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Curr Neurol Neurosci Rep. Author manuscript; available in PMC 2016 January 07.
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Published in final edited form as:

Curr Neurol Neurosci Rep. 2014 March ; 14(3): 433. doi:10.1007/s11910-013-0433-5.

Dreaming and Offline Memory Consolidation

Erin J. Wamsley
Department of Psychiatry, Harvard Medical School, Boston, MA 02115, USA

Department of Psychology, Furman University, Johns Hall 206K, 3300 Poinsett Highway,
Greenville, SC 29613, USA
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Abstract
Converging evidence suggests that dreaming is influenced by the consolidation of memory during
sleep. Following encoding, recently formed memory traces are gradually stabilized and
reorganized into a more permanent form of long-term storage. Sleep provides an optimal
neurophysiological state to facilitate this process, allowing memory networks to be repeatedly
reactivated in the absence of new sensory input. The process of memory reactivation and
consolidation in the sleeping brain appears to influence conscious experience during sleep,
contributing to dream content recalled on awakening. This article outlines several lines of
evidence in support of this hypothesis, and responds to some common objections.

Keywords
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Sleep; Dreaming; Rapid eye movement; Non-rapid eye movement; Memory consolidation; Offline
processing; Memory reactivation; Replay; Consciousness

Introduction
Accumulating evidence suggests that memories are “consolidated” in the sleeping brain.
Across a wide variety of memory domains—including verbal [1–4], emotional [5–7], motor
[8, 9], perceptual [10], and spatial learning [11–14]—postlearning sleep has been shown to
benefit later memory performance. Could dreams be a reflection of this memory processing
during sleep? It has long been recognized that daily experience influences dreaming.

Even prior to psychoanalytic dream theory, early experimentalists postulated a systematic

connection between waking experience and the content of our nightly dreams [15, 16]. As
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reviewed here, recent research confirms that dream content transparently reflects recently
encoded memory. Meanwhile, recent discoveries in psychology and neuroscience suggest
that the reactivation of memory networks during sleep aids consolidation, with both animal

[Link]@[Link].
This article is part of the Topical Collection on Sleep
Compliance with Ethics Guidelines
Human and Animal Rights and Informed Consent This article does not report original research findings. Some studies described
here were performed in our laboratory at Beth Israel Deaconess Medical Center, where all human subjects signed informed consent
prior to participation.
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and human studies demonstrating that patterns of brain activity first seen during learning are
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later “replayed” in sleep. Together, these data suggest that dreaming arises, at least in part,
from the reactivation and consolidation of memory in the sleeping brain. In the following
section, we will examine the evidence for this hypothesis, and explore several potential
counterpoints.

Evidence Linking Sleep-Dependent Memory Consolidation with Dreaming

New Learning Experiences Are Incorporated into Dreams—A first clue to the
relationship between learning, memory, and dreaming comes from the manner in which
waking experience influences dream content. Although it may seem patently obvious that
we dream about things experienced during the day, quantifying this phenomenon in the
laboratory initially proved to be difficult. In the 1960s and 1970s, investigators endeavored
to manipulate and predict the effects of specific waking experiences on dream content. But
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early attempts to influence dreams using manipulations such as films and images [17–20]
were near uniform in their failure to detect statistically significant direct1 effects of presleep
stimuli on dream content (for a review, see [21]).

But these early studies were nonetheless informative. First, the laboratory setting itself was
consistently found to have a powerful influence on dreams. Illustrating this robust effect, in
an analysis of 813 rapid eye movement (REM) mentation reports collected across several
studies, Dement et al. [22] reported that 22% of reports unambiguously incorporated either
isolated elements (10%) of the laboratory situation (i.e., the experimenter, the sleep
laboratory itself, electrodes, etc.) or more complete representations (12%) of the
experimental setting (i.e., a combination of elements). In retrospect, this observation seems
to indicate that the most salient aspects of our presleep experience are preferentially
incorporated into dreaming. For research participants, the novel experience of sleeping in a
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strange laboratory with wires attached to their head may be a much more impactful event
than any particular stimulus used as part of a psychological experiment.

