Current Biology 21, 1833–1837, November 8, 2011 ª2011 Elsevier Ltd All rights reserved DOI 10.1016/j.cub.2011.09.

029

Report
Dreamed Movement Elicits Activation
in the Sensorimotor Cortex

Martin Dresler,1,5 Stefan P. Koch,2,5 Renate Wehrle,1,5 case, dreamed hand clenching. Predecided eye movements
Victor I. Spoormaker,1 Florian Holsboer,1 Axel Steiger,1 served as temporal markers for the onset of hand clenching
Philipp G. Sämann,1 Hellmuth Obrig,2,3,4 and for hand switching. Previous studies have shown that
and Michael Czisch1,* muscle atonia prevents the overt execution of dreamed hand
1Max Planck Institute of Psychiatry, Kraepelinstrasse 2-10, movements, which are visible as minor muscle twitches at
80804 Munich, Germany most [3, 9].
2Berlin Neuroimaging Center, Charité University Hospital, Subjects were instructed to make series of left and right hand
Humboldt University of Berlin, 10099 Berlin, Germany movements separated by sets of left-right-left-right (LRLR)
3Max Planck Institute for Human Cognitive and Brain eye movements upon becoming lucid (see Experimental
Sciences, Stephanstrasse 1, 04103 Leipzig, Germany Procedures below; see also Supplemental Information avail-
4Clinic for Cognitive Neurology, University Hospital Leipzig, able online). Out of six highly experienced lucid dreamers
04103 Leipzig, Germany who participated in the study, two subjects succeeded in
performing the task during lucid REM sleep under combined
electroencephalography (EEG)-fMRI or combined EEG-NIRS
Summary conditions, respectively. As a control condition, the subjects
additionally performed both an actually executed and an imag-
Since the discovery of the close association between rapid ined hand-clenching task during wakefulness. Increases in
eye movement (REM) sleep and dreaming, much effort has blood oxygen level-dependent (BOLD) signal in the sensori-
been devoted to link physiological signatures of REM sleep motor cortex contralateral to the indicated movement side
to the contents of associated dreams [1–4]. Due to the im- were found in the successful fMRI experiments during the
possibility of experimentally controlling spontaneous dream two lucid REM states in one subject (Figure 2). Given the short
activity, however, a direct demonstration of dream contents duration of the lucid dreams, the activation of the sensorimotor
by neuroimaging methods is lacking. By combining brain cortical regions was very specific compared to wakefulness:
imaging with polysomnography and exploiting the state of executed hand clenching during wakefulness (WE) elicited
‘‘lucid dreaming,’’ we show here that a predefined motor widespread activation clusters in the contralateral pre- and
task performed during dreaming elicits neuronal activation postcentral gyrus (pFWE < 0.005). Collection threshold for
in the sensorimotor cortex. In lucid dreams, the subject is both imagined hand movements during wakefulness (WI) and
aware of the dreaming state and capable of performing pre- dreamed hand movements (LD) were set at puncorr < 0.005,
defined actions while all standard polysomnographic criteria with voxel extent 50. Cluster significance was assessed in
of REM sleep are fulfilled [5, 6]. Using eye signals as temporal a volume of interest defined by the activation during executed
markers, neural activity measured by functional magnetic hand movement in wakefulness. The strength of the activation
resonance imaging (fMRI) and near-infrared spectroscopy was evaluated by analyzing the difference between the maxi-
(NIRS) was related to dreamed hand movements during lucid mum and minimum BOLD signal amplitude in the peak voxel’s
REM sleep. Though preliminary, we provide first evidence time course of the respective motor regions. Strongest BOLD
that specific contents of REM-associated dreaming can be signal fluctuations of the peak voxel’s time course can be
visualized by neuroimaging. seen during WE (left hand: 5.62%; right hand: 4.49%), whereas
WI and LD showed mean fluctuations of only 1.3%–1.4% and
Results 1.8%–2.5%, respectively (Figure 2).
FMRI results were confirmed by an independent imaging
Lucid dreaming is a rare but robust state of sleep that can method in a second subject: NIRS data showed a typical
be trained [5]. Phenomenologically, it comprises features of hemodynamic response pattern of increased contralateral
both waking and dreaming [7]: in lucid dreams, the sleeping oxygenation over the sensorimotor region during successful
subject becomes aware of his or her dreaming state, has full task performance in lucid REM sleep (Figure 3; Figure 4).
access to memory, and is able to volitionally control dreamed Notably, during dreaming, the hemodynamic responses were
actions [6]. Although all standard polysomnographic criteria of smaller in the sensorimotor cortex but of similar amplitude in
rapid eye movement (REM) sleep [8] are maintained and REM the supplementary motor area (SMA) when compared to overt
sleep muscle atonia prevents overt motor behavior, lucid motor performance during wakefulness.
dreamers are able to communicate their state by predefined
volitional eye movements [6], clearly discernable in the electro- Discussion
oculogram (EOG) (Figure 1). Combining the techniques of lucid
dreaming, polysomnography, and brain imaging via functional The discovery of the close relationship between REM sleep
magnetic resonance imaging (fMRI) or near-infrared spectros- and dream reports [10, 11] boosted decades of neuroscientific
copy (NIRS), we demonstrate the possibility to investigate research into the brain state potentially evoking vivid and in-
the neural underpinnings of specific dream contents—in this tense dreams. Neurophysiological studies suggest that during
REM sleep, the brain functions as a closed loop system, in
which activation is triggered in pontine regions while sensory
5These authors contributed equally to this work input is gated by enhanced thalamic inhibition and motor
*Correspondence: [email protected] output is suppressed by atonia generated at the brain stem
Current Biology Vol 21 No 21
1834

