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Spectral Correlates of Arousal and Activation

Despite reports of electrical activity in animal brains (Caton, 1895), the existence of brain potentials remained unknown and unexplored in humans until Hans Berger (1930) published his first report on electrical activity of the human brain, which he designated the electroencephalogram, or EEG (Remond & Lairy, 1972; Gevins & Schaffer, 1980). Five years passed before his findings were confirmed in the English-speaking world (Adrian & Matthews, 1934; Remond & Lairy, 1972; Petsche, 1989).

Discovery and characterization of the alpha rhythm

In his first report, Berger (1930) characterized the "waves of the first order" in human EEG, which for the sake of brevity he designated the alpha rhythm. Alpha activity consisted of large (up to 200 æV) sinusoidal waveforms 90 ms to 120 ms in duration which appeared against smaller background waves, "waves of the second order" or beta rhythm. Alpha waves were pronounced in posterior regions but could also be recorded from central and frontal areas. Most importantly, alpha activity diminished in response to stimulation and mental effort.
Between 1929 and 1937 Berger published his investigation of the relationship between mental processing and alpha activity (Remond & Lairy, 1972). He described the phenomenon of alpha blocking, an abrupt suspension of alpha waveforms in ongoing EEG, which can readily be observed when an individual with eyes closed in a relaxed state opens his or her eyes. Berger (1930) claimed that alpha blocking was independent of respiratory, vascular, or motoric responses (confirmed by Bohdanecky, Indra, Lansky, & Radil-Weiss, 1984; Valentine, 1954; Ray & Cole, 1985) and resulted when individuals attended to objects in their field of view. When individuals open their eyes in a darkened room, there is little or no change in alpha activity (Bohdanecky et al., 1984) and, conversely, alpha suppression can occur to stimulation or task demands when one's eyes are closed (Etevenon, 1986; Adrian & Matthews, 1934).
Berger (1930) concluded that directing attention toward a stimulus was responsible for fluctuations in alpha activity and the amount of alpha activity reflected the extent of mental processing (Petsche, 1989). He speculated that interhemispheric EEG synchrony was mediated by the thalamus and rejected the claim that the alpha rhythm was generated in occipital cortex exclusively (Remond & Lairy, 1972). Beyond these conjectures, he remain curiously silent on the physiological mechanisms responsible for the production of alpha activity.

Physiological mechanisms of EEG rhythmicity

Moruzzi and Magoun (1949) shed light on the origins of rhythmicity in EEG. Stimulation of the reticular formation of the brainstem produces fast low-amplitude waves in cortical recordings, usually with concomitant behavioral arousal. They concluded that the reticular formation regulated gross changes in alpha activity (i.e., nonspecific arousal). Other experiments in animals, particularly the dog, revealed that specific reactivity observed in the alpha rhythm in response to task and stimulation depended on thalamocortical networks. Alpha rhythms recorded simultaneously from visual cortex and associated thalamic nuclei (lateral geniculate, pulvinar) exhibit identical peak frequency and bandwidth, high degrees of coherence, and similar responses to stimulation (Lopes da Silva, 1978; Lopes da Silva, Van Lierop, Schrijer, & Storm van Leeuwen, 1973). The biocircuitry of the reticular thalamic nuclei (RTN) acts as a pacemaker, recruiting thalamic nuclei into 8-13 Hz (alpha) rhythms by means of powerful inhibitory postsynaptic potentials (Andersen & Andersson, 1968; Steriade & Llinas, 1988; Steriade, Gloor, Llinas, Lopes da Silva, & Mesulam, 1990). During information processing activation of specific thalamic nuclei by sensory afferences, along with localized basal forebrain excitation of RTN, which itself inhibits thalamic nuclei, results in preferential activation of certain thalamic cells. Visual perception, for example, activates neurons in pulvinar nuclei, yielding diverse (desynchronized) rates of oscillations for these cells while other thalamic nuclei remain relatively synchronized. Nonspecific afferents from the brainstem, namely the pedunculopontine tegmental and laterodorsal tegmental nuclei, also act on thalamocortical networks, affecting general tonic activity of these networks (Steriade et al., 1990). Electrical potentials measured from the scalp are presumed to be a function of postsynaptic potentials of millions of pyramidal cells in cortical layers IV and V (Lopes da Silva, 1991; Gevins, 1986).
The physiology of rhythmic activity has been summarized into a general model (Andersen & Andersson, 1968; Steriade & Llinas, 1988; Sterman & Bowersox, 1981; Steriade et al. 1990; Lopes da Silva, 1991). According to this model, metabolic characteristics of (isolated) neurons generate instrinic oscillations usually between 1 to 20 Hz. In an intact brain, activity of single neurons are regulated by pacemaking biocircuitry (RTN) which incorporates actions of single cells into larger ensembles, uniting random discharges of individual cells into simultaneous uniform volleys. Synchronization of cellular firing results in a summation of electrical potentials that can be recorded at the scalp as high-amplitude oscillations of slow frequencies (i.e., alpha rhythm). Desynchronization arises when individual neurons or neural groups are recruited out of these uniform volleys and become dedicated to specific information processing. As neuronal ensembles are uncoupled from synchronized firing into their own (diverse) oscillatory frequencies, large, slower waveforms are replaced by faster frequencies of lower amplitude.
In neuropsychological terms, intrinsic (alpha) rhythms become partially or wholly desynchronized when sensory information is anticipated, attended to, or otherwise processed (Pfurtscheller, 1986; Pfurtscheller, 1989). Uncommitted cortical areas can remain in an "idling" state while other areas are desynchronized (Lopes da Silva, 1991; Pfurtscheller, 1992). Desynchronization can be localized to a single electrode or involve several electrodes and cortical areas (Pfurtscheller & Klimesch, 1990; Sterman et al., 1994). Regional patterns of simultaneous desynchronization and synchronization may characterize specific cognitive or behavioral states such as sensorimotor performance (Pfurtscheller & Klimesch, 1990; Pfurtscheller, 1992). EEG may prove useful in identifying processing strategies employed by different subjects (Tucker, 1976; Dunn & Reddix, 1991).

