com/locate/ynimg NeuroImage 41 (2008) 561 – 574

EEG default mode network in the human brain: Spectral regional field powers
Andrew C.N. Chen,⁎ Weijia Feng, Huixuan Zhao, Yanling Yin, and Peipei Wang
Center for Higher Brain Functions, Capital Medical University, Beijing 100069, China Received 17 September 2007; revised 12 December 2007; accepted 19 December 2007 Available online 15 January 2008

Eyes-closed (EC) and eyes-open (EO) are essential behaviors in mammalians, including man. At resting EC-EO state, brain activity in the default mode devoid of task-demand has recently been established in fMRI. However, the corresponding comprehensive electrophysiological conditions are little known even though EEG has been recorded in humans for nearly 80 years. In this study, we examined the spatial characteristics of spectral distribution in EEG field powers, i.e., sitting quietly with an EC and EO resting state of 3 min each, measured with high-density 128-ch EEG recording and FFT signal analyses in 15 right-handed healthy college females. Region of interest was set at a threshold at 90% of the spectral effective value to delimit the dominant spatial field power of effective energy in brain activity. Low-frequency delta (0.5–3.5 Hz) EEG field power was distributed at the prefrontal area with great expansion of spatial field and enhancement of field power (t = – 2.72, p b 0.02) from the EC to the EO state. Theta (4–7 Hz) EEG field power was distributed over the fronto-central area and leaned forward from EC to the EO state but with drastic reduction in field power (t = 4.04, p b 0.01). The middle-frequency alpha-1 (7.5– 9.5 Hz) and alpha-2 (10–12 Hz) EEG powers exhibited bilateral distribution over the posterior areas with an anterior field in lower alpha-1. Both showed significantly reduction of field powers (respectively, W = 120, p b 0.001 for alpha-1; t = 4.12, p b 0.001 for alpha-2) from EC to the EO state. Beta-1 (13–23 Hz) exhibited a similar spatial region over the posterior area as in alpha-2 and showed reduction of field power (t = 4.42, p b 0.001) from EC to the EO state. In contrast, high-frequency beta-2 and gamma band exhibited similar, mainly prefrontal distribution in field power, and exhibited no change from EC to the EO state. Corresponding correlation analyses indicated significant group association between EC and EO only in the field powers of delta (r = 0.95, p b 0.001) and theta (r = 0.77, p b 0.001) band. In addition, the great inter-individual variability (90 folds in alpha-1, 62 folds in alpha-2) in regional field power was largely observed in the EC state (10 folds) than the EO state in subjects. To summarize, our study depicts a network of spectral EEG activities simultaneously operative at well defined regional fields in the EC state, varying specifically between EC and EO states. In contrast to transient EEG spectral rhythmic dynamics, current study of long-lasting (e.g. 3 min)

spectral field powers can characterize state features in EEG. The EEG default mode network (EEG-DMN) of spectral field powers at rest in the respective EC or EO state is valued to serve as the basal electrophysiological condition in human brain. In health, this EEGDMN is deemed essential for evaluation of brain functions without task demands for gender difference, developmental change in age span, and brain response to task activation. It is expected to define brain dysfunction in disease at resting state and with consequences for sensory, affective and cognitive alteration in the human brain. © 2008 Published by Elsevier Inc. Keywords: EEG field power; 90% effective power; Regional distribution; Default mode network; EC-EO state; Individual differences

Introduction EEG and Brain Function/Dysfunction EEG reflects vital brain activities, in fetal (Preissl et al., 2004) and from neonate (Vanhatalo and Kaila, 2004) to aging (Cummins and Finnigan, 2007) in health and in disease. The function of EEG in the sleep/awake state (Chornet-Lurbe et al., 2002), perception (e.g. Basar et al., 2006; Andrade and Bhattacharya, 2003), attention (Vuilleumier and Driver, 2007), memory (Klimesch et al., 2006a; Wilding and Herron, 2006), emotion (Cacioppo, 2004), thinking (Srinivasan, 2007), as well as dysfunction in epilepsy (Benbadis, 2006), autism (Kagan-Kushnir et al., 2005), dyslexia (Arns et al., 2007), AD/HD (Barry et al., 2007; Hobbs et al., 2007), anxiety (Smit et al., 2007), depression (Hunter et al., 2007), MCI (van der Hiele et al., 2007; Buchem et al., 2007), dementia (Babiloni et al., 2006; Rossini et al., 2006), schizophrenia (Prichep, 2007), coma (Husain, 2006), vegetative state (Guerit, 2005), and even death (Sediri et al., 2007) is largely documented. In fact, EEG dynamics is closely impacted in man's whole life. In additional to genetic factors (Begleiter and Porjesz, 2006), one of the main effects on the EEG is the state of eyes-closed (EC) and eyes-open (EO) differentiation at rest. In the course of one's life, we often consciously or unconsciously lapse into an eyes-closed state, sometimes transiently

