Teemu Rinne

Separate Time Behaviors of the Temporal and Frontal Mismatch Negativity Sources

It has been proposed that mismatch negativity (MMN) is generated by temporal and frontal lobe sources, the former being associated with change detection and the latter with involuntary switching of attention to sound change. If this switching of attention is triggered by the temporal cortex change-detection mechanism, one would expect that the frontal component of MMN is activated later than the temporal one. This was studied by using 64-channel electroencephalography (EEG) and 122-channel magnetoencephalography (MEG) with realistically shaped head models to determine the source current distribution in different lobes as a function of time. Minimum-norm estimation (MNE) was performed, constraining the solution to the reconstructed cortical sheet. The results support the hypothesis that the frontal MMN generator is activated later than the auditory cortex generator.

Differential Contribution of Frontal and Temporal Cortices to Auditory Change Detection: fMRI and ERP Results

The present study addresses the functional role of the temporal and frontal lobes in auditory change detection. Prior event-related potential (ERP) research suggested that the mismatch negativity (MMN) reflects the involvement of a temporofrontal network subserving auditory change detection processes and the initiation of an involuntary attention switch. In the present study participants were presented with repetitive spectrally rich sounds. Infrequent changes of either small (10% change), medium (30% change), or large (100% change) magnitude were embedded in the stimulus train. ERPs and fMRI measures were obtained in the same subjects in subsequent sessions. Significant hemodynamic activation in the superior temporal gyri (STG) bilaterally and the opercular part of the right inferior frontal gyrus was observed for large and medium deviants only. ERPs showed that small deviants elicited MMN when presented in silence but not when presented with recorded MR background noise, indicating that small deviants were hardly detected under fMRI conditions. The MR signal change in temporal lobe regions was larger for large than for medium deviants. For the right fronto-opercular cortex the opposite pattern was observed. The strength of the temporal activation correlated with the amplitude of the change-related ERP at around 110 ms from stimulus onset while the frontal activation correlated with the change-related ERP at around 150 ms. These results suggest that the right fronto-opercular cortex is part of the neural network generating the MMN. Three alternative explanations of these findings are discussed.

Maturation of cortical sound processing as indexed by event-related potentials

OBJECTIVES Childrens auditory event-related potentials (ERPs) are dominated by the P1 and N2 peaks, while the N1 wave emerges between 3 and 4 years of age. The neural substrates and the behavioral correlates of the protracted N1 maturation, as well as of the 10-year long predominance of the N2 are unclear. The present study utilized high-resolution electroencephalography to study the maturation of auditory ERPs from age 4 to adulthood and to compare the sources of the N1 and the N2 peaks in 9-year-old children and adults. METHODS Three partial harmonic tones were delivered with short (700 ms) and long (mean of 5s) stimulus onset asynchrony (SOA), with only 700 ms SOA used with 4-year-olds. RESULTS With a short SOA, 4- and 9-year-old children displayed P1 and N2 peaks, whereas adults showed P1, N1, P2, and N2 waves. With a long SOA, 9-year-olds also displayed an N1 peak, which was frontal in scalp distribution to that in adults who showed P1, N1, and P2 peaks. After filtering out the slow N2 activity, the N1 wave was also revealed in the short-SOA data in 9-year-old but not in 4-year-old children. In adults and in 9-year-olds, the neural sources of the N2 and N1 mapped onto the superior aspects of the temporal lobes, the sources of the N2 being anterior to those of the N1. CONCLUSIONS The results indicated that childrens N1 is composed of differently weighted components as that in adults, and that in both children and adults the N1 and N2 are generated by anatomically distinct generators. A protracted ontogeny of the N1 could be linked with that of auditory sensitivity and orienting, whereas the P1 and N2 peaks are suggested to reflect auditory sensory processes.

Analysis of speech sounds is left-hemisphere predominant at 100-150 ms after sound onset

Hemispheric specialization of human speech processing has been found in brain imaging studies using fMRI and PET. Due to the restricted time resolution, these methods cannot, however, determine the stage of auditory processing at which this specialization first emerges. We used a dense electrode array covering the whole scalp to record the mismatch negativity (MMN), an event-related brain potential (ERP) automatically elicited by occasional changes in sounds, which ranged from non-phonetic (tones) to phonetic (vowels). MMN can be used to probe auditory central processing on a millisecond scale with no attention-dependent task requirements. Our results indicate that speech processing occurs predominantly in the left hemisphere at the early, pre-attentive level of auditory analysis.

