Please cite this article in press as: Bonnefond and Jensen, Alpha Oscillations Serve to Protect Working Memory
Maintenance against
Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
Current Biology 22, 1–6, October 23, 2012 ª2012 Elsevier Ltd All rights reserved
Alpha Oscillations Serve to Protect
Working Memory Maintenance
against Anticipated Distracters
Mathilde Bonnefond1,* and Ole Jensen1,*
1Radboud University Nijmegen, Donders Institute for Brain,
Cognition and Behaviour, Centre for Cognitive Neuroimaging,
Kapittelweg 29, 6525 EN Nijmegen, The Netherlands
Summary
When operating in a complex world, it is essential to have
mechanisms that can suppress distracting information
[1, 2]. Such mechanisms might be related to neuronal
oscillations, which are known to be involved in gating of
incoming information [3]. We here apply a working memory
(WM) task to investigate how neuronal oscillations are
involved in the suppression of distracting information that
can be predicted in time. We used a modified Sternberg
WM task in which distracters were presented in the retention
interval, while we recorded the ongoing brain activity
using magnetoencephalography. The data revealed a robust
adjustment of the phase of alpha oscillations in anticipation
of the distracter. In trials with strong phase adjustment,
response times to the memory probe were reduced. Further,
the power of alpha oscillations increased prior to the dis-
tracter and predicted performance. Our findings demon-
strate that the doors of perception close when a distracter
is expected. The phase adjustment of the alpha rhythm
adds to the computational versatility of brain oscillations,
because such a mechanism allows for modulating neuronal
processing on a fine temporal scale.
Results
To investigate the role of oscillatory activity in distracter
suppression in working memory (WM) tasks, we designed
a modified version of the Sternberg paradigm. A memory set
of four consonants was presented to the subject followed by
a probe. In the retention interval, we presented either weak
(a symbol) or strong (a letter) distracters in different blocks
such that the subjects could anticipate the nature of the dis-
tracter (Figure 1A). To optimize the chances for identifying
a phase effect in the 10 Hz band, we flashed the distracters
for 33 ms. Importantly, the onset of the distracter at t = 1.1 s
was predictable across trials. The ongoing brain activity was
recorded using magnetoencephalography (MEG) from 18
subjects performing this task.
Reaction Times Longer for Strong than Weak Distracters
Trials
It is well established that the reaction time to the probe
increases with memory load in the Sternberg task [4]. Thus
the potential incorporation of a distracter in WM should be
reflected in longer reaction times. A two-way repeated-
measures ANOVA analysis with the factors ‘‘distracter’’
(weak versus strong) and ‘‘probe type’’ (old versus new)
*Correspondence: mathilde.bonnefond@donders.ru.nl (M.B.), ole.jensen@
donders.ru.nl (O.J.)
http://dx.doi.org/10.1016/j.cub.2012.08.029
Report
revealed that the reaction times for strong distracters
(725 6 35 ms) was significantly longer than for weak dis-
tracters (699 6 31 ms; p < 0.05; Figure 1B). The findings
suggest that strong distracters are incorporated into WM in
some trials.
Occipitotemporal Alpha Power Increases Prior to the
Anticipation of Strong Distracters
To characterize the oscillatory activity reflecting the antici-
pation of distracters, we quantified the properties of the
MEG data in the interval before the strong versus the weak
distracters (0 < t < 1.1 s). The time-frequency representation
(TFR) of power revealed larger 8–12 Hz alpha activity over
posterior sensors prior to the distracter onset (Figures 2A
and 2B). A cluster-based randomization test (see Supple-
mental Information available online) showed that the effect
was significant in the alpha band in the 0.6–1.1 s interval
over left-lateralized parieto-occipital sensors (p < 0.05;
Figure 2B). The sources reflecting the alpha power increase
were located in the occipital cortex extending into the left
temporal cortex (Figure 2C).