⁎ Corresponding author. E-mail address: ac@cpums.edu.cn (A.C.N. Chen). Available online on ScienceDirect (www.sciencedirect.com). 1053-8119/$ - see front matter © 2008 Published by Elsevier Inc. doi:10.1016/j.neuroimage.2007.12.064

Sorg et al.g.. Neuper et al.N. Awealth of information is perhaps under-exploited regarding the spectral energy changes at regional brain areas. Zhao et al. 2006). and the study was approved by the local ethical committee... vegetative state or death. 2007) and is of great interest in that element for the default-mode network (DMN) or connectivity is readily measured from PET (Raichle et al. (b) how these regional spectral field powers varies from the EC to EO state.. 2001). 2006). 2007. Castellanos et al.. 2001. anxiety disorder (Zhao et al. The 3D 128-ch montage and nomenclature is depicted in Fig. Klimesch et al. much less up to now as a set of spectral field energy in the spatial domain. Measurement in health can be related to: (a) stable oxygen extraction function of organized resting brain function and suspended upon specific goal-directed behaviors (Raichle et al. 2007. in that order. Issues When one closes his/her eyes. 2007). in press). 2006. After rejection of EOG contamination and non-specific artifacts.C. 2001).. “Bad electrodes” were replaced with the extrapolated virtue values from the neighboring electrodes.. each set of EEG data (2-s epoch) was subjected to Fast-Fourier Transform (FFT) analysis to obtain the absolute EEG band power (μv2) at each electrode in the . the fMRI-DMN alternations in brain malfunctions are now demonstrated in fragile X syndrome (Menon et al. Garrity et al.. Geday et al..562 A... in a quiet room (temperature: 22–24 °C) at shaded daylight. EEG data were sampled at 512 Hz and electrode impedance was kept lower than 5 kΩ.. Experimental design Each subject was asked to sit in an armchair. the distinction of the eyes-closed condition is well studied but the eyes-open resting condition has been infrequently made. Each subject was not given any instruction but asked to stay fully relaxed. 2007). in a dysfunctional mode of coma. 1996. Subsequently. All the EEG channels were recorded in an averaged reference but were offline bilaterally re-referenced (averaged A1 + A2). 2001). (i) essential patterns in the DMN (Damoiseaux et al. in press.. 2005. “The Default Mode Brain State” defines a baseline state in the human brain lying quietly with eyes-closed (Raichle et al. and (j) light-sleep.. Each subject was asked to keep the eyes-closed for 3 min and eyes-open for another 3 min. 2007. 2004). (c) the individual characteristics of regional spectral field powers in both states. EEG has largely been dealt with as a set of isolated frequency fluctuations in the temporal waveform domain. throughout the duration of the experiment. respectively with the A.. Tourette's syndrome (Marsh et al. mild cognitive impairment (Rombouts et al.N. using Ag/AgCl electrodes and two channels of electro-oculogram (EOG) were continuously gathered for 3 min during the EC and EO states. The electrodes were mounted according to the 10/5 montage system (Oostenveld and Praamstra. Bellec et al.. Enschede. in press. Chen et al. 2007). (f) intrinsic visual. Yet the majority of time in the remaining hours we live in an eyes-open state. 2006. The portion of EOG contamination of each scalp trace was removed in the off-line analysis. 2003). 2008). At the resting eyes-closed state.. it was argued recently that the brain is in a “default mode” when devoid of any external task (Raichle et al.T. schizophrenia (Harrison et al. The artifact rejection methods consisted of exclusion in epoch with large amplitude (over ±80 μV). unless under adequate anesthesia. and (d) if some associated networks of regional spectral field powers exist in either EC or the EO state. and slow eye movement coincident with EOG. in press... epilepsy (Laufs et al. Damoiseaux et al. major depression (Greicius et al. 2004). From the literature on the default mode of brain function in fMRI.5–100 Hz bandpass filter off-line and were subjected to epoching (2 s each). and artefact rejection pre-processing. in press). 2006). 2006.. bipolar disorder (Calhoun et al. somatosensory and olfactory sensory cortical activation in silence at rest (Hunter et al. Materials and methods Subjects Fifteen healthy female volunteers (age: 20–26 years old) from the University community participated in the study. Rombouts et al. our knowledge of EC and EO in nearly 80 years of studying EEG is limited to a few transient classical “alpha block” (Pollen and Trachtenberg. largely at the visual field of the occipital brain upon EC to EO state. attention disorder (Helps et al. 2001. Netherlands).. 2007) and fMRI (e... 2007). Bilateral EOG was recorded from the horizontal and vertical sites to monitor blinking or eye movements. 2007). most of the study conditions comparing the fMRI-DFN are relied on the EC but little work on EO state. Damoiseaux et al. (e) slow oscillation between internal introspection and external extrospection of brain state at rest (Fransson.. However. Written informed consent was obtained from each subject in accordance with the Helsinki Declaration.... Esposito et al. (h) age-related DMN (Grady et al.. Subjects were excluded from the study if they had any physical/psychological problem or were using medication. EEG System (A.. 1. 2006).. 1972) or contemporary “alpha desynchronization” (Pfurtscheller et al. 2001). 2007). 2006. Calhoun et al. a different opinion on the nature of fMRI-DFN is also recently raised (Fransson. one never ceases feeling and thinking. 2005) and Alzheimer's disease (Wang et al. 2007)... EEG data were filtered with 0. or the ventral medial prefrontal cortex and posterior cingulate cortex exhibiting largest activation at rest and deactivation upon tasks (Greicius and Menon. 2006) studies. (d) network connectivity of ventral anterior cortex and post cingulate cortex enhanced during resting state (Greicius et al. Study aims Taking into consideration the aforementioned. other than the alpha-block studies. auditory. 2001)... Raichle and Synder. Wilson et al. this study was aimed at examining (a) the full spatial distribution of the spectral field powers at the basic EC state. 2006).. DC bias. 2006). In contrast since Hans Berger. EEG recordings and pre-treatment of EEG A high-density 128-ch EEG recording. In state of eyes-open. In neuroimaging.N. blinks.. even in the absence of conscious awareness during light sleep (Horovitz et al. (g) attention and memory processing (Fransson. briefly in minutes or even in fraction of hour not counting napping-sleep.. linear-detrend. (c) dorsal mPFC increased in internal feeling judgment and ventral mPFC reduced in external cognitive judgment (Gusnard et al. or slow respiratory activity (Birn et al. the brain is bombarded with tremendous bits of visual stimuli. / NeuroImage 41 (2008) 561–574 within a second. (b) both medial prefrontal cortex and anterior cingular cortex are embedded in an organized network for initiation of the functional “self” (Gusnard et al. while EEG was recorded.T. 2007).

