistic sounds, we found superior temporal responses beginning ∼55 ms post-stimulus onset, spreading to extratemporal cortices by ∼100 ms. Early regions were distinguished less by onset/peak latency than by functional properties and overall temporal response profiles. Early acoustically-dominated representations trended systematically toward category dominance over time (after ∼200 ms) and space (beyond primary cortex). Semantic category representation was spatially specific: Vocalizations were preferentially distinguished in frontotemporal voice-selective regions and the fusiform; scenes and objects were distinguished in parahippocampal and medial place areas. Our results are consistent with real-world events coded via an extended auditory processing hierarchy, in which acoustic representations rapidly enter multiple streams specialized by category, including areas typically considered visual cortex.

Cortical processing of distracting speech in noisy auditory scenes depends on perceptual demand.

Selective attention is essential for the processing of multi-speaker auditory scenes because they require the perceptual segregation of the relevant speech ("target") from irrelevant speech ("distractors"). For simple sounds, it has been suggested that the processing of multiple distractor sounds depends on bottom-up factors affecting task performance. However, it remains unclear whether such dependency applies to naturalistic multi-speaker auditory scenes. In this study, we tested the hypothesis that increased perceptual demand (the processing requirement posed by the scene to separate the target speech) reduces the cortical processing of distractor speech thus decreasing their perceptual segregation. Human participants were presented with auditory scenes including three speakers and asked to selectively attend to one speaker while their EEG was acquired. The perceptual demand of this selective listening task was varied by introducing an auditory cue (interaural time differences, ITDs) for segregating the target from the distractor speakers, while acoustic differences between the distractors were matched in ITD and loudness. We obtained a quantitative measure of the cortical segregation of distractor speakers by assessing the difference in how accurately speech-envelope following EEG responses could be predicted by models of averaged distractor speech versus models of individual distractor speech. In agreement with our hypothesis, results show that interaural segregation cues led to improved behavioral word-recognition performance and stronger cortical segregation of the distractor speakers. The neural effect was strongest in the δ-band and at early delays (0 - 200 ms). Our results indicate that during low perceptual demand, the human cortex represents individual distractor speech signals as more segregated. This suggests that, in addition to purely acoustical properties, the cortical processing of distractor speakers depends on factors like perceptual demand.

An online headphone screening test based on dichotic pitch.

Online experimental platforms can be used as an alternative to, or complement, lab-based research. However, when conducting auditory experiments via online methods, the researcher has limited control over the participants' listening environment. We offer a new method to probe one aspect of that environment, headphone use. Headphones not only provide better control of sound presentation but can also "shield" the listener from background noise. Here we present a rapid (< 3 min) headphone screening test based on Huggins Pitch (HP), a perceptual phenomenon that can only be detected when stimuli are presented dichotically. We validate this test using a cohort of "Trusted" online participants who completed the test using both headphones and loudspeakers. The same participants were also used to test an existing headphone test (AP test; Woods et al., 2017, Attention Perception Psychophysics). We demonstrate that compared to the AP test, the HP test has a higher selectivity for headphone users, rendering it as a compelling alternative to existing methods. Overall, the new HP test correctly detects 80% of headphone users and has a false-positive rate of 20%. Moreover, we demonstrate that combining the HP test with an additional test-either the AP test or an alternative based on a beat test (BT)-can lower the false-positive rate to ~ 7%. This should be useful in situations where headphone use is particularly critical (e.g., dichotic or spatial manipulations). Code for implementing the new tests is publicly available in JavaScript and through Gorilla (gorilla.sc).

Magnetic resonance diffusion tensor tractography of a midbrain auditory circuit in Alligator.

