Ryusuke Kakigi

Musical Training Enhances Automatic Encoding of Melodic Contour and Interval Structure

In music, melodic information is thought to be encoded in two forms, a contour code (up/down pattern of pitch changes) and an interval code (pitch distances between successive notes). A recent study recording the mismatch negativity (MMN) evoked by pitch contour and interval deviations in simple melodies demonstrated that people with no formal music education process both contour and interval information in the auditory cortex automatically. However, it is still unclear whether musical experience enhances both strategies of melodic encoding. We designed stimuli to examine contour and interval information separately. In the contour condition there were eight different standard melodies (presented on 80 of trials), each consisting of five notes all ascending in pitch, and the corresponding deviant melodies (20) were altered to descending on their final note. The interval condition used one five-note standard melody transposed to eight keys from trial to trial, and on deviant trials the last note was raised by one whole tone without changing the pitch contour. There was also a control condition, in which a standard tone (990.7 Hz) and a deviant tone (1111.0 Hz) were presented. The magnetic counterpart of the MMN (MMNm) from musicians and nonmusicians was obtained as the difference between the dipole moment in response to the standard and deviant trials recorded by magnetoencephalography. Significantly larger MMNm was present in musicians in both contour and interval conditions than in nonmusicians, whereas MMNm in the control condition was similar for both groups. The interval MMNm was larger than the contour MMNm in musicians. No hemispheric difference was found in either group. The results suggest that musical training enhances the ability to automatically register abstract changes in the relative pitch structure of melodies.

Preferential stimulation of Aδ fibers by intra-epidermal needle electrode in humans

&NA; We recorded evoked potentials (EPs) induced by conventional transcutaneous electrical stimulation (TS), laser stimulation (LS) and epidermal electrical stimulation (ES) using a specially made needle electrode. We evaluated the activated fibers by epidermal stimulation by assessing the conduction velocity (CV) of the peripheral nerves. The EPs were recorded from Cz electrode (vertex) of the International 10–20 system in 12 healthy subjects. For the ES, the tip of a stainless steel needle electrode was inserted in the epidermis of the skin (0.2 mm in depth). Distal and proximal sites of the upper limb were stimulated by the LS and ES with an intensity which induced a definite pain sensation. Similar sites were stimulated by TS with an intensity of two times the sensory threshold. A major EP positive response (P1) was obtained by stimulation by all three types of stimuli. The P1 latency for the TS (245±22 ms) was significantly shorter than that for the ES (302±17 ms, P<0.0001) and LS (341±21 ms, P<0.0001) and the peak latency P1 by the LS was also significantly longer, approximately 40 ms, than that by the ES (P<0.0001). The CVs were 15.1, 15.3 and 44.1 m/s obtained by ES, LS and TS, respectively. The CV indicated that the fibers activated by the ES were mainly A&dgr; fibers, which corresponded to the fibers stimulated by the LS. We considered that the ES with our newly developed needle electrode was a very convenient method for the selective stimulation of the A&dgr; fibers, since it was very simple, not requiring any special apparatus, did not cause bleeding or burns and caused minimum uncomfortable feeling.

An ERP Study of Second Language Learning after Childhood: Effects of Proficiency

Whether there is an absolute critical period for acquiring language is a matter of continuous debate. One approach to address this issue is to compare the processes of second language (L2) learning after childhood and those of first language (L1) learning during childhood. To study the cortical process of postchildhood L2 learning, we compared event-related brain potentials recorded from two groups of adult Japanese speakers who attained either high or intermediate proficiency in English after childhood (J-High and J-Low), and adult native English speakers (ENG). Semantic anomalies embedded in English sentences evoked a clear N400 component in all three groups, with only the time course of the brain activation varying among the groups. Syntactic violations elicited a left-lateralized negativity similar to the left anterior negativity in ENG and J-High, but not in J-Low. In ENG, a P600 component was additionally found. These results suggest that semantic processing is robust from early on in L2 learning, whereas the development of syntactic processing is more dependent on proficiency as evidenced by the lack of the left-lateralized negativity in J-Low. Because early maturation and stability of semantic processing as opposed to syntactic processing are also a feature of L1 processing, postchildhood L2 learning may be governed by the same brain properties as those which govern childhood L1 learning. We argue that these processes are qualitatively similar in many respects, with only restricted domains of language processing being subject to absolute critical period effects.