An Increase in Alpha Activity Protects against Anticipated
Distracters
We set out to determine whether alpha power is associated
with distracter suppression as assessed by the reaction
times to the probe. We computed the average alpha
power (8 Hz < f < 12 Hz; 0.6 < t < 1.1 s) for the cluster of sensors
shown in Figure 2B for fast and slow reaction times. This was
done by sorting the trials according to reaction times in five
bins. Trials from the second and fourth bin were considered
fast and slow in order to ignore trials with either too long or
too short reaction times because they are likely to be domi-
nated by premature button presses or lapses in attention
(see also Supplemental Information). A two-way repeated-
measures ANOVA with factors ‘‘distracter type’’ and ‘‘reaction
time’’ revealed a significant interaction between reaction time
and distracter type (p < 0.05). A post hoc tests showed that
alpha power was significantly higher for fast than for slow
reaction times trials in the strong distracter condition (p <
0.05; Figure 2D). These results suggest that stronger alpha
activity in anticipation of a strong distracter leads to a better
distracter suppression.
Alpha Power Predicts Individual Differences in
Performance
We performed an analysis comparing individual abilities to
suppress distracters in relation to the ability to adjust the alpha
activity prior to a distracter. Over subjects we correlated the
difference in reaction time and the difference in alpha power
(data derived from left occipital-temporal cortex using a beam-
former spatial filter; see Supplemental Information) between
weak and strong distracters. We observed a negative correla-
tion: the stronger the difference in power between anticipated
distracter types, the smaller the difference in reaction times
(r = 20.56; p = 0.02; Figure 2E). These data suggest that indi-
viduals with a better ability to modulate their alpha activity
are less impaired by strong distracters.
 Please cite this article in press as: Bonnefond and Jensen, Alpha Oscillations Serve to Protect Working Memory Maintenance against
Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
Current Biology Vol 22 No 20
2
Figure 1. The Modified Sternberg Paradigm
(A) A set of four consonants was presented sequentially at a rate of 1,133 ms
per letter. As in the conventional Sternberg paradigm, subjects indicated
by button press whether the probe was part of the memory set (‘‘old’’ or
‘‘new’’). In the retention interval, either a ‘‘weak’’ (a symbol) or ‘‘strong’’ (a
consonant) distracter was presented. The distracter strength could be
anticipated by the subjects because weak and strong distracters were
grouped in blocks of trials.
(B) The hit rate and reactions times in response to the memory probe (error
bars represent the SEM). Although the hit rate was independent of distracter
type, the response times were significantly longer for the strong compared
to weak distracters. This indicates that strong distracters are encoded in
working memory in some trials and interferes with retrieval.
Alpha Oscillations Align in Phase Prior to Anticipated
Strong Distracters
Anticipatory phase adjustment could provide a mechanism for
gain regulation in association with temporal expectation [3].
We aimed to determine whether alpha oscillations can adjust
in phase to optimally suppress the distracter. To this end, we
computed the phase-locking factor (PLF) over trials for each
condition. The TFR of the PLF (thresholded at 0.087; see
Supplemental Information) over the cluster of sensors shown
Figure 2B revealed phase locking in the alpha band prior to
the distracter onset (Figure 3A). The topography of the PLF
(8 Hz < f < 12 Hz; 0.6 < t < 1.1 s) considering all sensors revealed
a left-lateralized distribution that matched the power differ-
ence between conditions (cf. Figures 2B and Figures 3B).
Further analyses revealed that the phase effects are not
trivially explained by systematic variations in power (see
Supplemental Information). Next, we considered the differ-
ence in PLF for strong versus weak distracters (Figure 3C)
(the PLF was transformed to Rayleigh Z values to control for
biases due to differences in trial numbers; see Supplemental
Information). The difference in phase locking increased in
anticipation of the strong distracter (Figure 3C). This excludes
Figure 2. The Power of Alpha Band Activity Prior to the Distracter Onset
(A) Time-frequency representation of power when comparing strong and
weak distracters. The alpha power was higher for strong compared to
weak distracters. The arrow indicates the onset of the distracter.
(B) The topography of the alpha power (8–12 Hz; 0.6–1.1 s; combined planar
gradient) when comparing strong to weak distracters. The cluster random-
ization analysis revealed a significant cluster (marked by asterisks; p < 0.05)
in posterior sensors.
(C) The source analysis of the alpha activity (8–12 Hz; 0.6–1.1 s) for strong
compared to weak distracters revealed the strongest contributions from
left occipitotemporal cortices (MNI coordinates: [252 262 26]; t values
are masked corresponding to a p value of 0.05). To further support
this result, we also compared predistracter period (0.6 s < t < 1.1 s) to prep-
robe period (1.833 < t < 2.233 s). We found overlapping sources is occipito-
parietal cortex.
