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Kimmo Alho

Publications and source records attributed to Kimmo Alho.

At least 19 recordsLinked to original sources

Human brain activity associated with audiovisual perception and attention.

Coherent perception of objects in our environment often requires perceptual integration of auditory and visual information. Recent behavioral data suggest that audiovisual integration depends on attention. The current study investigated the neural basis of audiovisual integration using 3-Tesla functional magnetic resonance imaging (fMRI) in 12 healthy volunteers during attention to auditory or visual features, or audiovisual feature combinations of abstract stimuli (simultaneous harmonic sounds and colored circles). Audiovisual attention was found to modulate activity in the same frontal, temporal, parietal and occipital cortical regions as auditory and visual attention. In addition, attention to audiovisual feature combinations produced stronger activity in the superior temporal cortices than attention to only auditory or visual features. These modality-specific areas might be involved in attention-dependent perceptual binding of synchronous auditory and visual events into coherent audiovisual objects. Furthermore, the modality-specific temporal auditory and occipital visual cortical areas showed attention-related modulations during both auditory and visual attention tasks. This result supports the proposal that attention to stimuli in one modality can spread to encompass synchronously presented stimuli in another modality.

Adolescent↗

Selective attention to sound location or pitch studied with fMRI.

We used 3-T functional magnetic resonance imaging to compare the brain mechanisms underlying selective attention to sound location and pitch. In different tasks, the subjects (N = 10) attended to a designated sound location or pitch or to pictures presented on the screen. In the Attend Location conditions, the sound location varied randomly (left or right), while the pitch was kept constant (high or low). In the Attend Pitch conditions, sounds of randomly varying pitch (high or low) were presented at a constant location (left or right). Both attention to location and attention to pitch produced enhanced activity (in comparison with activation caused by the same sounds when attention was focused on the pictures) in widespread areas of the superior temporal cortex. Attention to either sound feature also activated prefrontal and inferior parietal cortical regions. These activations were stronger during attention to location than during attention to pitch. Attention to location but not to pitch produced a significant increase of activation in the premotor/supplementary motor cortices of both hemispheres and in the right prefrontal cortex, while no area showed activity specifically related to attention to pitch. The present results suggest some differences in the attentional selection of sounds on the basis of their location and pitch consistent with the suggested auditory "what" and "where" processing streams.

Acoustic Stimulation↗

Two separate mechanisms underlie auditory change detection and involuntary control of attention.

We used behavioral and event-related potential (ERP) measures to study the neural mechanisms of involuntary attention switching to changes in unattended sounds. Our subjects discriminated two equiprobable sounds differing in frequency (fundamental frequency 186 or 196 Hz) while task-irrelevant intensity decrements or increments (-3, -6, -9, +3, +6, or +9 dB, standard intensity 60 dB HL) infrequently occurred in the same sounds. In line with the results of previous studies, discrimination performance deteriorated with increasing magnitude of the task-irrelevant intensity change. However, these distraction effects were dissimilar for intensity increments and decrements: while there were no differences in reaction time (RT) between intensity decrements and increments, hit rates (HR) were lower for large intensity increments than for large decrements. ERPs to task-irrelevant intensity increments and decrements were also distinctly different: the response to intensity increments consisted of an N1 enhancement, mismatch negativity (MMN), and P3a, while the response to intensity decrements consisted only of MMN. These results are consistent with the assumption that two separate mechanisms (indexed by N1 and MMN) underlie auditory change detection. However, the finding that distinct distraction effects were obtained for both intensity decrements and increments but that the P3a is elicited only by the intensity increments seems to suggest that P3a may not be regarded as a general index of attentional shift but rather it is only generated in conditions in which an enhanced N1 is elicited, too.

Adult↗

Selective attention to human voice enhances brain activity bilaterally in the superior temporal sulcus.

