Ariane E. Rhone

Direct physiologic evidence of a heteromodal convergence region for proper naming in human left anterior temporal lobe.

Retrieving the names of friends, loved ones, and famous people is a fundamental human ability. This ability depends on the left anterior temporal lobe (ATL), where lesions can be associated with impaired naming of people regardless of modality (e.g., picture or voice). This finding has led to the idea that the left ATL is a modality-independent convergence region for proper naming. Hypotheses for how proper-name dispositions are organized within the left ATL include both a single modality-independent (heteromodal) convergence region and spatially discrete modality-dependent (unimodal) regions. Here we show direct electrophysiologic evidence that the left ATL is heteromodal for proper-name retrieval. Using intracranial recordings placed directly on the surface of the left ATL in human subjects, we demonstrate nearly identical responses to picture and voice stimuli of famous U.S. politicians during a naming task. Our results demonstrate convergent and robust large-scale neurophysiologic responses to picture and voice naming in the human left ATL. This finding supports the idea of heteromodal (i.e., transmodal) dispositions for proper naming in the left ATL.

Differential activation of human core, non-core and auditory-related cortex during speech categorization tasks as revealed by intracranial recordings.

Speech perception requires that sounds be transformed into speech-related objects with lexical and semantic meaning. It is unclear at what level in the auditory pathways this transformation emerges. Primary auditory cortex has been implicated in both representation of acoustic sound attributes and sound objects. While non-primary auditory cortex located on the posterolateral superior temporal gyrus (PLST) is clearly involved in acoustic-to-phonetic pre-lexical representations, it is unclear what role this region plays in auditory object formation. Additional data support the importance of prefrontal cortex in the formation of auditory objects, while other data would implicate this region in auditory object selection. To help clarify the respective roles of auditory and auditory-related cortex in the formation and selection of auditory objects, we examined high gamma activity simultaneously recorded directly from Heschls gyrus (HG), PLST and prefrontal cortex, while subjects performed auditory semantic detection tasks. Subjects were patients undergoing evaluation for treatment of medically intractable epilepsy. We found that activity in posteromedial HG and early activity on PLST was robust to sound stimuli regardless of their context, and minimally modulated by tasks. Later activity on PLST could be strongly modulated by semantic context, but not by behavioral performance. Activity within prefrontal cortex also was related to semantic context, and did co-vary with behavior. We propose that activity in posteromedial HG and early activity on PLST primarily reflect the representation of spectrotemporal sound attributes. Later activity on PLST represents a pre-lexical processing stage and is an intermediate step in the formation of word objects. Activity in prefrontal cortex appears directly involved in word object selection. The roles of other auditory and auditory-related cortical areas in the formation of word objects remain to be explored.

Mapping the temporal pole with a specialized electrode array: technique and preliminary results

Temporopolar cortex plays a crucial role in the pathogenesis of temporal lobe epilepsy and subserves important cognitive functions. Because of its shape and position in the middle cranial fossa, complete electrode coverage of the temporal pole (TP) is difficult to achieve using existing devices. We designed a novel TP electrode array that conforms to the surface of temporopolar cortex and achieves dense electrode coverage of this important brain region. A multi-pronged electrode array was designed that can be placed over the surface of the TP using a straightforward insertion technique. Twelve patients with medically intractable epilepsy were implanted with the TP electrode array for purposes of seizure localization. Select patients underwent cognitive mapping by electrocorticographic (ECoG) recording from the TP during a naming task. Use of the array resulted in excellent TP electrode coverage in all patients. High quality ECoG data were consistently obtained for purposes of delineating seizure activity and functional mapping. During a naming task, significant increases in ECoG power were observed within localized subregions of the TP. One patient developed a transient neurological deficit thought to be related to the mass effect of multiple intracranial recording arrays, including the TP array. This deficit resolved following removal of all electrodes. The TP electrode array overcomes limitations of existing devices and enables clinicians and researchers to obtain optimal multi-site recordings from this important brain region.

Sequence learning modulates neural responses and oscillatory coupling in human and monkey auditory cortex

Learning complex ordering relationships between sensory events in a sequence is fundamental for animal perception and human communication. While it is known that rhythmic sensory events can entrain brain oscillations at different frequencies, how learning and prior experience with sequencing relationships affect neocortical oscillations and neuronal responses is poorly understood. We used an implicit sequence learning paradigm (an “artificial grammar”) in which humans and monkeys were exposed to sequences of nonsense words with regularities in the ordering relationships between the words. We then recorded neural responses directly from the auditory cortex in both species in response to novel legal sequences or ones violating specific ordering relationships. Neural oscillations in both monkeys and humans in response to the nonsense word sequences show strikingly similar hierarchically nested low-frequency phase and high-gamma amplitude coupling, establishing this form of oscillatory coupling—previously associated with speech processing in the human auditory cortex—as an evolutionarily conserved biological process. Moreover, learned ordering relationships modulate the observed form of neural oscillatory coupling in both species, with temporally distinct neural oscillatory effects that appear to coordinate neuronal responses in the monkeys. This study identifies the conserved auditory cortical neural signatures involved in monitoring learned sequencing operations, evident as modulations of transient coupling and neuronal responses to temporally structured sensory input.