(D) The difference in alpha power (0.6 < t < 1.1s; 8 < f < 12 Hz) when
comparing trials with fast and slow reaction times for, respectively, weak
and strong distracters (error bars represent the SEM). The log-ratio of alpha
power was significantly larger for strong compared to weak distracters.
(E) Differences in alpha power (strong compared to weak distracters) corre-
lated negatively with differences in reaction times (strong compared to weak
distracters) over subjects, i.e., subjects in which the alpha power increased
in anticipation of a strong distracter were also subjects that were less
slowed down during memory probing.
that the phase locking was explained by sustained alpha
oscillations elicited by the last memory item. The difference
was significant when averaged over the 0.6–1.1 s interval
(p < 0.01; Figure 3D). These findings demonstrate that alpha
phase is adjusted in anticipation of a predictable strong
distracter.
To further substantiate our findings, we investigated the
phase adjustment using a spatial filter aimed at extracting
the activity from the occipital-temporal source. Using a linearly
 Please cite this article in press as: Bonnefond and Jensen, Alpha Oscillations Serve to Protect Working Memory Maintenance against
Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
Alpha Oscillations Protect Working Memory
3
Figure 3. Phase Adjustment of Alpha Activity Prior to Distracter Onset
(A) The time-frequency representation of the phase-locking factor (PLF) in
the retention interval over sensors of interest (see Figure 2B for sensors
selection; PLF thresholded at 0.087 corresponding to p < 0.01 in the Ray-
leigh test) reveals that alpha oscillations are adjusted in phase prior to the
onset of the distracter. The strong PLF values observed in the theta band
(5–8 Hz) a t = w0.4 s and t = w1.1 s reflect activity evoked by, respectively,
the last memory item and the distracter. Note that the acausal temporal
smoothing associated with the phase estimation was approximately three
cycles long, i.e., 300 ms for 10 Hz and 500 ms for 6 Hz. The arrow indicates
the onset of the distracter. We further observed a significant PLF in 14 of 18
subjects (0.6 < t < 1.1 s; 8 < f < 12 Hz; Rayleigh test: p < 0.01).
(B) The topography of the PLF (0.6 < t < 1.1 s; 8 < f < 12 Hz) reveals the stron-
gest values in posterior sensors.
(C) Time-frequency representation of the difference in PLF (transformed to
Rayleigh Z values where Z = number of trials 3 PLF2) comparing strong
and weak distracters (for sensors marked by asterisks in Figure 2B). The
difference is most pronounced in the alpha band just prior to the onset of
the distracter; i.e., the phase adjustment is not explained by sustained oscil-
lations elicited by the last memory item.
(D) The PLF is significantly higher in the strong than in the weak distracter
condition over the sensors of interest (p< 0.01; error bars represent
the SEM).
(E) The time-locked responses (event-related fields) prior to the distracter.
The single trials from the left occipital-temporal source were derived using
a beam forming approach. The time-locked responses were averaged over
subjects. A clear modulation in the alpha band is seen prior to the distracter
(as indicated by small green arrows). a.u., arbitrary units.
See also Figure S1.
constrained minimum variance beamformer [5], we obtained
the time course for each trial from seven grid points in the
left occipital-temporal cortex (see Figure 2C). The phase
adjustment should as well be observable in the trial average
just prior to distracter onset. The grand average over trials
and subjects is shown in Figure 3E (baseline: 20.233 < t <
20.033 s, low-pass filtered at 30 Hz). The phase-adjusted
Figure 4. Assessing Difference in Predistracter Phase for Slow and Fast
Reaction Times
(A) The difference in phase between fast and slow trials was quantified using
a phase-difference index (error bars represent the SEM). We found a signif-
icant phase difference index for the strong but not the weak distracter trials.
(B) The time-locked responses (event-related fields) prior to the distracter
according to trials with fast (in red) and slow (in blue) reaction times at
source level. A clear lag can be observed for three cycles of alpha
activity prior to the distracter (as indicated by small green arrows). a.u. =
arbitrary units.
See also Figure S2.
alpha activity prior to the distracter was clearly apparent in
this averaged trace (marked by green arrows). Note that the
phase-adjusted alpha activity occurred just prior to the dis-
tracter and cannot be explained by sustained oscillations eli-
cited by the last memory item at t = 0 s. We also found
evidence for frontal sources in which the phase adjustment
of alpha activity was more pronounced just prior to strong
compared to weak distracters (see Supplemental Information).