Somatosensory evoked magnetic fields following median nerve stimulation.

Topography of somatosensory evoked magnetic fields (SEFs) following stimulation of the median nerve were investigated in normal subjects. N20m-P30m-N40m-P60m and their counterpart, P20m-N30m-P40m-N60m, were identified in the hemisphere contralateral to the stimulated median nerve. Their equivalent current dipoles (ECDs) were considered to be located in the hand area of area 3b in the primary sensory cortex (SI). Restricted deflections, P25m and N25m, were considered to be generated in area 1 in SI. Therefore, short-latency deflections less than 40 ms were considered to be hybrids of ECDs generated in areas 3b and 1. Middle-latency deflections, N90m-P90m, were considered to be generated in the second sensory cortex (SII), but they were greatly affected by the much stronger fields generated in SI. The N30m deflection, which was a magnetic reflection of the N30 potential of somatosensory evoked potentials (SEPs), were widely recorded in the frontal area. The generator site of N30 of SEPs is considered to be the supplementary motor area (SMA). However, ECDs of N30m were located in SI, and no significant ECD generated in the frontal area including SMA was detected. No significant deflections other than small N90m-P90m in SII were identified in the hemisphere ipsilateral to the stimulated nerve. No significant deflections whose ECDs were generated in the mid-parietal area were identified. In conclusion, short- and middle-latency SEFs are mainly generated in area 3b in SI contralateral to the stimulated median nerve, and responses generated in area 1 of SI and SII affect the SEFs to some degree.

Automatic Encoding of Polyphonic Melodies in Musicians and Nonmusicians

In music, multiple musical objects often overlap in time. Western polyphonic music contains multiple simultaneous melodic lines (referred to as voices) of equal importance. Previous electrophysiological studies have shown that pitch changes in a single melody are automatically encoded in memory traces, as indexed by mismatch negativity (MMN) and its magnetic counterpart (MMNm), and that this encoding process is enhanced by musical experience. In the present study, we examined whether two simultaneous melodies in polyphonic music are represented as separate entities in the auditory memory trace. Musicians and untrained controls were tested in both magnetoencephalogram and behavioral sessions. Polyphonic stimuli were created by combining two melodies (A and B), each consisting of the same five notes but in a different order. Melody A was in the high voice and Melody B in the low voice in one condition, and this was reversed in the other condition. On 50 of trials, a deviant final (5th) note was played either in the high or in the low voice, and it either went outside the key of the melody or remained within the key. These four deviations occurred with equal probability of 12.5 each. Clear MMNm was obtained for most changes in both groups, despite the 50 deviance level, with a larger amplitude in musicians than in controls. The response pattern was consistent across groups, with larger MMNm for deviants in the high voice than in the low voice, and larger MMNm for in-key than out-of-key changes, despite better behavioral performance for out-of-key changes. The results suggest that melodic information in each voice in polyphonic music is encoded in the sensory memory trace, that the higher voice is more salient than the lower, and that tonality may be processed primarily at cognitive stages subsequent to MMN generation.

Organizing sound sequences in the human brain: the interplay of auditory streaming and temporal integration

The present study examined the relationship between two of the early brain processes of sound organization: auditory streaming and the temporal window of integration (TWI). Presented at a fast stimulus delivery rate, two tones alternating in frequency are perceived as separate streams of high and low sounds. However, when two sounds are presented within a ca. 200 ms temporal window, they are often processed as a single auditory event. Both stream segregation and temporal integration occur even in the absence of focused attention as was shown by their effect on the mismatch negativity (MMN) event-related potential. The goal of the present study was to determine the precedence between these two sound organization processes by using the stimulus-omission MMN paradigm. Infrequently omitting one stimulus from a homogeneous tone sequence only elicits an MMN when the stimulus onset asynchrony separating successive tones is shorter than 170 ms. This demonstrates the effect of the TWI. Magnetic brain responses elicited by infrequent stimulus omissions appearing in a sequence of two alternating tones were recorded. The magnetic MMN was elicited by tone omission when the alternating tones formed a single stream (with no or only small frequency separation between the two tones) but not when separate high and low streams emerged in perception (large frequency separation between the two alternating tones). This result shows that auditory streaming takes precedence over the processes of temporal integration.