Fig. Beta-2 (24– 34 Hz) and Gamma (35–45 Hz). were subjected to further analyses. EEG recording electrode sites and nomenclatures. These broad bands were defined by the conventional IFCN guideline (Nuwer et al.5–3.N.5 Hz).C. 1999). 1.A. Out of them. but marked reduction of theta field power were respectively shown from EC to EO state. .. theta (4–7 Hz). each 3-min period EEG was analyzed in 2 sec epochs. on average. 2. Superior-anterior view (left panel) and superior-posterior view (right panel) perspectives of the 128-ch EEG electrode nomenclature in 10/5 montage. Alpha-2 (10–12 Hz).5–9.. following 7 bands: Delta (0. Increase and expansion of delta field power.N. For each EC and EO state the valid epochs were averaged after procedure of fast Fourier Transform (FFT) performed by ASA 3.5 Hz). EO states.0 software (A.T. Beta-1 (13–23 Hz). resulting in 90 epochs. For each EC and EO state. Alpha-1 (7. Klimsch. Chen et al. / NeuroImage 41 (2008) 561–574 563 Fig. Delta field power was distributed over the prefrontal area and the theta field power at the fronto-central area. Topographic spectral mapping of low-frequency delta and theta field powers in EC vs. Doppelmayr et al. around 70–80 valid epochs at the EC state and 60–80 epochs at the EO state across the 15 subjects. 1999) but not according to individuals' spectral characteristics (cf. 1998.