In future research, it would be of interest to investigate whether
these frontal regions are involved in exercising a top-down
drive that adjusts the alpha activity in posterior regions ([6,
7]; see Supplemental Information; Figure S1A).
Alpha Phase Predicts the Efficiency of the Suppression
of the Processing of the Distracter
We further tested whether the phase-locking values for slow
and fast trials were associated with a difference in absolute
alpha phases. This was done using a phase-difference index
[8, 9]. The phase difference index in Figure 4A shows the differ-
ence in PLFs when comparing true versus randomized trials
for fast versus slow reaction times trials. For the strong
distracter, the significant phase difference index suggests
that fast and slow distracter trials are associated with different
phases of the alpha activity at the onset of the distracter.
Figure 4B shows the averaged traces for fast and slow reaction
times trials for data from the occipitotemporal source in the
strong distracter condition. This analysis confirms that fast
and slow reaction time trials are associated with different
alpha phases. Our key interpretation is that if the distracter
arrives at a certain alpha phase, it will be more intrusive
Please cite this article in press as: Bonnefond and Jensen, Alpha Oscillations Serve to Protect Working Memory Maintenance against
Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
Current Biology Vol 22 No 20
4
as compared to the reverse phase. Further, the results
demonstrate that the anticipatory phase adjustment has
functional implications, and that the phase adjustment is not
trivially a consequence of the power increase prior to the
distracter.
Also note the slow modulation of the ERFs in Figure 3E. This
modulation is reflected in the delta band but is similar for high
and low distracter trials (see Figure S2). This could suggest
that the phase of the delta rhythm is involved in controlling
the temporal modulation, but not magnitude, of the alpha
oscillations. We conclude that anticipatory phase adjustments
mainly in the alpha band serve to protect WM against
distracters.
Unpredictable Compared to Predictable Distracter Onsets
Result in Longer Reaction Times
If the temporal predictability of the distracter allows for
suppression by a phase-adjustment of the alpha oscillations,
then a predistracter delay varying by w100 ms (matching the
duration of an alpha cycle) should reduce performance. In
a behavioral study conducted outside the MEG system (see
Supplemental Information), we compared fixed (1.1.s) to vari-
able delays (from 1.03 to 1.16 s; 33 ms steps) between the
last memory item and the distracter. The total duration of the
retention interval was kept identical for both conditions.
Fixed intervals resulted in significantly shorter reaction times
(664 6 30 ms versus 687 6 33 ms; 12 participants; p < 0.05).
These data suggest that when subjects can precisely antici-
pate the distracter they are better at suppressing it.
Discussion
The present experiment was designed to identify the mecha-
nism responsible for suppressing anticipated intruding infor-
mation. In a task where subjects could anticipate the strength
and exact timing of distracters, we observed both a stronger
power increase and phase adjustment of alpha activity in
occipitotemporal areas prior to strong compared to weak dis-
tracter onsets. Importantly, we demonstrated that trials with
fast and slow reaction times were associated with different
absolute predistracter alpha phases and a stronger alpha
power in the faster trials. These findings suggest that a failure
to adjust the alpha phase and power just prior to a distracter
will disturb WM processes.
These results underscore that alpha activity is responsible
for active functional inhibition of sensory regions [10, 11]. For
instance, previous studies have shown that when attention is
allocated to the right hemifield, the alpha activity decreases
over the left visual system while it increases relatively in the
right hemisphere. The same holds for the somatosensory
system [12–21]. These modulations in alpha activity are pre-
dictive of performance [15, 18]. Decreases in alpha activity
have also been shown to correlate with temporal expectations
[22, 23]. We here show for the first time that alpha is modulated
according to the strength of distracters and that alpha power is
linked to the suppression of distracting information (see also
[24]). The source of the alpha activity included the left fusiform
gyrus, an area known to be engaged in letter processing
[25–27]. Moreover, the activity in this area predicted individual
differences in reaction time between the strong and the weak
distracter conditions. This result indicates that distracter
suppression in particular relies on the inhibition of representa-
tional specific areas. Further it extends results from a previous
study showing that subjects with a stronger posterior alpha
increase during the retention period in the Sternberg task
also perform faster [28].