Knowledge of brain circuitry is critical for understanding the organization, function, and evolution of central nervous systems. Most commonly, brain connections have been elucidated using histological and experimental methods that require animal sacrifice. On the other hand, magnetic resonance diffusion tensor imaging and associated tractography have emerged as a preferred method to noninvasively visualize brain white matter tracts. However, existing studies have primarily examined large, heavily myelinated fiber tracts. Whether tractography can visualize fiber bundles that contain thin and poorly myelinated axons is uncertain. To address this question, the midbrain auditory pathway to the thalamus was investigated in Alligator. This species was chosen because of its evolutionary importance as it is the reptilian group most closely related to birds and because its brain contains many thin and poorly myelinated tracts. Furthermore, this auditory pathway is well documented in other reptiles, including a related crocodilian. Histological observations and experimental determination of anterograde connections confirmed this path in Alligator. Tractography identified these tracts in Alligator and provided a 3-dimensional picture that accurately identified the neural elements of this circuit. In addition, tractography identified one possible unrecognized pathway. These results demonstrate that tractography can visualize circuits containing thin, poorly myelinated fibers. These findings open the door for future studies to examine these types of pathways in other vertebrates.

Auditory Perceptual History Is Propagated through Alpha Oscillations.

Perception is a proactive, "predictive" process, in which the brain relies, at least in part, on accumulated experience to make best guesses about the world to test against sensory data, updating the guesses as new experience is acquired. Using novel behavioral methods, the present study demonstrates the role of alpha rhythms in communicating past perceptual experience. Participants were required to discriminate the ear of origin of brief sinusoidal tones that were presented monaurally at random times within a burst of uncorrelated dichotic white noise masks. Performance was not constant but varied with delay after noise onset in an oscillatory manner at about 9 Hz (alpha rhythm). Importantly, oscillations occurred only for trials preceded by a target tone to the same ear, either on the previous trial or two trials back. These results suggest that communication of perceptual history generates neural oscillations within specific perceptual circuits, strongly implicating behavioral oscillations in predictive perception and with formation of working memory.

Ternary meter from spatial sounds: Differences in neural entrainment between musicians and non-musicians.

The present study explores the relationship between the rhythmic structure of music and the spatial dimension of sound. We study how the brain interacts with spatially-separated sounds to build up a metrical structure. Participants listened to sequences of isochronous sounds that came from different positions on the azimuth plane: 0° (control condition), ±30°, ±60° or ±90° (spatial conditions). Ternary meter was signaled by the alternation of one sound on one side and two sounds on the symmetrical side. In Experiment 1, musicians and non-musicians paid attention to the spatial sounds. In Experiment 2, participants paid attention to a visual distractor. We recorded their electroencephalograms and performed frequency-tagging analyses. In both experiments, the isochronous beat elicited steady-state evoked-potentials at the frequency of the beat (2.4 Hz). While in Experiment 1 the alternation produced clear responses at the frequency of the ternary meter (0.8 Hz), in Experiment 2 these responses were only significant in the Spatial 90° condition, and mainly in musicians. This suggests that top-down attentional mechanisms are in play for meter induction. Besides, musicians showed stronger responses to beat and meter than non-musicians, suggesting that formal musical training enhances the neural entrainment to spatially-defined rhythms.

The impact of task difficulty on the lateralization of processing in the human auditory cortex.

Perception of complex auditory stimuli like speech requires the simultaneous processing of different fundamental acoustic parameters. The contribution of left and right auditory cortex (AC) in the processing of these parameters differs. In addition, activity within the AC can vary positively or negatively with task performance depending on the type of task. This might affect the allocation of processing to the left and right AC. Here we studied with functional magnetic resonance imaging the impact of task difficulty on the degree of involvement of the left and right AC in two tasks that have previously been shown to differ in hemispheric involvement: categorization and sequential comparison of the direction of frequency modulations (FM). Task difficulty was manipulated by changing the speed of modulation and by that the frequency range covered by the FM. To study the impact of task-difficulty despite covarying the stimulus parameters, we utilized the contralateral noise procedure that allows comparing AC activation unconfounded by bottom-up driven activity. The easiest conditions confirmed the known right AC involvement during the categorization task and the left AC involvement during the comparison task. The involvement of the right AC increased with increasing task difficulty for both tasks presumably due to the common task component of categorizing FM direction. The involvement of left AC varied with task difficulty depending on the task. Thus, task difficulty has a strong impact on lateralized processing in AC. This connection must be taken into account when interpreting future results on lateralized processing in the AC.