Sound identification in human auditory cortex: Differential contribution of local field potentials and high gamma power as revealed by direct intracranial recordings

High gamma power has become the principal means of assessing auditory cortical activation in human intracranial studies, albeit at the expense of low frequency local field potentials (LFPs). It is unclear whether limiting analyses to high gamma impedes ability of clarifying auditory cortical organization. We compared the two measures obtained from posterolateral superior temporal gyrus (PLST) and evaluated their relative utility in sound categorization. Subjects were neurosurgical patients undergoing invasive monitoring for medically refractory epilepsy. Stimuli (consonant-vowel syllables varying in voicing and place of articulation and control tones) elicited robust evoked potentials and high gamma activity on PLST. LFPs had greater across-subject variability, yet yielded higher classification accuracy, relative to high gamma power. Classification was enhanced by including temporal detail of LFPs and combining LFP and high gamma. We conclude that future studies should consider utilizing both LFP and high gamma when investigating the functional organization of human auditory cortex.

The use of auditory and visual context in speech perception by listeners with normal hearing and listeners with cochlear implants

There is a wide range of acoustic and visual variability across different talkers and different speaking contexts. Listeners with normal hearing (NH) accommodate that variability in ways that facilitate efficient perception, but it is not known whether listeners with cochlear implants (CIs) can do the same. In this study, listeners with NH and listeners with CIs were tested for accommodation to auditory and visual phonetic contexts created by gender-driven speech differences as well as vowel coarticulation and lip rounding in both consonants and vowels. Accommodation was measured as the shifting of perceptual boundaries between /s/ and /∫/ sounds in various contexts, as modeled by mixed-effects logistic regression. Owing to the spectral contrasts thought to underlie these context effects, CI listeners were predicted to perform poorly, but showed considerable success. Listeners with CIs not only showed sensitivity to auditory cues to gender, they were also able to use visual cues to gender (i.e., faces) as a supplement or proxy for information in the acoustic domain, in a pattern that was not observed for listeners with NH. Spectrally-degraded stimuli heard by listeners with NH generally did not elicit strong context effects, underscoring the limitations of noise vocoders and/or the importance of experience with electric hearing. Visual cues for consonant lip rounding and vowel lip rounding were perceived in a manner consistent with coarticulation and were generally used more heavily by listeners with CIs. Results suggest that listeners with CIs are able to accommodate various sources of acoustic variability either by attending to appropriate acoustic cues or by inferring them via the visual signal.

Electrocorticographic Activation within Human Auditory Cortex during Dialog-Based Language and Cognitive Testing

Current models of cortical speech and language processing include multiple regions within the temporal lobe of both hemispheres. Human communication, by necessity, involves complex interactions between regions subserving speech and language processing with those involved in more general cognitive functions. To assess these interactions, we utilized an ecologically salient conversation-based approach. This approach mandates that we first clarify activity patterns at the earliest stages of cortical speech processing. Therefore, we examined high gamma (70–150 Hz) responses within the electrocorticogram (ECoG) recorded simultaneously from Heschl’s gyrus (HG) and lateral surface of the superior temporal gyrus (STG). Subjects were neurosurgical patients undergoing evaluation for treatment of medically intractable epilepsy. They performed an expanded version of the Mini-mental state examination (MMSE), which included additional spelling, naming, and memory-based tasks. ECoG was recorded from HG and the STG using multicontact depth and subdural electrode arrays, respectively. Differences in high gamma activity during listening to the interviewer and the subject’s self-generated verbal responses were quantified for each recording site and across sites within HG and STG. The expanded MMSE produced widespread activation in auditory cortex of both hemispheres. No significant difference was found between activity during listening to the interviewer’s questions and the subject’s answers in posteromedial HG (auditory core cortex). A different pattern was observed throughout anterolateral HG and posterior and middle portions of lateral STG (non-core auditory cortical areas), where activity was significantly greater during listening compared to speaking. No systematic task-specific differences in the degree of suppression during speaking relative to listening were found in posterior and middle STG. Individual sites could, however, exhibit task-related variability in the degree of suppression during speaking compared to listening. The current study demonstrates that ECoG recordings can be acquired in time-efficient dialog-based paradigms, permitting examination of language and cognition in an ecologically salient manner. The results obtained from auditory cortex serve as a foundation for future studies addressing patterns of activity beyond auditory cortex that subserve human communication.