Topographic specificity of alpha attenuation

Alpha activity is sensitive to stimulus properties and demonstrates topographic specificity associated with modality. Visual stimulation activates occipital and parietal cortex and acoustic stimulation activates temporal cortex (Grillon & Buchsbaum, 1986; Pfurtscheller, Maresh, & Schuy, 1977). Similarly, sensory and motor event-related potentials (ERPs) are largest over corresponding primary and association areas (Gevins, 1986). Stimulus intensity, complexity, familiarity, and meaningfulness can influence alpha activity, presumably due to attentional factors in response to each of these properties (Gale & Edwards, 1983; Berlyne & McDonnell, 1965; Baker & Franken, 1967; Remond & Lairy, 1972; Boiten, Sergeant, & Geuze, 1992). The degree of alpha attenuation also shows habituation, diminishing after repeated presentations of the same stimulus (Gevins & Schaffer, 1980).

Baseline conditions

The most reliable and well-known finding in EEG research occurs when an individual resting with eyes closed opens his or her eyes. When eyes are opened in a lit room, alpha blocking occurs: the dominant alpha rhythm is displaced by faster, lower amplitude waveforms. This attenuation in alpha activity is widespread, incorporating most or all cortical areas -- a good example of nonspecific arousal (Legewie, Simonova, & Creutzfeldt, 1969; Etevenon, 1986; Etevenon et al., 1990; Pockberger, Petsche, Rappelsberger, Zidek, & Zapotoczky, 1985; Sterman et al., 1994).
Statistical results of topographic EEG lend themselves to two general physiological characterizations. A significant main effect occurs when one condition elicits a global change in attention or resource allocation that is not localized to any brain region (i.e., nonspecific arousal). Task by site interactions indicate when specific cortical functions are activated by one task but not others -- in other words, selective attention.

Rationale for additional spectral parameters

Gross physiological rhythms such as ultradian and circadian rhythms are well-known (e.g., Lavie, 1989) and may affect EEG activity (Meneses-Ortega & Corsi-Cabrera, 1990). Systematic fluctuation of shorter duration is also present in the EEG signal. Consistent activation without brief alpha bursts or similar disengagements is not seen for even brief periods of time. Alpha periodicity was noted as early as Berger (1930) who believed the phenomenon reflected attentional modulation. Tasks can temporarily attenuate the alpha periodicity associated with the "idling" or neutral state, but constant fluctuations are never extinguished completely by even very high or very low task demands. Periodicity in alpha activity is acknowledged by most EEG scientists, yet nearly all studies ignore trend dynamics and average across fluctuation. A significant fraction of information within the EEG signal is thus lost. When an individual becomes engaged in stimulus or task processing, the natural wax and wane of attention is inhibited and the degree of inhibition is reflected in alpha variability. Standard deviation of epoch magnitude and residual variance of a linear regression are useful measures of this task-induced variability.