1987). The grand averages of FFT maps of different stages in topographic maps (256 hues) of the mean amplitude of the surface EEG power were calculated on a 3D “quasi-realistic” cortical model by a spline interpolating function (Tandonnet et al.9 = 36.60 62. In this manner..1 = 9. Subjects were ordinally listed from the EC state.0/3. as dependent measure.70 20.59 36.9/1. alpha-1.9 = 12. we choose to use the 5th percentile ranking value. From each spectral EEG band. Right panels: Correlation analyses indicated significant associations of the values between EC and EO states for delta and theta. Netherlands).7 = 4.00 20.91/10.7 = 3.86 = 0. In this study. 3. from the zero to maximum value of all the 128 EEG sites.00 22. Through automatic scaling.N.18 EO max/min = ratio 582/90 = 10.69 calculated.15 5. but radical reduction of theta field power from EC to EO. a scalp field power spectral map was interpolated from the 128-ch EEG and the site/electrode of maximal power was isolated with its value calculated.16/7. alpha-2. 2006.C.7/1.1 = 9.3/4.3 = 90.6/9.5 = 7. which might occur.54 20. the use of 5th percentile ranking value for EV was more conservative and would effectively exclude the probable outlier maximum electrode (such as from a bad electrode site with a spurious effect on the EEG). the 90% EV (effective value) was the sum of all EEG powers from the electrodes in the ROI.5/2. In the study.5/2.16 33.12 = 7. As in the mapping of brain activity by PETfMRI.34 62. The EV was defined by the 5th upper percentile ranking value of the 128 electrodes.6/3. Thus. Region of Interest (ROI) The total field power (sum of the electrode spectral power) in each spectral EEG bands within a region of interest (ROI) was Table 1 Magnitudes of the individual differences at the EC and EO state (field power in μV2) EC max/min = ratio Delta Theta Alpha-1 Alpha-2 Beta-1 Beta-2 Gamma 845/151 = 5.3/6. beta-2 and . the range of maximum and minimum values in EEG spectral power was established.20 90. EEG activity in this study is based on the threshold criterion.62 = 1. The ROI was spatially delimited by the area exhibiting the band power value in electrodes exceeding a critical value that equaled to 90% of the effective value (90% EV). Chen et al.62 29.564 A.0 = 2.18/21.3 = 21. For each of the 7 EEG bands. respectively in each of the 7 broad bands at the EC and EO state. From the threshold criterion.. 2005.40 230/2.54 = 10. we defined the ROI field power as the dominant spectral EEG field power.4/1.10 5.7/8. instead of the maximum value (the EEG site of the highest rank).86 12. the maximal value has been used in the calibration range. Enschede. Greater inter-individual variability was noted in EC than in EO state for both field powers. above 90% of the effective value. beta-1. However. for the sake of preventing the outlier electrode value. the effective size of spatial area and magnitude of activity can be defined and measured. Left panels: t-tests of EC and EO field power showed significant increase of delta. while the measured magnitude was the “dominant EEG power”..0 = 62. termed effective energy.4/2. Often. / NeuroImage 41 (2008) 561–574 Fig.91 29. the 90% EV of spectral EEG well delimited the dominant regional field power respectively for delta. Perrin et al.3/5.1 = 5.59 12.15 = 1.42 EC/EO 5.6/2. Babiloni et al.42 = 1. theta. The measured area in this study was the “region of interest”. separate spectral EEG power was calculated based on the sum of the measured power of all electrodes in the ROI.7/0.54 11.54 4.2 = 2.

4. If not. Chen et al.05 was accepted as significant. the expansion of the delta field power in the left fronto-lateral region was noted. These effects were verified and presented in Fig. the dominant delta field power was found to be at the prefrontal area for both the EC and the EO states. the Wilcoxon test for nonparametric comparison of ranks and Spearman correlation of ranks were employed to evaluate the probability significance. if the data sets were confirmed to normality distribution. In our past experience. In this manner. / NeuroImage 41 (2008) 561–574 565 Fig. while the upper alpha-2 field power was restricted at the posterior area. 4.A. Chicago. the spatial template of delta field power was applied by the EO state since it exhibited higher 90% EV than that of the EC state. Thus. while the rest of spectral powers were applied by the EC state as these values were higher at the EC than EO state. 3. the ROI territory that contains the largest electrode array was selected as a template. In comparison. dependent t-test of means and the Pearsons correlation test on the association of spectral field powers between these two states were used. Topographic spectral mapping of the middle-frequency alpha-1 and alpha-2 field powers. p b 0. The alpha-1 field power was distributed mainly at the posterior area with an anterior extension. 1. i. the sum of the electrode values within the ROI in each subject was calculated for statistical analysis.04v (SPSS Inc. Results Low-frequency EEG field power Spatial Mapping In Fig.e. 2. Both showed marked reduction in the field power from EC to EO state. The dominant 90% EV of the spectral field power in an EEG delimited a spectral regional area with 10-spacings set in each spectral field power range: below 10% in blue and above 90% in red between the range of zero and the top 90% EV (see the calibration bars in Figs. The statistical program SigmaStat 3. In all tests. As a general principle. The t-tests independently demonstrated a marked enhance- . 6). 2007). gamma broad bands in one of the two states. to rely on the original recording for straightforward interpretation of EEG values.C. but not a power value of a single electrode. In this study. theta power was distributed at the fronto-central area.g. Statistical analysis Statistical comparisons were performed on the scalp EEG field power by paired t-test for comparison of EC and EO states. A grand average of the resting EC and EO states was computed then the statistical effects of the state change were conducted on the effective spectral field power at the regional areas in each of the 7 bands. without logarithmic transformation of the EEG values (e. For statistical comparison between states or across conditions.N. Effect of the change of states From the EC to the EO state. but reduction of theta field power in amplitude and space at the fronto-central area was clearly shown. the dependent variable of interest was the field power of array of electrodes. We elected to use the raw values. van Albada and Robinson. log transformation of EEG values cannot completely eliminate the problem of statistical normality in the distribution of EEG values. USA) was used for statistical procedures. centered over the vertex for both (EC and EO) states.