in the SMA was not observed, the NIRS-measured hemody-

namic responses during dreaming were smaller in the sen-
sorimotor cortex but of similar amplitude in the SMA when
compared to overt motor performance during wakefulness.
The SMA is involved in timing, preparation, and monitoring of
movements [21], and linked to the retrieval of a learned motor
sequence especially in the absence of external cues [22]. The
SMA has been attributed as a programming area active only
during complex movements [23]; however, our NIRS data
Figure 1. Exemplary Lucid REM Sleep as Captured by Polysomnography speak for an activation of SMA even during simple movements
during Simultaneous fMRI in this subject. This is in line with several PET and fMRI studies
Note high-frequency electroencephalogram (EEG) and minimal electromyo- reporting SMA activations for simple tasks such as hand
gram (EMG) amplitude due to muscle atonia characteristic of rapid eye clenching, single finger-tapping, and alternated finger-tapping
movement (REM) sleep (left), with wakefulness for comparison (right). (see Supplemental References and Supplemental Discussion
Subjects were instructed to communicate the state of lucidity by quick of the NIRS and fMRI mismatch).
left-right-left-right (LRLR) eye movements. Filter settings are as follows:
In general, our data support the assumption that the pattern
EEG, bandpass filter 0.5270 Hz, with additional notch filter at 50 Hz; elec-
trooculogram (EOG), bandpass filter 0.1–30 Hz; EMG, bandpass filter
of activation in motor imagery largely overlaps with activity
16–250 Hz. corresponding to motor execution [24]. Moreover, the finding
of a similar action representation for hand clenching during
lucid dreaming extends the view of a common neural substrate
level [4, 12]. Cortical synthesis of internally generated sensori- for actions also for covert movements during this particular
motor perceptions may explain partially coherent narrative dreaming state. In this regard, dreamed motor activity might
dream mentation [13]. REM sleep is associated with increased help to elucidate the controversy about the locus of control
cerebral blood flow in visual association areas and limbic for the inhibition of motor output in motor imagery: because
regions as well as attenuated metabolism in dorsolateral pre- it is known that sleep atonia is generated at the brainstem level
frontal cortex, primary visual cortex, and precuneus [14, 15]. and spinal cord [4], the similarity between imagined and
Such a pattern of activation has been proposed to underlie dreamed motor activity might be interpreted as confirma-
general dream features like vivid imagery, emotional experi- tive for explanations in terms of a blocking mechanism down-
ences, and lack of insight and volition [4, 16]. stream of the motor cortex in motor imagery rather than a
The neural correlates of specific dream mentation, however, prefrontal blockade [24]. Controversial reports exist on the
still had to be disclosed. Efforts have been made to correlate involvement of M1 activation in motor imagery [23, 25, 26]:
REMs to gaze direction during dreams—the ‘‘scanning hy- although it is typically smaller compared with that in execution
pothesis’’ [1, 2]—and indeed similar cortical areas are involved [26], transient responses might be interpreted to reflect the
in eye movement generation in wake and REM sleep [17]. In onset of a specific motor imagination not leading to sustained
a similar vein, small muscle twitches during REM sleep were M1 activation [23]. The close interaction between somatosen-
presumed to signal a change in the dream content [3]. In sory and motor cortices during the performance of a move-
REM behavior disorder, patients even seem to enact dream ment differs between waking and dreaming. Lack of sensory
fragments [18], suggesting that dreamed motor actions in- feedback due to REM sleep atonia may further reduce the
volve activation of similar brain regions as during waking ac- activation in M1 and primary somatosensory cortex. However,
tions. Also in non-REM sleep, dreaming of task-related content small muscle twitches might well occur during dreamed hand
after an intense skill-learning session hints to the possibility clenching [3, 9], potentially leading to residual M1 activation.
that the same neural systems control executed and merely Because several limitations of our study complicate the
dreamed motor activity [19]. Nonetheless, the main obstacle interpretation of the activation pattern, the data have to be
in the direct neuroscientific study of specific dream content considered as very preliminary. Due to the scarcity of the
is that spontaneous dream activity cannot be experimentally phenomenon and the complexity of the methods, we basically
controlled because subjects typically cannot perform prede- present two case studies of two dreams each, assessed with
cided mental actions during sleep. Hence, dream research different methods, which naturally leaves the possibility of in-
methodology mostly relies on the evaluation of subjective re- dividual differences strongly influencing the results. Based on
ports of very diverse dream contents. However, such memory this, no statistical comparisons were made between condi-
traces collected after awakening are often imprecise. There- tions (WE, WI, LD) or regions. The lack of hand electromyo-
fore it seems difficult, if not impossible, to pinpoint specific gram (EMG) data further complicates the interpretation, and
dream content in a precise temporal frame, a prerequisite for the mismatch between the two subjects concerning SMA acti-
the analysis of imaging data acquired during dreaming. vation can hardly be explained satisfactorily. Hence, our data
In the present study, we employed the skill of lucid dreaming could not and should not be interpreted as a thorough clarifi-
to overcome some of these obstacles. In line with the concep- cation of the neural correlates of motor activity during dreams
tion that sleep atonia is generated at the brainstem level [4] and but rather as a methodological proof of concept on how to
confirming earlier EEG studies [20], fMRI BOLD responses measure neural correlates of dream content with neuroimag-
were observed bilaterally in the same sensorimotor cortical ing methods.
regions for lucid dreaming and wakefulness. However, during Although lucid dreaming comprises all defining markers of
dreaming, activation was much more localized in small clus- REM sleep proper [8] and all basal dream features such as
ters representing either generally weaker activation or focal hallucinations, it differs from nonlucid dreaming in its metacog-
activation of hand areas only, with signal fluctuations only in nitive insight into the hallucinatory nature of the dream state
the order of 50% as compared to the actually executed task and full access to cognitive capabilities [7, 27]. Recent sophis-
during wakefulness. Although in the fMRI analysis activation ticated EEG analyses have shown increased gamma band
Dreamed Movement Activates the Sensorimotor Cortex
1835