Superior temporal and inferior frontal cortices are activated by infrequent sound duration decrements: an fMRI study

Functional magnetic resonance imaging (fMRI) was used to examine the processing of infrequent changes occurring in an unattended sound sequence. In event-related brain potentials (ERPs), such sound changes typically elicit several responses, including an enhanced N1, the mismatch negativity (MMN), and the P3a. In the present study, subjects were presented with a repeating sound of 75 ms in duration, which was occasionally replaced, in separate blocks, by a 15-ms, 25-ms, or 35-ms sound (large, medium, and small change, respectively). In the baseline block, only the frequent 75-ms sound was presented. During the scanning, the subjects were instructed to ignore the sounds while watching a silent wildlife documentary. We assumed that in this condition, the MMN mechanism would contribute more to the observed activation than the other change-related processes. We expected sound changes to elicit fMRI activation bilaterally in the supratemporal cortices, where the electric MMN is mainly generated, and that the magnitude of this activation would increase with the magnitude of sound duration change. Unexpectedly, however, we found that only blocks with medium duration changes (25 ms) showed significant activation in the supratemporal cortex. In addition, as reported in some previous EEG and fMRI studies, contrasts between different levels of sound duration change revealed additional activation in the inferior frontal cortex bilaterally. This activation tended to be greater for the small and medium changes than for the large ones.

Variability and replicability of the mismatch negativity

The interindividual variation and test-retest stability of the mismatch negativity (MMN) and N1 components of the event-related potential (ERP) were investigated by presenting standard (85%) and deviant tones (15%) to 10 young subjects in 2 sessions separated by 1 month. Deviant tones in different blocks were either frequency or duration changes with interstimulus intervals (ISIs) of 0.5 and 1.5 sec. The results showed a fairly good test-retest stability of the MMN amplitude for both types of changes with each ISI at the group level. The amplitude of the duration MMN showed significant individual test-retest stability. The N1 amplitude showed high stability at both the group and individual levels. Both the MMN and N1 showed considerable interindividual variation. The results suggest that MMN and N1 can be used in follow-up studies not only at the group level but possibly at the individual level also.

Functional Maps of Human Auditory Cortex: Effects of Acoustic Features and Attention

Background While human auditory cortex is known to contain tonotopically organized auditory cortical fields (ACFs), little is known about how processing in these fields is modulated by other acoustic features or by attention. Methodology/Principal Findings We used functional magnetic resonance imaging (fMRI) and population-based cortical surface analysis to characterize the tonotopic organization of human auditory cortex and analyze the influence of tone intensity, ear of delivery, scanner background noise, and intermodal selective attention on auditory cortex activations. Medial auditory cortex surrounding Heschls gyrus showed large sensory (unattended) activations with two mirror-symmetric tonotopic fields similar to those observed in non-human primates. Sensory responses in medial regions had symmetrical distributions with respect to the left and right hemispheres, were enlarged for tones of increased intensity, and were enhanced when sparse image acquisition reduced scanner acoustic noise. Spatial distribution analysis suggested that changes in tone intensity shifted activation within isofrequency bands. Activations to monaural tones were enhanced over the hemisphere contralateral to stimulation, where they produced activations similar to those produced by binaural sounds. Lateral regions of auditory cortex showed small sensory responses that were larger in the right than left hemisphere, lacked tonotopic organization, and were uninfluenced by acoustic parameters. Sensory responses in both medial and lateral auditory cortex decreased in magnitude throughout stimulus blocks. Attention-related modulations (ARMs) were larger in lateral than medial regions of auditory cortex and appeared to arise primarily in belt and parabelt auditory fields. ARMs lacked tonotopic organization, were unaffected by acoustic parameters, and had distributions that were distinct from those of sensory responses. Unlike the gradual adaptation seen for sensory responses, ARMs increased in amplitude throughout stimulus blocks. Conclusions/Significance The results are consistent with the view that medial regions of human auditory cortex contain tonotopically organized core and belt fields that map the basic acoustic features of sounds while surrounding higher-order parabelt regions are tuned to more abstract stimulus attributes. Intermodal selective attention enhances processing in neuronal populations that are partially distinct from those activated by unattended stimuli.