12 folds in the EO state for the upper alpha-2 field power were noted (see Table 1). but 9. . p b 0.001.7 folds increase in EC state and a 8.54 fold at the EO state (see Fig. Middle-frequency EEG field power Spatial Mapping In Fig.91 folds in the delta power between the maximum and minimum values in the subjects at the EC state. median 4.3 μV2. For the theta field power.9 μV2. respectively. the great inter-individual variability in EEG powers of alpha-1 and alpha-2 was observed to be limited to the resting EC state unlike the EO state.001 and r = 0. t = 4. t = -2.77.4 μV2.00 at the EO state (see Fig.001). but only 0. EO: mean 6. marked greater inter-individual variability was shown respectively at EC than EO state for both field powers. From the Fig. p b 0.6 folds in the alpha-1 field power between the maximum and minimum values in the subjects at the EC state. The t-tests independently indicated a marked reduction of the alpha field power (EC: mean 54.72.7. Table 1). both alpha-1 and alpha-2 field powers were not discernible in the regional topography under the same scaling at the EC state. median 41. 3 depict the magnitude of the variability reached as high as 5. while a great reduction in the theta power (EC: mean 14. EO: mean 9. 3.9. EO: mean 407. Similar large values. p b 0. Individual's profile The individuals varied greatly in EEG powers. No correlation was shown in either alpha-1 or alpha-2 between the EC and the EO states.04. power was restricted at the posterior area only with a clear contour at the EC state.566 A. significant decrease of both alpha-1 and alpha-2 field power from EC to EO.N. the measured field powers also displayed a set of significant correlations between EC and EO states in both delta and theta powers (r = 0. b 001). respectively).1 μV2. / NeuroImage 41 (2008) 561–574 ment of the delta power (EC: mean 354. largely in the resting EC than EO state. the dominant alpha-1 field power was found to be at the posterior area along with a frontal field extension much eminently at the EC than the EO state. EO: mean 5.54 folds the EO state. 3. p b 0.6 μV2.95. 5). Subjects were ordinally listed from the EC state. Right panels: Correlation analyses indicated non-significant (ns) association of the values between EC and EO states for alpha-1 and alpha-2 field powers. Left panels: W-test (Wilcoxon rank test) and t-test of EC and EO power comparison showed. t = 4. a 62. In comparison.C. 5 right panels. 5. The right panels in Fig.7 μV2. Nevertheless.2). the upper alpha-2 field Fig. the magnitude of the variability reached to peak values of 90. the magnitude and spatial extent of the alpha-1 field power was greatly reduced (Fig.4 folds at the EC and 2.6 μV2. Chen et al. Individuals profile Again. 4. the ratios were 4.12.6 μV2. However. p b 0. left panel. W = 120.001) was found. Effect of the states From the EC to the EO state. All of the comparisons of the inter-individual variability are listed in the Table 1. left panel. while a similar large reduction of the upper alpha-2 field power was also evident (EC: mean 23. At the EO state. Table 1).

6. 7. Topographic spectral mapping of high-frequency beta-1 distributed at posterior area. the beta-1 field power was found to be over the posterior area as in the upper alpha-2 field power and exhibited reduction from the EC to the EO state. In Fig. High-Frequency EEG Field Power Spatial mapping In Fig.12.N.8 μV2. superior-anterior (sa) and superior-posterior (sp) to . In comparison. Effect of the states The observed statistical significant difference in beta-1 field power between the EC and EO states is shown in the top-left panel of Fig. Spectral co-habilitation and correlations among the Field Powers at each EC and EO State At each of the EC or EO state. The t-test independently indicated a reduction of the beta-1 field power (EC: mean 12. p b 0. t = 4. 8. Chen et al. Large topographic change in the beta-1 field power was discerned. while no significant effects were noted in both the beta-2 and gamma field powers. Fig. a composite model in each EC and EO state was respectively made from two viewing perspectives.9 μV2.A. beta-2 and gamma field powers (both distributed at the midline frontal and prefrontal areas) in EC vs. EO state. / NeuroImage 41 (2008) 561–574 567 Fig. the dominant regional EEG field powers are shown to be a distinct set of spectral activities distributed in the head space. All correlations between the EC and the EO states in these high-frequency beta-1. 6. EO: mean 6. beta-2 and gamma field powers were proved not significant (right panels. Profile of the individuals The individual distribution profile of the beta-1 field power was greatly dissimilar and distinguished the beta-1 from those of the beta-2 and gamma field powers. 7). the high-frequency beta-2 and gamma field powers were limited mainly at the prefrontal area.001).C.