Spectral parameters analyzed

The following four spectral parameters were analyzed in this study: log mean magnitude (LMAGN), standard deviation of epoch magnitude (SD), coefficient of a linear regression (SLOPE), and residual variance of a linear regression (RV). LMAGN is calculated by performing a natural logarithmic transformation of the average epoch magnitude (Gasser et al., 1982). SD, SLOPE, and RV assess temporal dynamics of alpha activity. SD reflects mean (global) variability associated with a task. RV is the error term of the linear regression and reflects variability not accounted for by a linear trend. SLOPE reflects the best linear trend of magnitude values across a number of seconds or minutes.
SLOPE values depend on the amplitude level at condition onset. For this reason, no demands were made of the subject for a short interval (approximately 10 s) prior to each condition. This "dead space" was used to reinstate a neutral attentional state, but without allowing too much time to pass so that a subject would become bored and restless. SLOPE values are also very sensitive to small changes in task demands, making it most useful for assessing brief intervals (e.g., 10 s) and least useful for assessing long duration tasks (e.g., 5 min). Conditions in the present study fall in between these extremes, so that SLOPE is mostly valuable when alpha trends are consistent across a task (i.e., RV is small).
LMAGN and SD are measured in microvolts (æV) and SLOPE and RV are measured in mean microvolt difference per epoch (ë æV per epoch). RV is always positive and is small when a strong linear relationship exists between amplitude and time. When the linear relationship is poor, RV is larger. Cortical engagement is associated with low amplitude, low variability, negative trend, and low trend instability.
LMAGN, SD, and RV reflect similar processes of activation and thus act together as a test of reliability. However, it must be kept in mind that SD can be influenced by SLOPE values and may not replicate results found in LMAGN or RV. On the other hand, SLOPE values do not reflect activation differences between tasks per se, but whether the attentional state varies within a task. Difficult tasks and high interest stimuli will usually result in more attentiveness with time (i.e., negative SLOPE values), but not always. In general, negative trends indicate task or stimulus habituation, or withdrawal from a task, and positive trends indicate concentration on task.

Summary of predictions

Three sets of eyes closed (EC) and eyes open (EO) baseline conditions were acquired to evaluate the four spectral parameters proposed above. No differences between replications are predicted for any spectral parameter or either condition. Opening one's eyes will result in nonspecific activation but no specific (topographic) attenuation in alpha activity compared to the EC condition (Sterman et al., 1994). As a more activated state, EO will show nonspecific reductions in amplitude, variability, and trend instability compared to EC. EC will consist of a stable state of relaxation and non- attentiveness whereas a prolonged eyes open condition without attentional demands will result in deteriorating subject attentiveness and vigilance (Sterman et al., 1994). Consequently, the slope for EO will be positive and higher than the slope for EC (approximately zero).

METHOD

Subjects

EEG was acquired from 27 healthy right-handed subjects. All subjects had mean alpha amplitude in EO conditions 10% to 50% below EC values. Seven subjects were excluded from the study due to excessive artifact or failure to meet the above criterion. EEG values were analyzed for 20 subjects (10 male, 10 female) between 21 and 40 years of age (mean age 29.45 y) while they viewed cinematic narratives and during baseline and control conditions. Subjects reported moderate to strong right-handedness as assessed by a modified Edinburgh Handedness Inventory (Appendix B), preferring the right hand or both hands equally on seven manual activities, with the right hand preferred on five or more manual activities (Oldfield, 1971). Hand dominance was also evident in writing samples from each hand. Subjects reported no history of neurological problems, use of a controlled substance 24 hr prior to the study, nor recent experiences which could alter one's normal psychophysiology (e.g., disturbed sleeping habits, atypical stress). Subjects were recruited through advertisements posted at the University of California at Los Angeles (UCLA) and the California State University, Northridge (CSUN) campuses. Informed consent was obtained from each subject prior to his or her participation, using a form approved by the Veterans' Administration Human Study Committee. Subjects were presented with a topographic image of their brain for participating in the study.