Regional cerebral blood flow was measured with positron emission tomography (PET) in 10 healthy male volunteers. They heard two binaurally delivered concurrent stories, one spoken by a male voice and the other by a female voice. A third story was presented at the same time as a text running on a screen. The subjects were instructed to attend silently to one of the stories at a time. In an additional resting condition, no stories were delivered. PET data showed that in comparison with the reading condition, the brain activity in the speech-listening conditions was enhanced bilaterally in the anterior superior temporal sulcus including cortical areas that have been reported to be specifically sensitive to human voice. Previous studies on attention to non-linguistic sounds and visual objects, in turn, showed prefrontal activations that are presumably related to attentional control functions. However, comparisons of the present speech-listening and reading conditions with each other or with the resting condition indicated no prefrontal activity, except for an activation in the inferior frontal cortex that was presumably associated with semantic and syntactic processing of the attended story. Thus, speech listening, as well as reading, even in a distracting environment appears to depend less on the prefrontal control functions than do other types of attention-demanding tasks, probably because selective attention to speech and written text are over-learned actions rehearsed daily.

Adult↗

Loudness summation and the mismatch negativity event-related brain potential in humans.

Abstract Infrequently omitting a sound from a repetitive sequence elicits the mismatch negativity (MMN) ERP response when the stimulus onset asynchrony (SOA) is less than 200 ms. We contrasted two alternative explanations of omission MMN. (1) Each sound starts a separate temporal integration process. Omissions violate the constancy of the temporal structure within the integration window. (2) Sounds preceding an omission are perceived to be louder than those followed by a sound within the integration period, because omissions allow the full stimulus aftereffect to be included in perceived loudness. We varied the SOA between 117 and 217 ms. For this case, the temporal structure explanation predicts that no MMN will be elicited, whereas the loudness summation explanation predicts that MMN will be elicited. MMN was elicited by tone omissions with random SOA, suggesting that loudness summation plays an important role in the elicitation of omission MMN.

Acoustic Stimulation↗

Does sleep quality affect involuntary attention switching system?

We studied the relationship between sleep quality and quantity and subsequently recorded automatically evoked event-related potential (ERP) responses. In previous studies decrement of attentional processing has been associated with changes in sleep. Sleep is shown to associate also with ERPs elicited by unattended sound stream, however, there is no consensus on these effects. A recent study suggested that the early anterior P3a to novel stimuli in attended stream is attenuated and the late parietal P3a is strengthened by total sleep deprivation. We carried out 72-h consecutive actigraphy measurements in a naturalistic setting to collect information about variation in sleep duration, sleep onset latency, sleep efficiency, and percentage of sleep. MMN and P3a deflections to infrequent changes in sound duration and pitch in unattended sound stream were obtained in a separate recording session from the same subjects when they were awake. No significant correlations were found between sleep and MMN parameters, indicating that MMN is resistant to normal variation in sleep. However, P3a to both pitch and duration changes correlated positively with sleep onset latency, and P3a to duration changes correlated negatively with sleep efficiency and percentage of sleep. The correlation was higher in the posterior scalp areas. Our results suggest that the involuntary attention switching system, reflected by the P3a is sensitized as a function of decreased sleep quality.

Acoustic Stimulation↗

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.

Acoustic Stimulation↗

Modulation of auditory cortex activation by sound presentation rate and attention.

We studied the effects of sound presentation rate and attention on auditory supratemporal cortex (STC) activation in 12 healthy adults using functional magnetic resonance imaging (fMRI) at 3 T. The sounds (200 ms in duration) were presented at steady rates of 0.5, 1, 1.5, 2.5, or 4 Hz while subjects either had to focus their attention to the sounds or ignore the sounds and attend to visual stimuli presented with a mean rate of 1 Hz. Consistent with previous observations, we found that both increase in stimulation rate and attention to sounds enhanced activity in STC bilaterally. Further, we observed larger attention effects with higher stimulation rates. This interaction of attention and presentation rate has not been reported previously. In conclusion, our results show both rate-dependent and attention-related modulations of STC indicating that both factors should be controlled, or at least addressed, in fMRI studies of auditory processing.

