Biomedical subjects

Ville Mäkinen

Publications and source records attributed to Ville Mäkinen.

10 recordsLinked to original sources

PubMed2005

Disentangling the effects of phonation and articulation: hemispheric asymmetries in the auditory N1m response of the human brain.

BACKGROUND: The cortical activity underlying the perception of vowel identity has typically been addressed by manipulating the first and second formant frequency (F1 & F2) of the speech stimuli. These two values, originating from articulation, are already sufficient for the phonetic characterization of vowel category. In the present study, we investigated how the spectral cues caused by articulation are reflected in cortical speech processing when combined with phonation, the other major part of speech production manifested as the fundamental frequency (F0) and its harmonic integer multiples. To study the combined effects of articulation and phonation we presented vowels with either high (/a/) or low (/u/) formant frequencies which were driven by three different types of excitation: a natural periodic pulseform reflecting the vibration of the vocal folds, an aperiodic noise excitation, or a tonal waveform. The auditory N1m response was recorded with whole-head magnetoencephalography (MEG) from ten human subjects in order to resolve whether brain events reflecting articulation and phonation are specific to the left or right hemisphere of the human brain. RESULTS: The N1m responses for the six stimulus types displayed a considerable dynamic range of 115-135 ms, and were elicited faster (approximately 10 ms) by the high-formant /a/ than by the low-formant /u/, indicating an effect of articulation. While excitation type had no effect on the latency of the right-hemispheric N1m, the left-hemispheric N1m elicited by the tonally excited /a/ was some 10 ms earlier than that elicited by the periodic and the aperiodic excitation. The amplitude of the N1m in both hemispheres was systematically stronger to stimulation with natural periodic excitation. Also, stimulus type had a marked (up to 7 mm) effect on the source location of the N1m, with periodic excitation resulting in more anterior sources than aperiodic and tonal excitation. CONCLUSION: The auditory brain areas of the two hemispheres exhibit differential tuning to natural speech signals, observable already in the passive recording condition. The variations in the latency and strength of the auditory N1m response can be traced back to the spectral structure of the stimuli. More specifically, the combined effects of the harmonic comb structure originating from the natural voice excitation caused by the fluctuating vocal folds and the location of the formant frequencies originating from the vocal tract leads to asymmetric behaviour of the left and right hemisphere.

Acoustic Stimulation↗

PubMed2005

Left-hemispheric brain activity reflects formant transitions in speech sounds.

Connected speech is characterized by formant transitions whereby formant frequencies change over time. Here, using magneto-encephalography, we investigated the cortical activity in 10 participants in response to constant-formant vowels and diphthongs with formant transitions. All the stimuli elicited prominent auditory N100m responses, but the formant transitions resulted in latency modulations specific to the left hemisphere. Following the elicitation of the N100m, cortical activity shifted some 10 mm towards anterior brain areas. This late activity resembled the N400m, typically obtained with more complex utterances such as words and/or sentences. Thus, the present study demonstrates how magnetoencephalography can be used to investigate the spatiotemporal evolution in cortical activity related to the various stages of the processing of speech.

Acoustic Stimulation↗

PubMed2005

Spatial processing in human auditory cortex: the effects of 3D, ITD, and ILD stimulation techniques.

Here, the perception of auditory spatial information as indexed by behavioral measures is linked to brain dynamics as reflected by the N1m response recorded with whole-head magnetoencephalography (MEG). Broadband noise stimuli with realistic spatial cues corresponding to eight direction angles in the horizontal plane were constructed via custom-made, individualized binaural recordings (BAR) and generic head-related transfer functions (HRTF). For comparison purposes, stimuli with impoverished acoustical cues were created via interaural time and level differences (ITDs and ILDs) and their combinations. MEG recordings in ten subjects revealed that the amplitude and the latency of the N1m exhibits directional tuning to sound location, with the amplitude of the right-hemispheric N1m being particularly sensitive to the amount of spatial cues in the stimuli. The BAR, HRTF, and combined ITD + ILD stimuli resulted both in a larger dynamic range and in a more systematic distribution of the N1m amplitude across stimulus angle than did the ITD or ILD stimuli alone. Further, the right-hemispheric source loci of the N1m responses for the BAR and HRTF stimuli were anterior to those for the ITD and ILD stimuli. In behavioral tests, we measured the ability of the subjects to localize BAR and HRTF stimuli in terms of azimuthal error and front-back confusions. We found that behavioral performance correlated positively with the amplitude of the N1m. Thus, the activity taking place already in the auditory cortex predicts behavioral sound detection of spatial stimuli, and the amount of spatial cues embedded in the signal are reflected in the activity of this brain area.