EEG Acquisition

A Neurosearch-24 acquisition unit (Lexicor Medical Technologies, Inc.) was used with an IBM-AT 486 compatible personal computer running Lexicor version 1.51 EEG acquisition software. Using a 12 bit A/D converter, EEG data were digitized at 512 samples per s during acquisition and was recorded and displayed 128 times per s, which resulted in 2 s epochs with 0.50 Hz frequency resolution. Lexicor software also performed an FFT of raw data for each epoch. High and low pass filters were set at 2 Hz and 16 Hz with rolloffs of 12 and 48 Db/octave, respectively. The common mode rejection ratio was greater than 90 dB at 60 Hz, notch filtering was applied at 60 Hz, and gain was set at 32,000. Topographic EEG was recorded from the scalp using an elastic lycra cap (Electro-Cap International Inc.) which consisted of 20 electrode ports containing pure tin electrodes arranged according to the international 10-20 placement system.

Preparation

After completion of a handedness questionnaire and film- viewing survey (see Chapter 4), subjects were fitted with an appropriate-sized Electro-Cap. Preparation of recording sites involved cleaning the scalp at each site with gentle scraping of a dowel (uncottoned Q-tip). Electrolytic gel was injected into each electrode port using a syringe with a blunted needle. Reference electrodes were attached to each ear lobe. All electrode impedances were below 7 K. All preparation instruments which came in direct physical contact with the subject were discarded after use.

Procedure

EEG was acquired at the Sepulveda Veterans Administration Medical Center Neuropsychology Laboratory. Subjects were tested individually in a dimly lit, electrically- shielded and sound-attenuated room approximately 2.5 by 3.5 m in size. Subjects were informed that subjective and psychophysiological reactions to film previews were being studied. Subjects were seated in a padded lounge chair which they could position for maximum comfort. A 21" screen television set was situated approximately 2 m from the subject at eye level. A small lamp controlled by the experimenter was used to illuminate the room whenever light was required. Data were acquired and stored to a hard disk situated in an adjacent room. Subject were monitored via close circuit video and two-way intercom systems during the study and EEG signals were monitored in real-time on a computer screen. All recording sessions began between 12 pm and 4 pm.

Baseline procedures

EC and EO baseline conditions of approximately 2 min were recorded three times during the study, spaced approximately 50 minutes apart (mean intervals of 51.1, 53.0 min, respectively). Between recordings, subjects watched short films (previews of coming attractions) and answered question about what they watched. Continuous topographic EEG was recorded from 19 sites (Appendix A).

Data Analysis

All EEG records were visually inspected on a computer monitor for artifacts. Any 2 s epoch contaminated by artifact was discarded prior to analysis. If artifact was present at any recording site, data from all 19 recording sites were rejected for an epoch.
For each condition, 50-64 epochs (100-128 s) were analyzed depending on the number of contaminated epochs. No record was used which had less than 80 percent of its epochs artifact-free. Each epoch underwent cosine tapering using a Blackman-Harris 4-term function which effectively reduces the frequency contributions from one-sixth of either end of the epoch.
EEG analysis focused on the 8-12 Hz frequency band. Four spectral parameters were calculated for each condition: amplitude (LMAGN), variability (SD), trend (SLOPE), and trend instability (RV).

RESULTS

Univariate Analyses of Variance (ANOVA) were used to assess topographic differences between replications and between baseline conditions. For initial comparisons, site values were pooled into six regions: frontal pole (Fp1, Fp2), frontal (F7, F3, Fz, F4, F8), central (C3, Cz, C4), temporal (T3, T4, T5, T6), parietal (P3, Pz, P4) and occipital (O1, O2) regions (cf. Klimesch et al., 1990, for similar treatment). Data were pooled to minimize computational complexity. The Huynh-Feldt correction of degrees of freedom were used to compensate for data sphericity. Individual site comparisons within an area were performed only if topographic interactions were significant. Functional laterality was assessed by comparing differences between homologous recording sites from each hemisphere (Fp1-Fp2, F7-F8, F3-F4, T3-T4, C3-C4, T5-T6, P3-P4, O1-O2).