Acoustic Stimulation↗

Attentional modulation of human auditory cortex.

Attention powerfully influences auditory perception, but little is understood about the mechanisms whereby attention sharpens responses to unattended sounds. We used high-resolution surface mapping techniques (using functional magnetic resonance imaging, fMRI) to examine activity in human auditory cortex during an intermodal selective attention task. Stimulus-dependent activations (SDAs), evoked by unattended sounds during demanding visual tasks, were maximal over mesial auditory cortex. They were tuned to sound frequency and location, and showed rapid adaptation to repeated sounds. Attention-related modulations (ARMs) were isolated as response enhancements that occurred when subjects performed pitch-discrimination tasks. In contrast to SDAs, ARMs were localized to lateral auditory cortex, showed broad frequency and location tuning, and increased in amplitude with sound repetition. The results suggest a functional dichotomy of auditory cortical fields: stimulus-determined mesial fields that faithfully transmit acoustic information, and attentionally labile lateral fields that analyze acoustic features of behaviorally relevant sounds.

Acoustic Stimulation↗

Linguistic processing in visual and modality-nonspecific brain areas: PET recordings during selective attention.

Positron emission tomography (PET) was used to investigate the neural basis of selective processing of linguistic material during concurrent presentation of multiple stimulus streams ("cocktail-party effect"). Fifteen healthy right-handed adult males were to attend to one of three simultaneously presented messages: one presented visually, one to the left ear, and one to the right ear. During the control condition, subjects attended to visually presented consonant letter strings and ignored auditory messages. This paper reports the modality-nonspecific language processing and visual word-form processing, whereas the auditory attention effects have been reported elsewhere [Cogn. Brain Res. 17 (2003) 201]. The left-hemisphere areas activated by both the selective processing of text and speech were as follows: the inferior prefrontal (Brodmann's area, BA 45, 47), anterior temporal (BA 38), posterior insular (BA 13), inferior (BA 20) and middle temporal (BA 21), occipital (BA 18/30) cortices, the caudate nucleus, and the amygdala. In addition, bilateral activations were observed in the medial occipito-temporal cortex and the cerebellum. Decreases of activation during both text and speech processing were found in the parietal (BA 7, 40), frontal (BA 6, 8, 44) and occipito-temporal (BA 37) regions of the right hemisphere. Furthermore, the present data suggest that the left occipito-temporal cortex (BA 18, 20, 37, 21) can be subdivided into three functionally distinct regions in the posterior-anterior direction on the basis of their activation during attentive processing of sublexical orthography, visual word form, and supramodal higher-level aspects of language.

Adolescent↗

Activation in the anterior left auditory cortex associated with phonological analysis of speech input: localization of the phonological mismatch negativity response with MEG.

The spatio-temporal dynamics of cortical activation underlying auditory word recognition, particularly its phonological stage, was studied with whole-head magnetoencephalography (MEG). Subjects performed a visuo-auditory priming task known to evoke the phonological mismatch negativity (PMN) response that is elicited by violations of phonological expectancies. Words and non-words were presented in separate conditions. In each of the 318 trials, the subjects first saw a word/non-word (e.g., 'cat') that was soon followed by a prime letter (e.g., 'h'). Their task was to replace mentally the sound of the first letter of the word/non-word with the prime letter, thus resulting in a new word/non-word (e.g., 'hat'). Finally, an auditory word/non-word either matching or mismatching with the anticipated item was presented. In most subjects, a PMNm followed by a later, N400m-like negativity was obtained in the left hemisphere to the mismatching auditory stimuli. A similar response pattern was obtained in the right hemisphere only in a few subjects. Source localization of the N1m, an index of acoustic analysis, and the PMNm and N400m-like responses was performed using L1 minimum-norm estimation. In the left hemisphere, the PMNm source for the words was significantly more anterior than the source of the N400m-like response; for the non-words, the PMNm source was significantly more anterior than the sources of the N1m and the N400m-like response. These results suggest that the left-hemisphere neuronal networks involved in sub-lexical phonological analysis are at least partly different from those responsible for the earlier (acoustic) and later (whole item) processing of speech input.