The magnitude of the individual differences. Table 2a. comparing the EC and EO states is presented in the rightist column of Table 1. the study data revealed a dramatic inter-individual variability in human EEG magnitude.001) and prefrontal . p b 0.b list the calculated cross-correlations among the 7 spectral field powers. Right panels: Correlation analyses indicated nonsignificant (ns) association of the values between EC and EO states for all the high-frequency spectral field powers. At the EC state.12 folds of alpha-1 (drastically different from that in the EC state) in the EO state. No other spatial EEG band powers in the EC state showed significant correlation (Table 2a). p b 0. Left panels: t-test of EC and EO field power comparison showed a significantly decrease of beta-1. respectively in EC and EO states. correlation analyses indicated that only the high frequency beta2 and gamma powers at the prefrontal head region were correlated (r = 0.4 of theta in the group at EC state was revealed. 8 was computerized by a virtual brain model in respective the EC and the EO state of 25 young male subjects studied in Denmark (data not published before).92. ranging from 0. the prefrontal delta field power exhibited significant correlation with the posterior beta-1 (r = 0. 7. Table 1 lists the maximum and minimum values of regional 7 spectral field powers.N.34 and 7. The ratios of maximum/minimum manifested to a lesser differentiation at the EO state.86 of the delta field power to 8. The ratios in variability between the EC and EO states for alpha-1 and alpha-2 were 10. Great inter-individual variability in each spectral field power was noted. Chen et al.001).568 A. demonstrating markedly greater variability in the group at the EC than in the EO state for alpha-1 and alpha-2 field powers. but W-tests (Wilcoxon rank test) of EC and EO power comparisons showed no significant (ns) effects of beta-2 and gamma field powers from EC to EO state. This Fig. Subjects were ordinally listed from the EC state. in contrast.C.86. Again. In 15 subjects a high end of 96.10 folds respectively. illustrate the coexistence of these spectral activities in space. At the EO state. / NeuroImage 41 (2008) 561–574 Fig.0 folds of alpha-1 and low end 4.

For example. Dominant EEG Spectral Field Power and Function The spatial nature of the field EEG power may reflect some underlying co-activity in structure-function relations.A. at EC vs.52. Though it is tempted to categorize it as some residue of blinking/eyesmovement generated in the prefrontal subcortical site. This spatial distribution of the field EEG powers has not fully been appreciated. the eminent prefrontal delta EEG field power (see Fig. In EEG studies.) perspectives.. 8. the relations of regional EEG field powers in listening to music compared to lecture (speech. Constellation of spectral field powers measured in a 3-min period depicts the regional distribution of the respective seven frequency band powers on the brain topography. while the author Chen was directing the Human Brain Mapping Laboratory. 3 min eyes-closed) and variety of emotional ratings are effectively revealed.036). p = 0. an important off-press publication indicated that bold fMRI can be correlated with EEG powers in six widely distributed resting state networks. it is necessary to define not only the spectral information but also the spatial characteristics of the EEG of subjective experience. 2) could indicate some prefrontal metabolic/functional significance. EO state. Denmark. gamma field powers (r = 0. characterized by EEG features involved in combination with different brain rhythms (Mantini et al. The resting EEG DFN was composed from data gathered in 25 healthy college young males from Aalborg. Data were not published previously. / NeuroImage 41 (2008) 561–574 569 Fig.C. Chen et al. termed EEG default mode network (EEG DMN). Discussion EEG DMN The main study findings indicate that the resting EEG in eyesclosed state devoid of stimulation or task is composed of a defined set of regional spectral activities in the classic 7 broad bands. After completing the current work. The likelihood of a blinking artifact on the prefrontal delta EEG was diminished since our artifact algorithm was set at a threshold to eliminate all epochs exceeding 80 μV such as .N. In addition. 2006).046). two facts argue against it.554. the frontocentral theta field power was correlated with the anterior-posterior alpha-1 field power (r = 0.) and superior-posterior (s. as to validate the nature of EEG DMN and brain functions. viewed from superior-anterior (s. p. while the prefrontal beta-2 and gamma field powers were significantly correlated (Table 2b). The distribution maps of spatial regional field powers are consistent with a previous observation using Danish subjects (Chen et al. a.. 2007). p = 0. submitted for publication). Extending from current work in a preliminary report (Feng and Chen.