Second, a handful of studies began to suggest that experimental manipulations involving

exposure to novel learning experiences more easily affect subsequent dream reports.
Roffwarg and coworkers [23, 24], for example, examined how the long-term use of red-
tinted goggles impacted dreaming, finding evidence that this shift in waking color
perception altered the reported perceptual qualities of dreaming. In analogous studies of
long-term perceptual alteration, subsequent research examined dream reports during
adaptation to the use of prism goggles that invert the visual field, similarly finding evidence
for incorporation of perceptual alterations into subsequent dreaming [25, 26]. As discussed
further later, studies by Fiss et al. [27] and De Koninck et al. [28] were likewise able to
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influence dreaming through the use of novel learning experience. The relative success of
these paradigms, as opposed to the passive viewing of experimental stimuli before sleep,
suggests that novel learning experiences have a particularly pronounced impact on dream
experience.

1As opposed to “symbolic”

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More recent studies have continued to demonstrate the effect of novel learning on dream
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content. Engaging, interactive video game tasks have shown particularly powerful effects.
For example, in one study, participants played the video game Tetris extensively prior to
dream report collection [29]. Mentation reports were then repeatedly elicited during early
non-REM (NREM) sleep at sleep onset—64% of participants reported unambiguous2 Tetris
images. This basic effect of Tetris on sleep-onset imagery was later independently replicated
by Kusse et al. [30]. Related investigations using a downhill skiing arcade game [31] and
virtual navigation tasks [32, 33] have similarly demonstrated the robust incorporation of a
presleep learning task into dream content. Although, for practical reasons, most of these
studies have examined dream reports only from the first few minutes of NREM sleep, we
have seen that this effect extends into later night dreaming during both stage 2 and REM
sleep [34]. The frequency of direct incorporation in these sleep studies—which is
dramatically higher than that observed in previous investigations—again suggests that
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engaging learning experiences may have a particularly robust influence on dream content,
relative to viewing static images or films.

Other research has sidestepped the difficulty of prospectively manipulating dream content
by asking participants to retrospectively identify the waking memory sources of dreams. For
example, in an analysis of the possible waking memory sources of 299 home-collected
dream reports, Fosse et al. [35] demonstrated that fragments of recent experience quite often
appear in dream reports. In this study, 51% of 299 dream reports were judged by subjects to
contain at least one feature with strong similarity to a recent waking event. Others have
taken a similar retrospective approach, establishing that participants frequently identify
recent memories as a source of dream content [36, 37], especially for dreams recalled from
NREM sleep [38, 39]. These investigations highlight that although attempts to predict or
control the waking source of specific dream elements have often failed, the influence of
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experience on dream content is nonetheless transparently clear.

Taken together, these data suggest that waking experiences are transparently represented in
the content of dreams. Furthermore, it may be that particularly salient daytime experiences
are preferentially incorporated into dreaming, with novel learning being a particularly strong
driver of dream content.

Memories Are “Reactivated” in the Sleeping Brain

It is increasingly clear that sleep facilitates the consolidation of memory. A large number of
studies have now replicated the robust finding that sleep following learning improves
memory performance at a subsequent test [1, 3, 5, 8, 10, 40–43]. Recent discoveries in the
neuroscience of learning and memory have been particularly enlightening in thinking about
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just how memory might be facilitated during sleep. Groundbreaking research in the last two
decades has established the remarkable fact that waking patterns of brain activity are
“replayed” during sleep in rodents [44–49]. For example, sequences of hippocampal place
cell activity seen while rats explore a spatial environment during wakefulness are again
observed when the rats fall asleep [48, 49]. As the firing of each of these hippocampal

2An example report from one participant: “[I'm] seeing in my mind how the game pieces kind of float down and fit into the other
pieces, and am also rotating them.”

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“place” cells is associated with a particular physical location in the environment, the
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reappearance of these sequences suggests that during sleep rats are retracing the paths they
have learned. Initially described in the hippocampus, this effect was later seen in cortical
regions as well [44, 46]. Although reactivation on the single neuron level has not yet been
observed in humans, imaging studies demonstrate that brain regions engaged during
presleep learning are again preferentially activated during sleep, and that this learning-
related activity predicts postsleep task improvement [50, 51].

