Egbert Juergens & Frank Roesler (1995) How do Alpha Oscillations Influence Gamma Band Activity?. Psycoloquy: 6(09) Brain Rhythms (10)

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PSYCOLOQUY (ISSN 1055-0143) is sponsored by the American Psychological Association (APA).
Psycoloquy 6(09): How do Alpha Oscillations Influence Gamma Band Activity?

Commentary on Pulvermueller et al. on Brain-Rhythms

Egbert Juergens & Frank Roesler
Department of Psychology
Philipps-University Marburg
Gutenbergstrasse 18
D-35032 Marburg


Gamma-band activity, as examined by Pulvermueller et al. (1994) is not necessarily a genuine and independent phenomenon but may be an epiphenomenon of activity changes in lower frequency bands. Recent experiments have shown a strong contribution of harmonics of alpha activity to activity in the beta and gamma ranges. The target article and the response to the commentary of Mueller & Jokeit (1994) do not present enough evidence to invalidate the hypothesis that the observed gamma-band effects are merely due to changes in alpha power.


brain theory, cell assembly, cognition, event related potentials (ERP), electroencephalograph (EEG), gamma band, Hebb, language, lexical processing, magnetoencephalography (MEG), psychophysiology, periodicity, power spectral analysis, synchrony
1. Pulvermueller et al. (1994) reported findings from an experiment in which subjects had to discriminate between words and pseudowords. In the event-related potential of the EEG, pseudowords elicited a more negative amplitude around 400 ms post-stimulus-onset than legal words. In addition, a reduction of Gamma activity in the 25-35 Hz range was found from a pre- to a post-stimulus epoch over the left hemisphere when pseudowords had to be processed. No such effect could be observed for words. While the pronounced negativity associated with pseudowords is well known (Kutas & Kluender, 1994), the gamma-reduction is a new finding. However, it is not clear whether this gamma reduction is a genuine and independent phenomenon or only an epiphenomenon of some other much better known effect, that is, a change of alpha-activity.

2. Alpha activity is known to be closely linked to cognitive processing (Michel, Kaufman & Williamson, 1994; Pfurtscheller & Klimesch, 1991; Vijn, 1992). A reduction of spectral power in the lower alpha range (8-11 Hz) was found in many psychophysiological experiments after onset of a stimulus, during cognitive processing, and with increased attention. A stimulus related increase of power in the upper alpha band (11-13 Hz) was also reported (Gale, Davies & Smallbone, 1977; Roesler, 1975). Therefore, it is most likely that stimulus related changes of alpha activity were also present in the experiments described by Pulvermueller et al. (1994). Moreover, it has to be expected that alpha band effects were different for words and pseudowords, because processing demands, as reflected by the phasic slow negative wave, were higher for pseudowords than for legal words.

3. As Mueller and Jokeit (1994) already noted, Fourier analysis of nonsinusoidal periodic activity does not only produce one single peak corresponding to the oscillation period but also peaks at integer multiples of the base frequency. Power coefficients at these harmonics depend on the shape of the periodic process. A symmetric triangular waveform, for example, has harmonics at odd multiples of the base frequency while the power at even multiples is zero (Brigham, 1975).

4. A recent experiment of our group on memory storage and retrieval confirms that strong interrelations can exist between power in the alpha and in higher frequency ranges (Juergens, Roesler, Hennighausen & Heil, 1995). In the first part of this experiment, associations between line drawings and two types of mediators (grid positions and concrete nouns) had to be learned. In the retrieval experiment subjects should decide whether two drawings were associated via a common mediator or not (Heil, Roesler & Hennighausen, 1994; Roesler, Heil & Hennighausen, 1995). During both experimental sessions, the EEG was recorded between 0 and 100 Hz from seven electrodes. Fourier spectra were calculated in successive overlapping time windows of 1 s length. For one subject, a stimulus-locked increase of power at 12 Hz was found, while in the other two subjects alpha power decreased at the individual peak frequencies of 9 and 11 Hz, respectively. The spectra of all subjects revealed corresponding changes of power in the higher frequency ranges but only at harmonics of the individual dominant alpha frequencies. These observations could be substantiated by an analysis of covariance of the time courses at harmonic and non-harmonic frequencies in the alpha, beta and gamma ranges.

5. The target article does not discuss the possible influence of alpha activity on the reported findings in the gamma range. In their response to Mueller & Jokeit (1994), Pulvermueller & Lutzenberger (1994) reported that an analysis of variance did not reveal a significant Wordness-effect or interactions of Wordness with other factors in the 10-20 Hz range. However, this finding does not invalidate the hypothesis that the changes in the gamma-range are due to equivalent Wordness-effects in the alpha range, for two reasons: Firstly, the lower alpha band (8-9 Hz) was not examined at all; and secondly, the filter used to restrict the frequency range was cosinus shaped (Lutzenberger et al., 1994). This implies that filter coefficients at the edges of the passband (10, 11, 19 and 20 Hz) are small. Consequently, the respective frequencies could bear only very little influence on the power of the filtered signal. The exclusion of the lower alpha band may be the reason why possible differences in lower alpha activity between word/pseudoword conditions did not become manifest as significant statistical effects in the analysed frequency band (10-20 Hz). It should be noted that the influence of the first harmonics (16-22 Hz) of the lower alpha band might be non-significant because of the specific shape of the alpha oscillation. Power decreases at the first harmonics of lower alpha activity could also be masked by increases of higher alpha activity. The second harmonics of the lower alpha activity is located in the frequency range of 24-33 Hz overlapping the 25-35 Hz range where a significant Wordness effect was found. This might be the reason for the observed wordness effects in the gamma-band.

6. Further steps in data analysis are necessary for a decision whether effects in the gamma-band are genuine or just epiphenomena of alpha activity. In particular, it is necessary to evaluate how different sub- bands of the alpha range are related to the Wordness effect. In Juergens et al. (1995) it was found that stimulus related changes in the alpha-band are often restricted to very narrow frequency ranges with a bandwidth of one Hz only. Higher spectral resolution is a prerequisite for a more convincing analysis. Using narrow band filtering, frequency peaks at integer multiples of the dominant alpha frequency with high temporal covariance would reveal the existence of harmonics. Another problem is caused by the fact that alpha oscillations show substantial interindividual variation in frequency, depending on experimental manipulations and most likely also in the pattern of harmonics (Deakin & Exley, 1979; Gale, 1973). Therefore, alpha-gamma-interrelations have to be evaluated separately for each subject.

7. A further problem in Pulvermueller et al. (1994) concerns the fact that there were absolutely no differences in gamma-band activity from the pre- to the post-stimulus epoch when words had to be processed. Processing of words has a precise timing in relation to stimulus onset. Consequently the proposed assembly model would predict a difference in cell assembly activity between pre- and post-stimulus epoch. While in the prestimulus baseline "a continuous stream of thoughts" prevails, evoking search processes which are similar to the functional state of recognizing pseudowords, the presentation of a legal word should ignite a particular cell assembly only. This should become evident as a short increase of gamma-activity.


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