The Aesthetic Preference for Nature Sounds Depends on Sound Object Recognition.

People across the world seek out beautiful sounds in nature, such as a babbling brook or a nightingale song, for positive human experiences. However, it is unclear whether this positive aesthetic response is driven by a preference for the perceptual features typical of nature sounds versus a higher-order association of nature with beauty. To test these hypotheses, participants provided aesthetic judgments for nature and urban soundscapes that varied on ease of recognition. Results demonstrated that the aesthetic preference for nature soundscapes was eliminated for the sounds hardest to recognize, and moreover the relationship between aesthetic ratings and several measured acoustic features significantly changed as a function of recognition. In a follow-up experiment, requiring participants to classify these difficult-to-identify sounds into nature or urban categories resulted in a robust preference for nature sounds and a relationship between aesthetic ratings and our measured acoustic features that was more typical of easy-to-identify sounds. This pattern of results was replicated with computer-generated artificial noises, which acoustically shared properties with the nature and urban soundscapes but by definition did not come from these environments. Taken together, these results support the conclusion that the recognition of a sound as either natural or urban dynamically organizes the relationship between aesthetic preference and perceptual features and that these preferences are not inherent to the acoustic features. Implications for nature's role in cognitive and affective restoration are discussed.

Musical instrument categorization is highly sensitive to spectral properties of earlier sounds.

Auditory perception is shaped by spectral properties of surrounding sounds. For example, when spectral properties differ between earlier (context) and later (target) sounds, this can produce spectral contrast effects (SCEs; i.e., categorization boundary shifts) that bias perception of later sounds. SCEs affect perception of speech and nonspeech sounds alike (Stilp Alexander, Kiefte, & Kluender in Attention, Perception, & Psychophysics, 72(2), 470-480, 2010). When categorizing speech sounds, SCE magnitudes increased linearly with greater spectral differences between contexts and target sounds (Stilp, Anderson, & Winn in Journal of the Acoustical Society of America, 137(6), 3466-3476, 2015; Stilp & Alexander in Proceedings of Meetings on Acoustics, 26, 2016; Stilp & Assgari in Journal of the Acoustical Society of America, 141(2), EL153-EL158, 2017). The present experiment tested whether this acute context sensitivity generalized to nonspeech categorization. Listeners categorized musical instrument target sounds that varied from French horn to tenor saxophone. Before each target, listeners heard a 1-second string quintet sample processed by filters that reflected part of (25%, 50%, 75%) or the full (100%) difference between horn and saxophone spectra. Larger filter gains increased spectral distinctness across context and target sounds, and resulting SCE magnitudes increased linearly, parallel to speech categorization. Thus, a highly sensitive relationship between context spectra and target categorization appears to be fundamental to auditory perception.

Reduced late mismatch negativity and auditory sustained potential to rule-based patterns in schizophrenia.

Complex rule-based auditory processing is abnormal in individuals with long-term schizophrenia (SZ), as demonstrated by reduced mismatch negativity (MMN) to deviants in rule-based patterns and reduced auditory sustained potential (ASP) that appears when grouping tones together. Together, this suggests deficits later in the auditory processing hierarchy in Sz. Here, MMN and ASP were elicited by deviations from a complex zig-zag pitch pattern that cannot be predicted by simple linear rules. Twenty-seven SZ and 26 matched healthy controls (HC) participated. Frequent groups of patterns contained eight tones that zig-zagged in a two-up one-down pitch-based paradigm. There were two deviant patterns: the final tone was either higher in pitch than expected (creating a jump in pitch) or was repeated. Simple MMN to pitch-deviants among repetitive tones was measured for comparison. Sz exhibited a smaller pitch MMN compared to HC as expected. HC produced a late MMN in response to the repeat and jump-deviant and a larger ASP to the standard group of tones, all of which were significantly blunted in SZ. In Sz, the amplitude of the late complex MMN was related to neuropsychological functioning, whereas ASP was not. ASP and late MMN did not significantly correlate in HC or in Sz, suggesting that they are not dependent on one another and may originate within distinct processing streams. Together, this suggests multiple deficits later in the auditory sensory-perceptual hierarchy in Sz, with impairments evident in both segmentation and deviance detection abilities.