The discovery of memory reactivation during sleep certainly suggests a parallel with dreams
of recent experience. Perhaps rats are dreaming of the “maze” they traversed while awake.
Of course, we know that not all brain activity is consciously experienced. The neural activity
associated with regulation of respiration and heart rate, for example, does not influence our
stream of conscious awareness. Analogously, these cellular firing sequences in small regions
of rodent hippocampus and cortex may be irrelevant to understanding the stream of
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consciousness during sleep. But is this so-called memory “replay” during sleep at least a
good potential candidate for a neural correlate of dreaming? A complete review of the
memory reactivation literature is beyond the scope of this article, but it is noteworthy that
several key features of this phenomena parallel the structure of dream experience:

1. In rodent studies, memory reactivation has most often been reported to occur in
NREM sleep. As described earlier, NREM sleep is also the sleep stage during
which episodic memories are most often incorporated into dreaming.

2. Within NREM sleep, reactivation is expressed most strongly immediately after

learning. The strength of this reactivation effect then tends to decay quickly across
time [49, 52]. Similarly, sleep mentation may be most strongly related to recent
experience early in the sleep phase [29, 31, 53].
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3. Learning-related neural activity is not reactivated exactly in its original form.

Waking firing sequences are reactivated in intermittent bursts, on a faster timescale
than the original experience [49]. And rarely is a waking firing sequence reiterated
exactly—although reactivated sequences statistically resemble waking activity, the
“replay” is not always precise. Similarly, recent experiences are not “replayed” in
dream content in their original form. Instead, dreams intermingle fragments of
recent experience with other content [31, 35].

Dreaming About Learning Experiences Is Associated with Enhanced Memory

Memory reactivation during sleep is thought to lead to consolidation and enhancement of
postsleep memory performance [54]. If dreaming reflects memory reactivation, we might
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expect that dreams should “index” the consolidation process, tracking the extent to which a
particular memory is processed during sleep. In fact, evidence dating back to the 1970s
suggests that dreaming of a learning experience is associated with enhanced memory for that
information. Fiss et al. [27], for example, found that after reading the text of a short story,
participants who reported dreams related to the story exhibited superior memory for the text
the following morning. De Koninck et al. [28] also examined dreams and verbal learning,
exploring dream content as a corollary of language learning in an academic setting. Among
students enrolled in a French-immersion class, those with the strongest language acquisition

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across the 6-week course incorporated French into dream content more often than students
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who were less successful in the class. Most recently, our laboratory has demonstrated that
dreaming of a virtual maze navigation task is associated with enhanced consolidation of
spatial memory both across a nap [32] and across a full night of sleep [34]. Demonstrating
that dreaming of a recent learning experience is associated with enhanced memory, these
studies suggest that dream content does reflect the consolidation process.

Together, these observations lead us to the hypothesis that reactivation of memory traces
during sleep causes memory-related content to be incorporated into dreams. As an index of
memory reactivation, learning-related dreaming predicts subsequent memory performance.
In this sense, dreaming may be considered a reflection of the consolidation process. In the
following section, addressing several counterpoints will help us in further specifying the
hypothesis and its predictions.
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Objections to the Hypothesis

Dreams Are Not an Accurate Replay of Waking Experience
It is sometimes argued that dreams could not reflect an adaptive process of memory
consolidation because they fail to faithfully replicate waking experience [55, 56]. Vertes
[55], for example, has argued that “the hypothesis that sleep serves a role in memory
consolidation would be more appealing if dreams reproduced waking experiences (or even
approximately so), but they do not.” The late dream researcher Ernest Hartmann [56]
similarly argued that the content of dreams is incompatible with memory consolidation
because dream reports do not veridically replay waking experience.

It is true that episodic memories are rarely or never “replayed” in dreaming exactly as they
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occurred. Instead, dreams often incorporate only isolated fragments of a waking episode,
and these may be interleaved with remote and semantic memory material. For example,
although Fosse et al. [35] reported that more than half of dreams contained features similar
to a recent experience, less than 2% of these preserved the original setting, characters,
objects, and actions of the waking memory source. Consider the following example of a
waking memory source and corresponding report (taken from [35]):

Waking experience: “When I left Starbucks, we had so many leftover pastries and
muffins to throw away or take home. I couldn't decide which muffins to take and which
to toss...”