Acoustic Stimulation↗

Local landmark-based mapping of human auditory cortex.

Mammalian sensory cortex is functionally partitioned into cortical fields that are specialized for different processing operations. In theory, averaging functional and anatomical images across subjects can reveal both the average anatomy and the mean functional organization of sensory regions. However, this averaging process must overcome at least two obstacles: (1) the relative locations and sizes of cortical sensory areas vary in different subjects so that across-subject averaging introduces spatial smearing; (2) the relative locations and sizes of cortical areas vary between hemispheres, making it difficult to compare activations between hemispheres or to combine activations across hemispheres. These difficulties are particularly acute for small cortical regions such as auditory cortex. In whole-brain averaging procedures, considerable intersubject variance in the location and orientation of auditory cortex is introduced by variance of the size and shape of structures outside auditory cortex. Here, we compared these global methods with local landmark-based methods (LLMs) that use warping based on local anatomical landmarks. In comparison to maps made with global methods, LLMs produced anatomical maps of auditory cortex with clearer gyral and sulcal structure, and produce functional maps with improved resolution. These results suggest that LLMs have significant advantages over global mapping procedures in studying the details of auditory cortex organization.

Adult↗

Effects of auditory distraction on electrophysiological brain activity and performance in children aged 8-13 years.

Distractibility was investigated in three age groups of children (8-9, 10-11, and 12-13 years) with event-related brain potentials (ERPs) and performance measures in a forced-choice visual task. Distraction was reflected by increased reaction times (RTs) and decreased performance accuracy in the visual discrimination task following presentation of unexpected novel sounds. The amplitude of the late portion of the P3a elicited by novel sounds was largest for the youngest group and showed a centrally dominant scalp distribution and smallest for the oldest group with a frontal scalp distribution. A frontally dominant late negativity (LN) that was largest in the youngest group followed the P3a. Correlation between the RT increase caused by the distracting novel sounds and the amplitude of the LN elicited by these sounds suggested that the LN is associated with the degree of attention engaged by the distracting sounds.

Adolescent↗

Long-term exposure to noise impairs cortical sound processing and attention control.

Long-term exposure to noise impairs human health, causing pathological changes in the inner ear as well as other anatomical and physiological deficits. Numerous individuals are daily exposed to excessive noise. However, there is a lack of systematic research on the effects of noise on cortical function. Here we report data showing that long-term exposure to noise has a persistent effect on central auditory processing and leads to concurrent behavioral deficits. We found that speech-sound discrimination was impaired in noise-exposed individuals, as indicated by behavioral responses and the mismatch negativity brain response. Furthermore, irrelevant sounds increased the distractibility of the noise-exposed subjects, which was shown by increased interference in task performance and aberrant brain responses. These results demonstrate that long-term exposure to noise has long-lasting detrimental effects on central auditory processing and attention control.

Adult↗

Electric brain responses indicate preattentive processing of abstract acoustic regularities in children.

This study investigated the preattentive processing of abstract acoustic regularities in children aged 8-14 years. Event-related brain potentials (ERPs) were elicited by frequent (standard) pairs ascending in pitch (the second tone having a higher frequency than the first tone) and by infrequent (deviant) pairs descending in pitch. In the easy condition, the second tone of the pair was always one step higher (standard) or lower (deviant) than the first tone, while in the hard condition, the second tone was randomly 1-10 steps higher or lower than the first tone. In the easy condition we found the mismatch negativity (MMN) and a subsequent positive P3a-like deflection. In the hard condition, the amplitude of MMN was lower over frontal sites than in the easy condition, while the temporal component of MMN was not impaired by complexity of abstract regularities. These results suggest that the complexity of the auditory stimulation affects preattentive auditory change detection in children.