Human cortical area responding to stimuli in apparent motion.

APPARENT motion is the perception of the realistic smooth motion of an object which flashes first at one place then at another. To investigate human cortical responses to stimuli in apparent motion, we used a multi-channel biomagnetometer to record the magnetic fields evoked by these stimuli in four normal subjects. The results showed the presence of a localized cortical area exclusively sensitive to apparent motion stimuli that is identical to that for smooth motion. In three subjects this area corresponded to the human homologue of MT/V5. Moreover, the same region in the extrastriate cortex was involved in the short range (0.1°) apparent motion process as well as the long range (1.0°) process.

Effects of distraction on pain perception: magneto- and electro-encephalographic studies

After a painful CO2 laser stimulation to the skin, the magnetoencephalography (MEG) response (164 ms in average peak latency) was not affected by distraction, but the sequential electroencephalography (EEG) responses (240-340 ms), probably generated by a summation of activities in multiple areas, were markedly affected. We suspect that the MEG response, whose dipole is estimated in the bilateral second somatosensory cortex (SII) and insula, reflects the primary activities of pain in humans.

Disordered plasticity in the primary somatosensory cortex in focal hand dystonia

Interventional paired associative stimulation (PAS) can induce plasticity in the cortex, and this plasticity was previously shown to be disordered in the primary motor cortex in focal hand dystonia (FHD). This study aimed to test whether associative plasticity is abnormal in the primary somatosensory cortex (S1) in FHD and whether PAS modulates excitatory or inhibitory interneurons within the cortex. Ten FHD patients and 10 healthy volunteers were studied. We investigated the changes in single- and double-pulse somatosensory-evoked potentials before and after PAS, which consisted of peripheral electrical nerve stimulation and subsequent transcranial magnetic stimulation over S1. Four sessions of somatosensory-evoked potentials recordings were performed: before PAS, and immediately, 15 and 30 min after PAS. We compared the time course of the somatosensory-evoked potentials between the FHD and healthy groups. In the single-pulse condition, the P27 amplitudes were significantly higher in FHD immediately after PAS than before PAS, while no changes were observed in healthy subjects. In the double-pulse condition, significant differences in the suppression ratio of P27 were found immediately after and 15 min after PAS, while there were no significant differences in healthy subjects. The P27 suppression tended to normalize toward the level of the healthy volunteer group. In FHD, PAS transiently induced an abnormal increase in excitability in S1. In addition, intracortical inhibition in S1 was found to increase as well. This abnormal plasticity of the intracortical neurons in S1 may contribute to the pathophysiology of dystonia.

Temporal Dynamics of Neural Adaptation Effect in the Human Visual Ventral Stream

When the same visual stimulus is repeatedly presented with a brief interval, the brain responses to that stimulus are attenuated relative to those at first presentation [neural adaptation (NA)]. Although this effect has been widely observed in various regions of human brain, its temporal dynamics as a neuronal population has been mostly unclear. In the present study, we used a magnetoencephalography (MEG) and conducted a macrolevel investigation of the temporal profiles of the NA occurring in the human visual ventral stream. The combination of MEG with our previous random dot blinking method isolated the neural responses in the higher visual cortex relating to shape perception. We dissociated three dimensions of the NA: activation strength, peak latency, and temporal duration of neural response. The results revealed that visual responses to the repeated compared with novel stimulus showed a significant reduction in both activation strength and peak latency but not in the duration of neural processing. Furthermore, this acceleration of peak latency showed a significant correlation with reaction time of the subjects, whereas no correlation was found between the reaction time and the temporal duration of neural responses. These results indicate that (1) the NA involves the brain response changes in the temporal domain as well as the response attenuation reported previously, and (2) this temporal change is primarily observed as a rapid rising of “what” responses, rather than a temporal shortening of neural response curves within the visual ventral stream as considered previously.