Corresponding dream experience: “My dad and I leave to go shopping. We go from

room to room, store to store. One of the stores is filled with muffins, muffins, muffins
from floor to ceiling, all different kinds, I can't decide which one I want...”
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Here, it would certainly be maladaptive for the dreamer to falsely remember having been in
a room filled with muffins while shopping with her father—a fantastical event that never
occurred.

But increasingly, we are learning that the effect of sleep on memory is not merely to cement
experience in its original form. Instead, sleep transforms memory traces over time, allowing
us to extract generalizations [57, 58], integrate information [41, 59], and arrive at creative

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insights [40, 60]. Models of memory consolidation have long stressed the integration of new
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information into established networks—the influential “complementary learning systems”

model, for example, proposes sleep as a time when recently encoded information is
“interleaved” into related memory networks [61]. This could be achieved by simultaneously
reactivating recent and remote memory fragments, slowly associating new content into
cortical semantic networks. Lewis and Durrant's recent information overlap to extract model
of memory consolidation similarly postulates that sleep facilitates schema development
through the repeated reactivation of related information [62••]. As in the example above,
dream reports often bizarrely intermingle recent experience fragments with related material
—this may be exactly the type of process that facilitates information integration during
sleep.

The rodent memory reactivation literature also supports this conclusion. Studies of memory
“replay” in rodents do not demonstrate reactivation of experience in its original form. First,
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learning-related patterns of activity are reactivated with some degree of distortion. But also,
there is evidence that multiple related experiences are reactivated simultaneously during
sleep [49]. This recombination of waking experiences could create an amalgam that serves
to facilitate insight and generalization—in fact, it has recently been observed that replay can
include the playing out of novel spatial trajectories never experienced while awake, some of
which are “shortcuts” through a learned environment [63].

Thus, although dreams do not faithfully replay waking experience, we should not expect
them to. Memory consolidation is not thought to involve the repeated replay of experience in
its original form. To the contrary, sleep facilitates integration across multiple memories and
the extraction of generalities. The intermingling of memory fragments seen in dream reports
could well reflect this adaptive process.
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Memory Reactivation Is a Poor Candidate for the Neural Correlate of Dreaming Because It
Does Not Happen in “Real Time”
During rodent sleep, the reactivation of waking sequences occurs on a much faster timescale
than the original experience [44, 49]. In contrast, there is compelling indirect evidence that
dream events occur on the same timescale as waking experience [64]. This mismatch
suggests that the brain processes of memory consolidation cannot be strictly identical to
those that create dreaming. Still, it may be that memory reactivation influences dream
content. The brief, localized bursts of memory reactivation observed in rodent models could
feed into the dream creation process by affecting downstream mechanisms of dream
generation. In short, memory reactivation as observed in rodents is not likely to be “the”
neural mechanism of dreaming, but it may nonetheless be an important contributor.
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Not All Dreaming Is Even Tangentially Related to a Past Memory

Only a portion of dream content can be definitively traced back to a specific prior
experience [35, 38]. Dreams about possible futures, fantastical places, and people that we
have never met may show no evidence of being tied to any episodic memory. But in a
broader sense, all dreams (and all waking thought, for that matter) are most certainly
constructed from prior experience. The appearance of a complete stranger in a dream, for

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example, relies on knowledge of the visual characteristics of the human face and on schemas
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for typical human behavior. This semantic memory has been generalized and extracted over
the course of our lifetimes on the basis of thousands of individual past experiences. In a
sense, this form of generalized memory can be thought of as the end product of
consolidation. Through consolidation, individual experiences are integrated in order to
extract commonalities [38, 61]. Certainly, the activation of semantic memory networks
during sleep could occur independently of any consolidation-related process, perhaps
reflecting more or less random activation of distributed cortical networks [65]. But
alternatively, activation of semantic knowledge during sleep could reflect the “updating” of
cortical networks as new information is slowly incorporated into existing schemas (e.g., as
described in the information overlap to extract model [62••]).