Figure 2. Comparison of Sensorimotor Activation during Wakefulness and Sleep

Functional magnetic resonance imaging (fMRI) blood oxygen level-dependent (BOLD)-response increases were contrasted between left and right hand
movements (columns) in the three conditions (rows): executed hand movement during wakefulness (WE) (A), imagined hand movement during wakefulness
(WI) (B), and dreamed hand movement during lucid REM sleep (LD) (C). Effects of left (right) hand movements were calculated in a fixed-effects analysis as
a contrast ‘‘left > right’’ and ‘‘right > left,’’ respectively. Subpanels depict results in an SPM glass-brain view (sagital and coronal orientation) to demonstrate
the regional specificity of the associated cortical activation, along with sensorimotor activation overlaid on an axial slice of the subject’s T1-weighted
anatomical scan (position indicated on the glass brain for condition A). Clusters of activation in the glass-brain views are marked using the numbering given
in Table S1. Red outlines in the glass-brain views mark the extent of activation found in the WE condition. This region of interest (ROI) was derived from the
Current Biology Vol 21 No 21
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plane resolution 3.125 3 3.125 mm2, TR 2 s, TE 40 ms) and NIRS were per-
formed in the early morning hours during which REM sleep incidence is
highest. The NIRS device was a custom-built monitor with 22 measuring
channels unilaterally covering right central sensorimotor areas. Five of the
six subjects underwent NIRS measurements with concurrent polysomno-
graphic recordings during sleep for 1–3 nights, and four of the six subjects
were assessed with fMRI with concurrent polysomnographic recordings for
2–6 nights. Subjects were instructed to signal the onset of a lucid dream by
left-right-left-right (LRLR) eye movements and to immediately start clench-
ing their left hand in their dream. They were further instructed to repeat the
LRLR signal after ten clenches and start to clench the right hand. The alter-
nating hand-clenching task was repeated for as long as possible, with the
LRLR signal indicating each changeover. Whereas a flat electromyogram
(EMG) indicated REM sleep atonia, eye signals were clearly visible and
distinguishable from spontaneous REMs in the EOG, providing unequivocal
temporal markers for imaging analysis (Figure 1). Polysomnographic data
were monitored online by a trained experimenter throughout the recording
session. Immediately after participants awakened, a short dream report was
prompted to verify lucidity and task compliance. Task performance was
Figure 3. Near-Infrared Spectroscopy Topography
considered successful when the following three conditions were concur-
Concentration changes of oxygenated (D[HbO], upper panel) and deoxy- rently fulfilled: (1) the subject was in REM sleep, (2) EOG signals showed
genated hemoglobin (D[HbR], lower panel) during executed (WE) and imag- at least four sets of LRLR eye signals, and (3) both lucidity and dreamed
ined (WI) hand clenching in the awake state and dreamed hand clenching hand clenching were confirmed by the subject in the subsequent verbal
(LD). The optical probe array covered an area of w7.5 3 12.5 cm2 over the report. When the EEG showed clear signs of awakening after a LRLR signal,
right sensorimotor area. The solid box indicates the ROI over the right one of the investigators entered the scanner or NIRS room and inquired
sensorimotor cortex with near-infrared spectroscopy (NIRS)-channels sur- about task performance. Alternatively, when LRLR signals were followed
rounding the C4-EEG electrode position. NIRS channels located centrally by a period of about 40 s without any further signals, one of the investigators