Adaptive and Selective Time Averaging of Auditory Scenes.

To overcome variability, estimate scene characteristics, and compress sensory input, perceptual systems pool data into statistical summaries. Despite growing evidence for statistical representations in perception, the underlying mechanisms remain poorly understood. One example of such representations occurs in auditory scenes, where background texture appears to be represented with time-averaged sound statistics. We probed the averaging mechanism using "texture steps"-textures containing subtle shifts in stimulus statistics. Although generally imperceptible, steps occurring in the previous several seconds biased texture judgments, indicative of a multi-second averaging window. Listeners seemed unable to willfully extend or restrict this window but showed signatures of longer integration times for temporally variable textures. In all cases the measured timescales were substantially longer than previously reported integration times in the auditory system. Integration also showed signs of being restricted to sound elements attributed to a common source. The results suggest an integration process that depends on stimulus characteristics, integrating over longer extents when it benefits statistical estimation of variable signals and selectively integrating stimulus components likely to have a common cause in the world. Our methodology could be naturally extended to examine statistical representations of other types of sensory signals.

Acoustic and higher-level representations of naturalistic auditory scenes in human auditory and frontal cortex.

Often, in everyday life, we encounter auditory scenes comprising multiple simultaneous sounds and succeed to selectively attend to only one sound, typically the most relevant for ongoing behavior. Studies using basic sounds and two-talker stimuli have shown that auditory selective attention aids this by enhancing the neural representations of the attended sound in auditory cortex. It remains unknown, however, whether and how this selective attention mechanism operates on representations of auditory scenes containing natural sounds of different categories. In this high-field fMRI study we presented participants with simultaneous voices and musical instruments while manipulating their focus of attention. We found an attentional enhancement of neural sound representations in temporal cortex - as defined by spatial activation patterns - at locations that depended on the attended category (i.e., voices or instruments). In contrast, we found that in frontal cortex the site of enhancement was independent of the attended category and the same regions could flexibly represent any attended sound regardless of its category. These results are relevant to elucidate the interacting mechanisms of bottom-up and top-down processing when listening to real-life scenes comprised of multiple sound categories.

Effect of language experience on selective auditory attention: An event-related potential study.

Dual language experience has typically been shown to improve various executive control functions. We investigated with event-related brain potentials (ERPs) recorded from early (natively) bilingual speakers and control participants whether it also affects auditory selective attention. We delivered to our participants two tone streams, one to the left and one to the right ear. Both streams consisted of standard tones and two types of infrequent deviant tones which had either an enhanced duration or intensity. The participants were instructed to attend either to the right or left stream and to detect longer-duration deviants in the attended stream. The results showed that the early bilinguals did not outperform the controls in target detection accuracy or speed. However, the late portion of the attention-related ERP modulation (the negative difference, Nd) was larger over the left hemisphere in the early bilinguals than in the controls, suggesting that the maintenance of selective attention or further processing of selectively attended sounds is enhanced in the bilinguals. Moreover, the late reorienting negativity (RON) in response to intensity-deviant tones was larger in the bilinguals, suggesting more efficient disengagement of attention from distracting auditory events. Hence, our results demonstrate that brain responses associated with certain aspects of auditory attention are enhanced in the bilingual adults, indicating that early dual language exposure modulates the neuronal responsiveness of auditory modality.