Acoustic Stimulation↗

PubMed2005

Serotonergic modulation of mismatch negativity.

Neurochemical mechanisms mediating the interaction between emotional and cognitive processing are not yet fully understood. Here, we utilized acute tryptophan depletion (ATD) to reduce the brain synthesis of serotonin (5-HT), which is thought to have a central role in regulation of emotions and mood in humans. ATD effects on event-related potentials and magnetic fields were studied using a passive odd-ball paradigm in a randomized, double-blinded, controlled, cross-over design. Auditory responses were recorded simultaneously with high-resolution magnetoencephalography (MEG) and electroencephalography (EEG) in 14 healthy subjects, 5 h after ATD or a control condition. ATD significantly increased depressed mood and lowered plasma tryptophan concentration (total tryptophan decreased by 75%, free tryptophan decreased by 39%). As compared with the control condition, ATD increased the amplitudes of mismatch negativity (MMN) to duration and frequency changes and decreased the latencies of magnetic MMN to frequency changes in the hemisphere ipsilateral to the ear stimulated. Further, ATD modulated N1m latencies and decreased P2m source activity. ATD increased the interhemispheric latency difference of MMNm to frequency changes. No effects on P50 were observed. The present results suggest serotonergic modulation of preattentive auditory change detection, suggested to initiate involuntary attention shifting in the human brain.

Adult↗

PubMed2004

Auditory event-related responses are generated independently of ongoing brain activity.

For researchers and clinical practitioners alike, evoked and event-related responses measured with MEG and EEG provide the means for studying human brain function and dysfunction. However, the generation mechanism of event-related responses remains unclear, hindering our ability to formulate viable theories of neural information processing. Event-related responses are assumed to be generated either (1) separately of ongoing, oscillatory brain activity or (2) through stimulus-induced reorganization of ongoing activity. Here, we approached this issue through examining single-trial auditory MEG data in humans. We demonstrate that phase coherence over trials observed with commonly used signal decomposition methods (e.g., wavelets) can result from both a phase-coherent state of ongoing oscillations and from the presence of a phase-coherent event-related response which is additive to ongoing oscillations. To avoid this problem, we introduce a method based on amplitude variance to establish the relationship between ongoing oscillations and event-related responses. We found that auditory stimuli do not give rise to phase reorganization of ongoing activity. Further, increases in spectral power accompany the emergence of event-related responses, and the relationship between spectral power and the amplitude of these responses can be accounted for by a linear summation of the event-related response and ongoing oscillation with a stochastically distributed phase. Thus, on the basis of our observations, auditory event-related responses are unique descriptors of neural information processing in humans, generated by processes separate from and additive to ongoing brain activity.

Acoustic Stimulation↗

PubMed2004

Glides in speech fundamental frequency are reflected in the auditory N1m response.

The cortical dynamics underlying the perception of constant and gliding speech fundamental frequency (F0) was investigated in 10 subjects using magnetoencephalography (MEG). The stimuli comprised vowels having either constant, ascending or descending F0s and tones of corresponding frequencies, matched with the vowels in intensity or loudness. The amplitude of the N1m response was highly sensitive to F0 variation embedded in vowels and insensitive to corresponding variation in tones. The latency of the N1m elicited by the tones with respect to vowels was significantly delayed. Thus, the speech-specific behavior of the N1m arises out of cortical sensitivity to the acoustic structure of voiced speech, that is to the F0 and its harmonics, which underlie the perception of pitch and intonation in speech.