Modified locus equations categorize stop place in a perceptually realistic time frame

Slope and y-intercepts of locus equations have previously been shown to successfully classify place of articulation for English voiced stop consonants when derived from measurements at vowel onset and vowel midpoint. However, listeners are capable of identifying English voiced stops when less than 30 ms of vowel is presented. The present results show that modified locus equation measurements made within the first several pitch periods of a vowel following an English voiced stop were also successful at classifying place of articulation, consistent with the amount of vocalic information necessary for perceptual identification of English voiced stops /b d g/.

The Elicitation of Audiovisual Steady-State Responses: Multi-Sensory Signal Congruity and Phase Effects

Most ecologically natural sensory inputs are not limited to a single modality. While it is possible to use real ecological materials as experimental stimuli to investigate the neural basis of multi-sensory experience, parametric control of such tokens is limited. By using artificial bimodal stimuli composed of approximations to ecological signals, we aim to observe the interactions between putatively relevant stimulus attributes. Here we use MEG as an electrophysiological tool and employ as a measure the steady-state response (SSR), an experimental paradigm typically applied to unimodal signals. In this experiment we quantify the responses to a bimodal audio-visual signal with different degrees of temporal (phase) congruity, focusing on stimulus properties critical to audiovisual speech. An amplitude modulated auditory signal (‘pseudo-speech’) is paired with a radius-modulated ellipse (‘pseudo-mouth’), with the envelope of low-frequency modulations occurring in phase or at offset phase values across modalities. We observe (i) that it is possible to elicit an SSR to bimodal signals; (ii) that bimodal signals exhibit greater response power than unimodal signals; and (iii) that the SSR power at specific harmonics and sensors differentially reflects the congruity between signal components. Importantly, we argue that effects found at the modulation frequency and second harmonic reflect differential aspects of neural coding of multisensory signals. The experimental paradigm facilitates a quantitative characterization of properties of multi-sensory speech and other bimodal computations.

Electrocorticographic delineation of human auditory cortical fields based on effects of propofol anesthesia

ABSTRACT The functional organization of human auditory cortex remains incompletely characterized. While the posteromedial two thirds of Heschls gyrus (HG) is generally considered to be part of core auditory cortex, additional subdivisions of HG remain speculative. To further delineate the hierarchical organization of human auditory cortex, we investigated regional heterogeneity in the modulation of auditory cortical responses under varying depths of anesthesia induced by propofol. Non‐invasive studies have shown that propofol differentially affects auditory cortical activity, with a greater impact on non‐core areas. Subjects were neurosurgical patients undergoing removal of intracranial electrodes placed to identify epileptic foci. Stimuli were 50 Hz click trains, presented continuously during an awake baseline period, and subsequently, while propofol infusion was incrementally titrated to induce general anesthesia. Electrocorticographic recordings were made with depth electrodes implanted in HG and subdural grid electrodes implanted over superior temporal gyrus (STG). Depth of anesthesia was monitored using spectral entropy. Averaged evoked potentials (AEPs), frequency‐following responses (FFRs) and high gamma (70–150 Hz) event‐related band power were used to characterize auditory cortical activity. Based on the changes in AEPs and FFRs during the induction of anesthesia, posteromedial HG could be divided into two subdivisions. In the most posteromedial aspect of the gyrus, the earliest AEP deflections were preserved and FFRs increased during induction. In contrast, the remainder of the posteromedial HG exhibited attenuation of both the AEP and the FFR. The anterolateral HG exhibited weaker activation characterized by broad, low‐voltage AEPs and the absence of FFRs. Lateral STG exhibited limited activation by click trains, and FFRs there diminished during induction. Sustained high gamma activity was attenuated in the most posteromedial portion of HG, and was absent in all other regions. These differential patterns of auditory cortical activity during the induction of anesthesia may serve as useful physiological markers for field delineation. In this study, the posteromedial HG could be parcellated into at least two subdivisions. Preservation of the earliest AEP deflections and FFRs in the posteromedial HG likely reflects the persistence of feedforward synaptic activity generated by inputs from subcortical auditory pathways, including the medial geniculate nucleus. HighlightsAuditory cortex on Heschl’s gyrus (HG) was delineated using propofol anesthesia.Core auditory cortex on HG could be parcellated into two functional divisions.Short‐latency responses in posteromedial tip of HG persisted throughout induction.Frequency‐following responses in posteromedial tip of HG increased during induction.High gamma activity was attenuated during propofol induction.