Replication differences

Main effects were found between replications in both conditions for all spectral parameters. A topographic interaction between EC replications was found in SLOPE. No other topographic differences were found.
The first eyes closed replication (EC1) was lower in LMAGN, SD, and RV, compared to the second eyes closed replication (EC2) [FLMAGN(1,19)= 12.211, p<.01; FSD(1,19)= 11.097, p<.01; FRV(1,19)= 9.589, p<.01]. Similar differences were found between EC1 and the third eyes closed replication (EC3) [FLMAGN(1,19)= 13.011, p<.01; FSD(1,19)= 11.680, p<.01; FRV(1,19)= 13.249, p<.01]. SLOPE was higher in EC1 than in either EC2 and EC3 [FSLOPE(1,19)= 10.045, p<.01; F(1,19)= 15.145, p<.01, respectively]. No significant main effect was found between EC2 and EC3.
EO1 exhibited lower values in all parameters compared to EO3 [FLMAGN(1,19)= 10.162, p<.01; FSD(1,19)= 11.247, p<.01; FSLOPE(1,19)= 7.937, p<.01; FRV(1,19)= 11.022, p<.01]. Similar differences were observed between EO2 and EO3, though SLOPE did not differ [FLMAGN(1,19)= 20.517, p<.01; FSD(1,19)= 10.840, p<.01; FSLOPE(1,19)= 0.783, ns; FRV(1,19)= 13.630, p<.01]. No differences were found between EO1 and EO2 (see Table 3.1).

Table 3.1. Mean spectral values for replicated baseline conditions (19 sites, n=20)

        LMAGN      SD      SLOPE     RV
--------------------------------------------------------  
EC1     1.91*     3.12*     0.014*  0.021*
EC2     2.03      3.51     -0.025   0.023
EC3     2.06      3.56     -0.024   0.025
--------------------------------------------------------  
EO1     1.36      1.70      0.004   0.012 
EO2     1.38      1.89      0.012   0.013 
EO3     1.50*     2.17*     0.016+  0.016*
--------------------------------------------------------  
*     p<.01, all other replications
+     p<.01, first replication only

Topographic differences in EC replications were found in SLOPE [F(2,34)= 8.381, p<.001]. SLOPE values for six out of seven posterior sites (all except Pz) were higher in EC1 than EC2 [p<.01]. Fifteen of 19 sites (all except Fp1, Fp2, F7, and T4) had higher slopes in EC1 compared to EC3 [p<.01]. No slope differences were found between EC2 and EC3 [p>.01].
Task-induced laterality did not differ between replications in any parameter for EC or EO conditions [p>.05].

Baseline differences

Replication data were averaged for each spectral parameter and baseline condition (see Table 3.2). Main effects and topographic interactions were found between EC and EO.

Table 3.2. Mean spectral values for baseline conditions, averaged across replications (19 sites, n=20)

       LMAGN      SD       SLOPE     RV 
--------------------------------------------------------  
EC      2.00*     3.40*    -0.012*  0.023*
EO      1.41      1.92      0.011   0.014 
--------------------------------------------------------    
     * p<.01

Compared to EC, EO exhibited lower LMAGN, SD, and RV [FLMAGN(1,19)= 111.550, p<.01; FSD(1,19)= 103.384, p<.01; FRV(1,19)= 78.680, p<.01]. EC SLOPE was negative and smaller than EO SLOPE which was positive [FSLOPE(1,19)= 7.410, p<.01]. No laterality differences were found between conditions for any parameter [p>.01].
Topographic differences between EC and EO involved multiple sites from disparate functional areas, essentially duplicating main effects. Recording site by condition interactions were found in LMAGN, SD, and RV, and consisted of significant lower values at all 19 sites for EO [p<.01].

DISCUSSION

Subjects were more aroused during the first eyes closed replication and less aroused during last eyes open replication. The eyes open condition was also more aroused than the eyes closed condition.

Eyes closed replications

Contrary to predictions, subjects in the first eyes closed baseline were more aroused than during later conditions, probably due to residual attentiveness and low level anxiety displayed by the subjects. Subjects may not have been acclimated to the experimental setting and procedure during the first eyes closed condition (Rebert & Mahoney, 1978). Many subjects demonstrated low level anxiety over the first recording period by asking questions abruptly at any time during the condition. The lack of functional differences between replications parallel findings reported by Sterman et al. (1994) and Suyenobu (1994). Etevenon et al. (1989) and Etevenon et al. (1990) reported functional differences between eyes closed replications, but these results may not be reliable as neither study corrected for spatial dependence (Vasey & Thayer, 1987).
Eyes closed conditions appeared to represent two related psychological states: onset of relaxation, characterized by moderate activation that declines consistently over time (EC1); and thorough relaxation, a highly deactivated state characterized by fluctuations and intermittent increasing activation (EC2 and EC3; see Figure 3.1). However, no behavioral measures were analyzed to confirm these characterizations.