over midline and more anterior compared to sensorimotor ROI were chosen entered the scanner or NIRS room and inquired about task performance,
as ROI for the supplementary motor area (SMA, dotted box). thereby definitely awakening a potentially sleeping participant.
During fMRI, two subjects signaled and reported lucid dreams of suffi-
cient length, i.e., four sets of LRLR eye signals and hand clenchings. Poly-
activity over frontal and frontolateral areas when comparing
somnographic evaluation, however, revealed that one of the two subjects
lucid against nonlucid REM sleep, suggesting a neural corre- showed increased EMG amplitudes, which could indicate a transition to
late of this subjectively experienced cognitive control [28]. wakefulness. This subject was excluded from the analysis. The remaining
Hence, it cannot be excluded that neural activity associated subject showed two lucid dreams during which the task was correctly per-
with nonlucid dreams differs from that of dream content delib- formed, verified by polysomnographic data matching the dream report. Two
erately induced by a lucid dreamer. To assess regular dream further subjects reported lucidity during fMRI; however, one was not able to
perform the signaling or task as a result of sudden dream termination,
mentation in more detail, lucid dreamers might reenact certain
whereas the EEG of the other was compromised by strong sweating arte-
simple dream reports and the accordant activation patterns of facts, thereby making any objective evaluation of sleep stage, lucidity, or
lucid and nonlucid dreams could be compared. Although the task performance impossible. During NIRS, two subjects signaled and
neuroimaging of lucid dreaming is currently hampered by the reported lucidity, but only one subject was able to successfully conduct
scarcity of the phenomenon even in trained subjects, pro- the hand-clenching task (in two different dreams). We compared the activa-
mising attempts exist to enhance lucid dreaming frequency, tion pattern of lucid hand movements to executed and imagined hand
movements, collected during wakefulness with both neuroimaging modali-
e.g., by transcranial direct current stimulation [27]. ties (see Supplemental Information for details).
In summary, we provide the first demonstration of imaging
of specific dream contents by using the technique of lucid Supplemental Information
dreaming. This technique could be used to inversely infer
specific dream content from its underlying neural activity, Supplemental Information includes two tables, Supplemental Results,
allowing for true ‘‘dream reading.’’ Similar to first ‘‘brain Supplemental Experimental Procedures, and Supplemental Discussion
reading’’ experiments in the wake state [29], we chose a simple and can be found with this article online at doi:10.1016/j.cub.2011.09.029.
hand-clenching task because motor activity can be reliably
Acknowledgments
located by both imaging modalities applied. The combination
of lucid dreaming with neuroimaging and polysomnography We thank J. Allan Hobson for his advice and discussions on this topic.
is a promising technique to also transfer more sophisticated
‘‘brain reading’’ tasks [30] to the dreaming state. Future appli- Received: February 9, 2011
cations might therefore lead to the imaging of prototypical Revised: August 31, 2011
dream content like visual dream imagery or dreamed emotions. Accepted: September 15, 2011
Published online: October 27, 2011

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Dreamed Movement Activates the Sensorimotor Cortex
1837

Figure 4. Condition-Related NIRS Time Courses

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