Memory traces for words as revealed by the mismatch negativity.

Brain responses to the same spoken syllable completing a Finnish word or a pseudo-word were studied. Native Finnish-speaking subjects were instructed to ignore the sound stimuli and watch a silent movie while the mismatch negativity (MMN), an automatic index of experience-dependent auditory memory traces, was recorded. The MMN to each syllable was larger when it completed a word than when it completed a pseudo-word. This enhancement, reaching its maximum amplitude at about 150 ms after the word's recognition point, did not occur in foreign subjects who did not know any Finnish. These results provide the first demonstration of the presence of memory traces for individual spoken words in the human brain. Using whole-head magnetoencephalography, the major intracranial source of this word-related MMN was found in the left superior temporal lobe.

Effects of acoustic gradient noise from functional magnetic resonance imaging on auditory processing as reflected by event-related brain potentials.

The processing of sound changes and involuntary attention to them has been widely studied with event-related brain potentials (ERPs). Recently, functional magnetic resonance imaging (fMRI) has been applied to determine the neural mechanisms of involuntary attention and the sources of the corresponding ERP components. The gradient-coil switching noise from the MRI scanner, however, is a challenge to any experimental design using auditory stimuli. In the present study, the effects of MRI noise on ERPs associated with preattentive processing of sound changes and involuntary switching of attention to them were investigated. Auditory stimuli consisted of frequently presented "standard" sounds, infrequent, slightly higher "deviant" sounds, and infrequent natural "novel" sounds. The standard and deviant sounds were either sinusoidal tones or musical chords, in separate stimulus sequences. The mismatch negativity (MMN) ERP associated with preattentive sound change detection was elicited by the deviant and novel sounds and was not affected by the prerecorded background MRI noise (in comparison with the condition with no background noise). The succeeding positive P3a ERP responses associated with involuntary attention switching elicited by novel sounds were also not affected by the MRI noise. However, in ERPs to standard tones and chords, the P1, N1, and P2 peak latencies were significantly prolonged by the MRI noise. Moreover, the amplitude of the subsequent "exogenous" N2 to the standard sounds was significantly attenuated by the presence of MRI noise. In conclusion, the present results suggest that in fMRI the background noise does not interfere with the imaging of auditory processing related to involuntary attention.

Working memory of identification of emotional vocal expressions: an fMRI study.

The distribution of brain activation during working memory processing of emotional vocal expressions was studied using functional magnetic resonance imaging (fMRI) in eight female subjects performing n-back tasks with three load levels (0-back, 1-back, and 2-back tasks). The stimuli in the n-back tasks were the Finnish female name [Saara] uttered in an astonished, angry, frightened, commanding, and scornful mode, and the subjects were instructed to memorize the emotional connotation of the stimuli. Subregions in the prefrontal, parietal, and visual association areas were load-dependently activated during the performance of the n-back tasks. The most consistently activated areas in the prefrontal region were detected in the inferior frontal gyrus corresponding to Brodmann's areas (BAs) 44 and 45 and in the middle and superior frontal gyri (BAs 6/8). Activation was also found in the inferior parietal lobe and intraparietal sulcus (BAs 40/7) and visual association areas including the lingual and fusiform gyri. The results suggest that a distributed neuronal network in occipital, parietal, and frontal areas is involved in working memory processing of emotional content of aurally presented information.

Working memory of auditory localization.

To investigate brain mechanisms of sound location memory, we studied the distribution of brain activation with functional magnetic resonance imaging (fMRI) in subjects performing an audiospatial n-back task with three memory load levels. Working memory processing of audiospatial information activated areas in the superior, middle and inferior frontal gyri, and in the posterior parietal and middle temporal cortices. In a control experiment, fMRI during audio- and visuospatial 2-back task performances revealed only few differentially activated subregions between the two tasks. These results demonstrate that working memory processing of auditory locations involves a distributed network of brain areas and suggest that mnemonic processing of audio- and visuospatial information is directed along a common neural pathway in the posterior parietal and prefrontal cortices.

Activation of multiple cortical areas in response to somatosensory stimulation: combined magnetoencephalographic and functional magnetic resonance imaging.

We combined information from functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG) to assess which cortical areas and in which temporal order show macroscopic activation after right median nerve stimulation. Five healthy subjects were studied with the two imaging modalities, which both revealed significant activation in the contra- and ipsilateral primary somatosensory cortex (SI), the contra- and ipsilateral opercular areas, the walls of the contralateral postcentral sulcus (PoCS), and the contralateral supplementary motor area (SMA). In fMRI, two separate foci of activation in the opercular cortex were discerned, one posteriorly in the parietal operculum (PO), and one anteriorly near the insula or frontal operculum (anterior operculum, AO). The activation sites from fMRI were used to constrain the solution of the inverse problem of MEG, which allowed us to construct a model of the temporal sequence of activation of the different sites. According to this model, the mean onset latency for significant activation at the contralateral SI was 20 msec (range, 17-22 msec), followed by activation of PoCS at 23 msec (range, 21-25 msec). The contralateral PO was activated at 26 msec (range, 19-32 msec) and AO at 33 msec (range, 22-51 msec). The contralateral SMA became active at 36 msec (range, 24-48 msec). The ipsilateral SI, PO, and AO became activated at 54-67 msec. We conclude that fMRI provides a useful means to constrain the inverse problem of MEG, allowing the construction of spatiotemporal models of cortical activation, which may have significant implications for the understanding of cortical network functioning.

Distribution of cortical activation during visuospatial n-back tasks as revealed by functional magnetic resonance imaging.

Human neuroimaging studies conducted during visuospatial working memory tasks have inconsistently detected activation in the prefrontal cortical areas depending presumably on the type of memory and control tasks employed. We used functional magnetic resonance imaging to study brain activation related to the performance of a visuospatial n-back task with different memory loads (0-back, 1-back and 2-back tasks). Comparison of the 2-back versus 0-back tasks revealed consistent, bilateral activation in the medial frontal gyrus (MFG), superior frontal sulcus and adjacent cortical tissue (SFS/SFG) in all subjects and in six out of seven subjects in the intraparietal sulcus (IPS). Activation was also detected in the inferior frontal gyrus, medially in the superior frontal gyrus, precentral gyrus, superior and inferior parietal lobuli, occipital visual association areas, anterior and posterior cingulate areas and in the insula. Comparison between the 1-back versus 0-back tasks revealed activation only in a few brain areas. Activation in the MFG, SFS/SFG and IPS appeared dependent on memory load. The results suggest that the performance of a visuospatial working memory task engages a network of distributed brain areas and that areas in the dorsal visual pathway are engaged in mnemonic processing of visuospatial information.