Aging effects on auditory processing : An event-related potential study

Deviant tones randomly embedded in a sequence of standard tones elicit an event-related potential (ERP) called the mismatch negativity (MMN), which reflects automatic stimulus-change detection in the human auditory system. When the tones are attended, deviant tones elicit also an N2b component that partly overlaps the MMN. Sequences of standard and deviant (probability 0.15) tones were presented to 13 healthy younger and 13 older subjects. Deviant stimuli were, in separate blocks, either occasional shorter duration or higher frequency tones. The interstimulus interval (ISI) was, in separate blocks, either 0.5 s or 1.5 s, and in the frequency-change condition also 4.5 s. Aging affected neither frequency nor duration of MMN with the 0.5 s ISI. This finding indicates that automatic stimulus discrimination per se is not impaired with normal aging. However, with a 4.5-s ISI the MMN/N2b-complex attenuated significantly more in the older than younger subjects. This suggests that the stimulus trace decays faster or that involuntary attention switching is less sensitive with aging.

Brain networks of bottom-up triggered and top-down controlled shifting of auditory attention

During functional magnetic resonance imaging (fMRI), our participants selectively attended to tone streams at the left or right, and occasionally shifted their attention from one stream to another as guided by a centrally presented visual cue. Duration changes in the to-be-attended stream served as targets. Loudness deviating tones (LDTs) occurred infrequently in both streams to catch attention in a bottom-up manner, as indicated by their effects on reaction times to targets. LDTs activated the right temporo-parietal junction (TPJ), posterior parts of the left inferior/middle frontal gyrus (IFG/MFG), ventromedial parts of the superior parietal lobule (SPL), and left frontal eye field/premotor cortex (FEF/PMC). In addition, LDTs in the to-be-ignored sound stream were associated with enhanced activity in the ventromedial prefrontal cortex (VMPFC) possibly related to evaluation of the distracting event. Top-down controlled cue-guided attention shifts (CASs) activated bilateral areas in the SPL, intraparietal sulcus (IPS), FEF/PMC, TPJ, IFG/MFG, and cingulate/medial frontal gyrus, and crus I/II of the cerebellum. Thus, our results suggest that in audition top-down controlled and bottom-up triggered shifting of attention activate largely overlapping temporo-parietal, superior parietal and frontal areas. As the IPS, superior parts of the SPL, and crus I/II were activated specifically by top-down controlled attention shifts, and the VMPFC was specifically activated by bottom-up triggered attention shifts, our results also suggest some differences between auditory top-down controlled and bottom-up triggered shifting of attention.

RAPID COMMUNICATION Scalp-Recorded Optical Signals Make Sound Processing in the Auditory Cortex Visible

1982) have demonstratedtonotopic organization in the auditory cortex. Thus,these methods, especially fMRI, are able to map brainareas involved in auditory processing on a millimeterscale. However, fMRI is not able to track the fine-grained temporal dynamics of the central auditorysystem in encoding and analyzing sound informationbecause its time resolution is limited by the propertiesof the hemodynamic changes, lagging neural activity atleast by several hundred milliseconds. In contrast,MEG provides temporally detailed information on cen-tral auditory processing (Lu¨tkenho¨ner and Stein-stra¨ter, 1998) but has severe problems in separatingsimultaneously activated adjacent sources from eachother and cannot indicate the extent and pattern of theactivated area. The present paper reports the firstevent-related optical signals (EROS; Gratton and Fabi-ani, 1998) from the functioning human auditory cortex.Usingthisnovelnoninvasivemeasureofcorticalactiva-tion, we were able to locate, in both time and space,neuronal generators for two of the most importantauditory functions, sound detection and change detec-tion (Na¨a¨ta¨nen, 1992).The EROS method combines both high spatial andhigh temporal resolution in a single measure (Grattonand Fabiani, 1998). In an EROS recording, a source ofintensity-modulated near-infrared light and a detectorare placed on the scalp a few centimeters apart fromeach other. The low-intensity light emitted by thesource diffuses through the skin, bone, and brain, andsome photons exit the head, reaching the detector. Aspatially high-resolution signal is achieved by selectingthe photons on the basis of their time of flight; thosephotons that take similar amounts of time to migratethrough the medium are likely to follow relativelysimilar paths. EROS is a measure of phase-shifts in themodulationenvelopeofthelightasthephotonsmigratethrough the brain tissue, which is optically modified byneural activation. Some of the optical changes in thebrain tissue are related to various hemodynamic phe-nomena lasting relatively long times compared withneural electric activity (Cannestra