The finding that alpha phase adjusts prior to the distracter
is highly novel. Clearly, the alpha activity is rhythmic and
the phase of the oscillatory activity has been shown to
modulate processing [8, 29–33]. We now show that partici-
pants can adjust alpha phase when the timing of a distracter
is predictable. Such an adjustment of oscillatory activity has
been shown in attentional tasks in monkeys with respect to
delta oscillations [3]. In this study, the monkeys had to detect
a target either in the visual or the auditory domain both
presented in alternation at 1.5 Hz. The phase of the delta
activity (w1.5 Hz) was top-down adjusted in the visual cortex
for the attended visual stimulus (visual-attention condition)
to appear at a high-excitability phase and the unattended
visual stimulus (auditory-attention condition) to appear at
a low-excitability phase. Our findings extend this concept:
in the Lakatos et al. study [3], the frequency of the oscillations
was determined by the cadence of the task. We show that
an intrinsically generated cortical rhythm can adjust in
phase. Further, this alpha phase adjustment is consistent
with the behavioral experiment showing that distracters
predictable in time are associated with better performance
compared to distracters presented with a w100 ms jitter.
One might suspect that the phase adjustment just prior to
the distracter is explained by alpha activity evoked by the
last memory, which sustained in phase until the appearance
of the distracter. This concern is ruled out by the observation
that the behaviorally relevant alpha adjustment increases just
prior to the distracter onset (see Figures 3E and Figures 4B;
Figure S1B).
Our findings demonstrating alpha phase adjustment imply
that the human brain is able to predict events with a temporal
precision on the order of 20–30 ms or less. This is consistent
with various findings demonstrating that humans are able
to discriminate fine temporal durations [34–41]. We further
showed that alpha phase adjustment is predictive of fast and
slow reaction times to the probe. We propose that a trial in
which performance is poor is a result of an insufficient adjust-
ment of alpha phase possibly due to lapses in attention.
Although the duration of the distracter was 33 ms, we expect
that the results would be reproduced when using longer
lasting distracters because it is the stimulus onset that defines
the critical moment for when our attention is drawn.
From a mechanistic point of view, one might ask why an
adjustment of both alpha magnitude and phase is required
for optimal performance. It has been proposed that the alpha
rhythm is a consequence of pulses of inhibition that can vary
in magnitude [10, 11, 42, 43]. This notion has recently received
support from monkey recordings showing that alpha oscilla-
tions inhibit neuronal firing selectively at specific phases [44].
An adjustment in alpha magnitude only would provide a
suboptimal mechanism for functional inhibition, because
information appearing between the inhibitory pulses will not
be suppressed; thus, there is a need for alpha phase adjust-
ment as well. Our findings are consistent with the notion that
a key function of the central nervous system is to predict
upcoming events [45]. We here provide insight to the physio-
logical substrate of temporal predictions.
In this study, we demonstrated that alpha phase can be
adjusted in anticipation of an event in order to close the doors
of perception when intruding information might disturb
ongoing processes. This new principle adds considerable to
the versatility of the alpha rhythm in the context neuronal
Please cite this article in press as: Bonnefond and Jensen, Alpha Oscillations Serve to Protect Working Memory Maintenance against
Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
Alpha Oscillations Protect Working Memory
5
processing. The stage is now set for investigating how general
this principle is [46].
Supplemental Information
Supplemental Information includes two figures, Supplemental Results and
Discussion, and Supplemental Experimental Procedures and can be found
with this article online at http://dx.doi.org/10.1016/j.cub.2012.08.029.
Acknowledgments
The authors gratefully acknowledge The Netherlands Organization for
Scientific Research (NWO) VICI grant number 453-09-002, ‘‘The healthy
brain’’ funded by the Netherlands Initiative Brain and Cognition, a part of
the Organization for Scientific Research (NWO) under grant number
056-14-011 and the Fyssen funding scheme. We thank Paul Gaalman for
assistance with structural MRI collection and Sander Berends for assis-
tance with MEG recordings.
Received: January 31, 2012
Revised: July 21, 2012
Accepted: August 15, 2012
Published online: October 4, 2012
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Anticipated Distracters, Current Biology (2012), http://dx.doi.org/10.1016/j.cub.2012.08.029
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