2006).. 2007). However. Correlations of the averaged power of alpha EEG (8–12 Hz) and bold fMRI signals can be found in occipital.. It is proposed to call beta-2 as the high-frequency alpha-3. we observed a frontal extension along with a more prominent posterior distribution in the lower alpha-1 field power (Fig.. 2006b) has been described in function but little is known on the spatial differentiation. Also. Due to their tiny EEG power. a positive correlation of medial frontal cortical metabolism with delta EEG power has been established (Alper et al. Our finding of the theta EEG field power distributed eminently in the fronto-central area (see Fig. the observed beta-1 field power in regional distribution. We now are in the process to validate the cortical and subcortical structures of the observed regional scalp EEG field powers described in this study. submitted for publication). extending from the conventional occipital alpha EEG. no correlation of the blinking rate and the prefrontal EEG field power could be established (r = 0. 2007). In the middle-frequency alpha EEG. the neural source of alpha EEG is revealed in the brain region of distinct cortical areas... 2006). 2007). beta-2 and gamma EEG field powers. In high frequency beta-1. as it may explain 65% of the variance in the delta power (Boord et al. Of special interest is the site of eminent gamma field power at the dorso-lateral frontal areas in the left hemisphere.. 2) is consistent with the known frontal midline theta EEG activity (Inanaga. At resting state. This conjecture may be viable with the EO state. 2005) as well as the amplitude of delta (2–4 Hz) sources with the prefrontal white matter across the mild cognitive impairment and Alzheimer's patients (Babiloni et al.570 A. Recently. by current source imaging of LORETA on spectral EEG field powers in Talairach standard brain modeling.344.4/min).. a plausibility that further remains to be proven. The great similarity in distribution at the bilateral temporal area and the prefrontal area in the beta-2 and gamma EEG field powers (Fig. 2000). The regional preeminent theta EEG has been related to infant arousal (Grigg-Damberger et al. which is noted in the differentiation of music from speech perception (Feng and Chen. p-values in italics. attention (Sauseng et al. parietal and even frontal lobes (Goncalves et al.5–4 Hz) power correlates with brain atrophy in Alzheimer's disease patients (Fernandez et al. 1997. thought to reflect binding of perception in brain function (Bertrand and Tallon-Baudry. It is generated in the medial prefrontal cortex and/or anterior cingulate cortex in monkey (Tsujimoto et al. memory (Onton et al. 4). 2004). significant values in shaded field) the blinking on the EEG (150 μV). The transient beta activity in motor function is associated with fMRI bold activation of the sensorimotor cortex (Parkes et al. and visual alpha EEG in perceptual function is from occipital lobes (Goncalves et al. while the transient gamma activity in visual perception with bold activation of the calcarine sulcus (Hoogenboom et al. the distribution of these activities at some discrete field opens to new challenge for understanding the structurefunction relations in beta-gamma EEG. and cognitive aging (Brickman et al. In the theta EEG power. such that the sleep spindle alphaEEG is likely from thalamus. the regional territory of the eminent alpha-1 and alpha-2 field powers reside more on the parietal area than occipital area (see Fig.. 1998). 2006) and in man (Cahn and Polich. the major changes largely in reduction of the field power. little spatial information of the beta and gamma EEG field in human brain has been reported in literature. music (Feng and Chen. the delta (1. suggests that beta-1 as a spectral lingering of alpha-2 activity and belongs to the same family.C. 2006). 53% of the variance over age span is reported to be explained by cerebral metabolic rate (Boord et al. 2006). perception (Basar et al. Recently. 2005). the mu-rhythm in motor function is from central sulcus. 4) either in congruent or at variance with the resting anterior alpha asymmetry in emotion (Cacioppo. 2007). significant values in shaded field) .N. The spectral field power in lower alpha-1 differentiated the upper alpha-2 field power with no anterior extension and only the similar posterior portion is of interest. Klimesch et al. submitted for publication). p = 0. 2006). similar to that of alpha-2 at the posterior scalp area (Fig. 2005).... 2007). p-values in italics. 2006)...b) renders these beta-2 and gamma EEG in the same family. are the Table 2b EO state: cross correlations of the regional spectral field powers (correlation values in bold. The lower and upper alpha in EEG literature (Klimesch. In the past. State changes The spectral topographic distributions showing little changes attest the stability of the underlying structures associated with the EEG field powers.21). it has been shown that EEG is associated with cerebral metabolic rate.. This argument was also not tenable as nearly no blinking episode was found in the EEG record during the EC state (only one blink across the 15 subjects). Nevertheless. Chen et al.. 2007). emotion (Guntekin and Basar. in the range less than 3 μV in amplitude.. Instead. which exhibited high frequency of blinking (mean: 37.. 6) and their strong correlation at both the resting EC and EO states (Table 2a. This observed alpha-1 and alpha-2 field powers in territory may lead to reconsider the important aspect of frontal and parietal alpha EEG in source and function. 2006) but with wide individual variability at rest (de Munck et al. 2006). 6). / NeuroImage 41 (2008) 561–574 Table 2a EC state: cross correlations of the regional spectral field powers (correlation values in bold. except the increase of prefrontal delta.

