title: Disentangling the effects of phonation and articulation: hemispheric asymmetries in the auditory N1m response of the human brain.
abstract: 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.
  
  
  CONCLUSIONS
  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.
authors.affiliation: Apperception & Cortical Dynamics, Department of Psychology, University of Helsinki, POB 9, FIN-00014, Finland. hannu.tiitinen@helsinki.fi
authors.forename: Hannu
authors.initials: H
authors.lastname: Tiitinen
authors.forename: Anna Mari
authors.initials: AM
authors.lastname: Mäkelä
authors.forename: Ville
authors.initials: V
authors.lastname: Mäkinen
authors.forename: Patrick J C
authors.initials: PJ
authors.lastname: May
authors.forename: Paavo
authors.initials: P
authors.lastname: Alku
mesh_terms: D000161:Acoustic Stimulation / Q000379:methods; D000328:Adult; D001307:Auditory Perception / Q000502:physiology; D001921:Brain / Q000502:physiology*; D005072:Evoked Potentials, Auditory / Q000502:physiology*; D005260:Female; D007839:Functional Laterality / Q000502:physiology*; D006801:Humans; D008297:Male; D010699:Phonation / Q000502:physiology*; D013060:Speech / Q000502:physiology*