Acoustic Stimulation↗

Processing abstract auditory features in the human auditory cortex.

Using electric and magnetic brain responses we tested whether violations of an abstract auditory regularity are processed in auditory cortex and whether abstract auditory regularities are retained for at least 10 s. The mismatch negativity (MMN) event-related potential and its magnetic counterpart (MMNm) were recorded to infrequent tone pairs of descending pitch (the second tone having a lower frequency than the first one) embedded in a sequence of tone pairs of ascending pitch, whereas the absolute frequency of both ascending and descending tone pairs varied on seven levels. Results showed that the dominant generators of the electromagnetic activity elicited by violations of the pitch-ascension rule lie within auditory cortex. We also found that the memory representation of pitch-ascension is retained for at least 10 s. When short trains of ascending-pitched tone pairs were followed by a silent period of 8-12 s, descending-pitched probe tone pairs elicited the MMN component when a single reminding pair with ascending pitch was presented before the probe. The reactivating effect of the reminder was similar to what has been previously shown for concrete auditory regularities, such as the constancy of tone pitch. The present results support the view that auditory cortical functions can process sensory and categorical information in a similar manner.

Acoustic Stimulation↗

Spatiotemporal dynamics of the auditory novelty-P3 event-related brain potential.

The spatiotemporal dynamics of the cerebral network involved in novelty processing was studied by means of scalp current density (SCD) analysis of the novelty P3 (nP3) event-related brain potential (ERP). ERPs were recorded from 30 scalp electrodes at the occurrence of novel unpredictable environmental sounds during the performance of a visual discrimination task. Increased SCD was observed at left frontotemporal (FT3), bilateral temporoparietal (TP3 and TP4) and prefrontal locations (F8-F4 and F7-F3), suggesting novelty-P3 generators located in the left auditory cortex, and bilaterally in temporoparietal and prefrontal association regions. Additional increased SCD was found at a central location (Cz) and at superior parietal locations (P3-Pz-P4). The SCD of the nP3 was therefore generated at three successive, partially overlapping, stages of neuroelectric activation. At the central location, SCD started to be significant before the onset of the nP3 waveform, contributing solely to its early phase. At temporoparietal and left frontotemporal locations, nP3 electrophysiological activity was characterized by sustained current density, starting at about 210 ms and continuing during the full latency range of the response, including its early and late phases. At its late phase, the nP3 was characterized by sharp phasic current density at prefrontal and superior parietal locations, starting at about 290 ms and vanishing at around 385 ms. Taken together, these results provide the first evidence of the cerebral spatio-temporal dynamics underlying novelty processing.

Acoustic Stimulation↗

Hemispheric lateralization of cerebral blood-flow changes during selective listening to dichotically presented continuous speech.

Regional cerebral blood flow (rCBF) was measured with positron emission tomography (PET) while subjects were selectively listening to continuous speech delivered to one ear and ignoring concurrent speech delivered to the opposite ear, as well as concurrent text or letter strings running on a screen. rCBF patterns associated with selective listening either to the left-ear or right-ear speech message were compared with each other and with rCBF patterns in two visual-attention conditions in which the subjects ignored both speech messages and either read the text or discriminated the meaningless letter strings moving on the screen. Attention to either speech message was associated with enhanced activity in the superior temporal cortex of the language-dominant left hemisphere, as well as in the superior and middle temporal cortex of the right hemisphere suggesting enhanced processing of prosodic features in the attended speech. Moreover, enhanced activity during attention to either speech message was observed in the right parietal areas known to have an important role in directing spatial attention. Evidence was also found for attentional tuning of the left and right auditory cortices to select information from the contralateral auditory hemispace.

Acoustic Stimulation↗