regardless of the state. Using high-density 128-ch montage. The robust correlations of high-frequency beta-2 and gamma field powers in both states do suggest some invariant /function of these activities at the prefrontal area. Additionally. the EEG DMN is aimed at defining different EEG states. and Capital Medical University for Innovation Awards. Babiloni et al. it is an advantage to invoke states which may enhance field powers. Given the high magnitude of spectral field powers in the EC state. in the study. In current study.. This EC and EO differential states have recently not explicitly elucidated in the fMRI literature as the default mode of brain function. Implication Methodologically. time-episode. the current study strategy is ready and opens up a new avenue for the examination of the EEG DMN in gender differences. remain to be examined. our approach in the delimiting the dominant regional distribution of spectral EEG field power is an important extension of the conventional quantitative EEG. Acknowledgments This study was supported by several grants respectively from the Chinese National Science Foundation for a project on “Brain and Anesthesia” NSF-30770691. largely defined by the lack of explicit task demands on the brain in subjects in the EC state. By considering the individual normalization. as much as 90 folds in alpha-1 and 62 folds in alpha-2 field powers (see Table 1) shall not deter scientific investigation in EEG research. while these two high-frequency field powers are mutually correlated at the same area. Inter-individual variability in EEG field powers and the inter-state consistency Two aspects of the inter-subject variability deserve special attention. many variables used for processing the EEG (electrode density. The surprising markedly inter-individual variability. FFT. Beijing Municipal Government for Advancement of Sciences.. Chen et al. these changes may occur intrinsically from the EC to the EO state both in terms of physiological visual information processing and psychological terms of visual perception in seeing. the EEG default mode network (DFN) at rest with eyes-closed of the dominant spectral field powers entails a constellation of large low-frequency delta activity at the prefrontal area.A. as much as 90 folds in alpha-1 and 60 folds in alpha-2 in the group of 15 subjects. relative EEG transformation and use of non-parametric statistics with adequate sample size.N. It seems that non-parametric Wilcox-test of ranks can be justified and similarly non-parametric Spearman correlation test is appropriate for evaluating statistical features. 2005). . Firstly. In the eyes-closed state. 2007. In the field of EEG research. are in magnitudes higher than expected. Though there is some knowledge that a 10–15% of the normal healthy subjects have little or nearly no alpha activity. The EEG DMN of the spectral field power in the eyesclosed or in the eyes-open state at rest. is readily served as basal brain condition for the comparison of activation challenge or dysfunctional alteration in diseases of the brain. 1972) or modern “alpha desynchronization” (Rihs et al. In general. the delta field power is enhanced at the prefrontal area while the theta. EC or EO.. The reason for this in neurophysiology remains totally unknown. the observed changes in spectral EEG field powers at different regional distribution attests to the complex and consorted EEG changes from EC to EO at rest. the prefrontal delta field power is correlated with both beta-1 and gamma field powers. devoid of task demands. qEEG. Several reports of the brain state in the default mode from fMRI studies include “self-referential mental activity” (Gusnard et al. mapping (Duffy et al.. Greater inter-individual variability in field power can be seen in the eyes-closed than the eyes-open state. 2006).. “exteroceptive and interoceptive deployment of attention” (Nagai et al. epoch. The emphasis is on the field potential. and “inner-speech” (Hunter et al. we selected the 90% EV (see Methods) as a sensible threshold to delimit a reference cluster of field power for comparison of states. if is corresponding to the above hypothesis. frequency bands. over conventional use of selective electrode spectral power in an interpolated map. effective-energy threshold) require standardization to facilitate useful comparison across conditions and among laboratories. reference. recoding. the spectral dominant EEG field power can be extracted in region of interest (ROI) for robust comparisons and is based on the “effective-energy” concept. alpha-2 and beta-1 powers are reduced in the respective areas. sensible and viable EEG/brain functions are deemed fully worthy for future investigation. Conclusion In a group of 15 healthy females.. Such results of strong modulation at various spectral EEG field power greatly expand our understanding of the limited classical “alpha-block” in EEG from EC to EO (Pollen and Trachtenberg. effort should be directed toward the innovative method for standardized automatic quantification of spectral EEG powers and differentiation of EEG states (Chen et al. submitted for publication). 2001). 2007) from the state transaction. the large differences in alpha field powers noted. In fact. This work further entrusts the steady-state comparison of experience episode in the brain. much smaller theta activity at fronto-central area. these inter-individual difference ratios are 10 folds higher at the resting EC than EO state..C. To this end. it provides an advantage in invoking states with effective EEG reduction. and high-frequency beta-2 and gamma activities at the prefrontal area. the magnitude of such differences comes as a surprise. Our observed spectral EEG field powers in the regional distribution between the EC and EO states. Given the low magnitude of field powers in the EO state. The observed 10 folds of alpha variability in EC over EO shall be taken into account. alpha-1. Apparently. in a discrete area. 2001). lying at rest (Raichle et al. little spectral EEG field powers (except delta and theta) showing significant correlations may indicate two divergent networks between the EC and EO state. 2006). on the development of age-span in normal brain function as well as their changes upon experimental stimulation or task demand. / NeuroImage 41 (2008) 561–574 571 main manifestation of the EC to EO state. the EEG DMN is valuable in the examination of alterations in patients with diseases and the dysfunctional responses to their environment (cf. Since no task or instruction was given to the subjects at the resting states. Laufs et al. the subject variables include the choice of the resting basal state. Our study on listening to ‘happy' music /'sad' music compared to that of lecture speech bears it out (Feng and Chen.. expressed in EEG state. In the current study. alpha-2 and beta-1 at the posterior area. 1994). 2004). alpha-1 activity at the anterior-posterior area. Secondly.. “mind-readiness alert” (Fransson 2005). These results warrant consideration as a practical issue and call for re-consideration in statistical parametric comparisons of means. Compared to the eyes-open resting state. Likewise.

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