Acoustic Stimulation↗

PubMed2004

Transient brain responses predict the temporal dynamics of sound detection in humans.

The neural events leading up to the conscious experience of stimulus events have remained elusive. Here we describe stimulation conditions under which activation in human auditory cortex can be used to predict the temporal dynamics of behavioral sound detection. Subjects were presented with auditory stimuli whose energy smoothly increased from a silent to a clearly audible level over either 1, 1.5, or 2 s. Magnetoencephalographic (MEG) recordings were carried out in the passive and active recording conditions. In the active condition, the subjects were instructed to attend to the auditory stimuli and to press a response key when these became audible. In both conditions, the stimuli elicited a prominent transient response whose emergence is unexplainable by changes in stimulus intensity alone. This transient response was larger in amplitude over the right hemisphere and in the active condition. Importantly, behavioral sound detection followed this brain activation with a constant delay of 180 ms, and further the latency variations of the brain response were directly carried over to behavioral reaction times. Thus, noninvasively measured transient events in the human auditory cortex can be used to predict accurately the temporal course of sound detection and may therefore turn out to be useful in clinical settings.

Adult↗

PubMed2003

Visual short-term memory load affects sensory processing of irrelevant sounds in human auditory cortex.

We used whole-head magnetoencephalography (MEG) to investigate neural activity in human auditory cortex elicited by irrelevant tones while the subjects were engaged in a short-term memory task presented in the visual modality. As compared to a no-memory-task condition, memory load enhanced the amplitude of the auditory N1m response. In addition, the N1m amplitude depended on the phase of the memory task, with larger response amplitudes observed during encoding than retention. Further, these amplitude modulations were accompanied by anterior-posterior shifts in N1m source locations. The results show that a memory task for visually presented stimuli alters sensory processing in human auditory cortex, even when subjects are explicitly instructed to ignore any auditory stimuli. Thus, it appears that task demands requiring attentional allocation and short-term memory result in interaction across visual and auditory brain areas carrying out the processing of stimulus features.

Acoustic Stimulation↗

PubMed2002

Human cortical dynamics determined by speech fundamental frequency.

Evidence for speech-specific brain processes has been searched for through the manipulation of formant frequencies which mediate phonetic content and which are, in evolutionary terms, relatively "new" aspects of speech. Here we used whole-head magnetoencephalography and advanced stimulus reproduction methodology to examine the contribution of the fundamental frequency F0 and its harmonic integer multiples in cortical processing. The subjects were presented with a vowel, a frequency-matched counterpart of the vowel lacking in phonetic contents, and a pure tone. The F0 of the stimuli was set at that of a typical male (i.e., 100 Hz), female (200 Hz), or infant (270 Hz) speaker. We found that speech sounds, both with and without phonetic content, elicited the N1m response in human auditory cortex at a constant latency of 120 ms, whereas pure tones matching the speech sounds in frequency, intensity, and duration gave rise to N1m responses whose latency varied between 120 and 160 ms. Thus, it seems that the fundamental frequency F0 and its harmonics determine the temporal dynamics of speech processing in human auditory cortex and that speech specificity arises out of cortical sensitivity to the complex acoustic structure determined by the human sound production apparatus.

Adult↗

PubMed2002

Cortical processing of speech sounds and their analogues in a spatial auditory environment.

We used magnetoencephalographic (MEG) measurements to study how speech sounds presented in a realistic spatial sound environment are processed in human cortex. A spatial sound environment was created by utilizing head-related transfer functions (HRTFs), and using a vowel, a pseudo-vowel, and a wide-band noise burst as stimuli. The behaviour of the most prominent auditory response, the cortically generated N1m, was investigated above the left and right hemisphere. We found that the N1m responses elicited by the vowel and by the pseudo-vowel were much larger in amplitude than those evoked by the noise burst. Corroborating previous observations, we also found that cortical activity reflecting the processing of spatial sound was more pronounced in the right than in the left hemisphere for all of the stimulus types and that both hemispheres exhibited contralateral tuning to sound direction.

Acoustic Stimulation↗