Figure 3.1. Trend of mean magnitude values for three EC replications (n=20).

Extreme deactivation was not maintained during any EC resting condition. In fact, magnitude values for each replication converged across time. Unlike task conditions, attentional and cognitive processes during eyes closed baselines are elicited by the individual's internal state and are not contingent upon external stimulation. The psychological state associated with intermediate magnitude may be self-reinforcing and most relaxing. Measures of variability in eyes closed baselines also demonstrated a similar convergence to intermediate values.

Eyes open replications

Nonspecific differences were found between eyes open replications. All replications exhibited a positive trend, but the rate of deactivation for the third eyes open condition was especially extreme. An abrupt shift in subject's mind set or attentiveness seemed to occur during the third replication. Most subjects may have assumed (correctly) that no other demands were to be made upon them and they relaxed completely. An onset of fatigue cannot explain this finding in that the third eyes closed condition, recorded immediately after, resembled earlier conditions. In contrast to eyes closed conditions, eyes open replications diverged across time and a drop in motivation likely occurred in the final replication.
Suyenobu (1994) also found significantly less activation in later EO replications compared to earlier conditions. However, Dolce & Waldeier (1974) and Sterman et al. (1994) reported no differences between three sets of eyes open baselines in alpha activity, though Dolce & Waldeier (1974) did note differences in other spectral bands. Fernandez, Harmony, Rodriguez, Reyes, Marosi, & Bernal (1993) found topographic differences between eyes open replications (limited to four sites) which they attributed to anxiety during the first recording session. No topographic differences were found between eyes open replications in the present study.
As shown in Figure 3.2, eyes open replications represented two mental states: nominal attentiveness, characterized by moderate activation that declines regularly over time (EO1, EO2); and inattentiveness, characterized by modest activation that deteriorates rapidly and unevenly over time (EO3). Again, these characterizations are not confirmed.


Figure 3.2. Trends of magnitude values for three EO replications (n=20).

Comparison between baseline conditions

Replications were averaged to derive condition means. Although an argument can be made for averaging only like conditions (e.g., EC2 and EC3 only), increased variance was tolerated in light of expected substantial differences between eyes closed and eyes open conditions (Adrian & Matthews, 1934; Gevins & Schaffer, 1980; Sterman et al., 1994), and because arousal heterogeneity is suspected in most studies which employ similar baseline conditions (Gevins, 1984; Freeman & Maurer, 1989).
As expected, opening the eyes resulted in nonspecific arousal, a finding consistent with the literature (e.g., Legewie et al., 1969; Etevenon, 1986; Sterman et al., 1994). Evidence of nonspecific arousal was also reflected in reduced variability and trend instability. As shown in Figure 3.3, the eyes open baseline is comprised of a moderate arousal which deteriorates steadily over time and the eyes closed baseline is an very low arousal state that settles to a less extreme state of arousal. This is accomplished by severe and erratic fluctuations of attention.


Figure 3.3. Trends of magnitude values for replicated baseline conditions (n=20).

The differences between baselines is found not only in amplitude, but all spectral parameters. It is unfortunate that few EEG scientists analyze these additional parameters. Measures of variance, normality, and other indices of dispersion and asymmetry have proven reliable and meaningful in other disciplines (Cacioppo & Dorfman, 1987).

Cerebral asymmetries did not differ between baselines conditions for any spectral parameter. Butler & Glass (1974) also found no evidence of functional asymmetries in alpha power between baseline conditions. However, it should be noted that individual differences in functional laterality during baseline conditions have been reported (Tomarken, Davidson, & Henriques, 1990).

"Macrostate" assumption

An assumption underlying this experiment is the conception of psychological or physiological "macrostates". According to this model, the various perceptual and cognitive operations associated with a mental or behavioral condition is thought to constitute a single distinguishable neurophysiological state with a distinct and reliable spectral pattern (Gevins, 1984; Gevins, 1986). Eyes closed and eyes open resting conditions are very good tests of this model. Both conditions are uncontrolled, self-paced, and may include dissimilar processes between subjects. Assuming that EEG is sensitive enough to measure functional differences between conditions, the fact that replications of these conditions did not differ topographically for four spectral parameters is evidence which supports the macrostate assumption.

References

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