However, the fundamental claim of the memory consolidation hypothesis—that dreaming

reflects consolidation in the sleeping brain—does not require that all dream content result
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from memory-related activity. It may be that the activity of memory systems comprises only
a portion of the total brain activity underlying dreaming, with diffuse activity unrelated to
memory also influencing conscious experience during sleep. The most fundamental form of
the hypothesis would not claim that every part of every dream can be explained as a
memory-related process—only that memory consolidation is one major influence on
dreaming.

Dreams Do Not Seem Functional

The content of dreams often does not seem particularly helpful. The philosopher Owen
Flanagan [66] wrote that “since we rarely dream about what we need to remember, the
hypothesis that dreams themselves serve any memory enhancing function appears
unwarranted.” Certainly, as in the example above, dreams typically do not appear to be a
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rehearsal of things that are important to remember. Although anecdotal evidence of dreams
containing insight, creativity, and problem-solving abounds, empirical data are lacking. In
fact, most dream reports contain negative emotions, failures, or misfortunes [67]. No
problems are solved, no insights are arrived at, no fears are overcome. The bizarre and
apparently irrelevant content of dreams has inspired proposals that we dream in order to
forget spurious associations [68], or that the content of dreams is determined in an
essentially random fashion [65]. In short, the content of recalled dreams does not in itself
reveal any obvious function.

But a dream that arises from a functional brain process need not seem functional in itself.
Supposing that memory consolidation does influence dreaming, this would not necessarily
lead dreams to seem positive or helpful in any way. Memory consolidation involves the
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stabilization of recently formed memories against interference, as well as the functional

reorganization of memory representations in the brain—these processes should not be
expected to give rise to conscious experiences that mimic the waking experience of
memorization, problem-solving, or creative insight. It is important to distinguish the
functionality of memory processing during sleep from the notion that the content of dreams
itself is functional—the former may be true without the latter.

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It is not clear whether dreaming per se has a function at all. The data reviewed here suggest
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only that dreaming is influenced by brain processes that have a clearly demonstrated
function—as memories are processed in the sleeping brain, this functional activity
influences the ongoing content of a dream. To understand whether dreaming itself can be
described as having a function, we will need first to understand the degree to which the brain
basis of memory consolidation overlaps with the neural substrate for dreaming. Beyond this,
a “function” for dreaming additionally hinges on the difficult question of whether conscious
experience in general serves any function. Here, we suggest only that conscious experience
during sleep is influenced by a neurophysiological function of the sleeping brain.

Conclusions
More than 50 years after the discovery of REM sleep launched the field of sleep research,
the study of dreaming remains in its infancy. In recent years, however, new insights into the
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neurobiology of memory have led to the hypothesis that dreaming reflects the reactivation
and consolidation of memory in the sleeping brain. In its most plausible form, this
hypothesis asserts that:

1. Waking experiences do not appear in dreams in their original form, and we should
not expect them to. Memory consolidation involves the integration of multiple
experiences and the extraction of generalities. In dreams, the intermingling of
memory fragments into novel and sometimes bizarre combinations could reflect
this adaptive process.

2. The brain mechanisms of dreaming are likely not identical to those responsible for
memory consolidation. Although the content of dreams is influenced by memory
consolidation, it may be that not every element of every dream is related to this
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process. If conscious experience during sleep is the emergent result of neural

activity distributed across much of the brain, only a portion of this activity would
be expected to be influenced by the activity of memory systems.

3. Dreaming reflects the functional brain process of memory consolidation, but this
does not mean that dreams, per se, have a function.

As for waking conscious experience, a single cognitive or brain system is unlikely to

provide a complete explanation for the complexity of subjective experience during sleep.
Ultimately, it may be that only a portion of dream experience originates from the
consolidation of memory. And even for this portion, we have not yet arrived at a satisfying
explanation. The evidence described here tells us that memory consolidation likely
influences dream experience, but says little about exactly how this might occur. To explain
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the role of memory in dreaming, we need to arrive at a more complete understanding of how
memory consolidation is instantiated in the brain and how it interacts with the neural
substrate for dreaming. Even given these qualifications, however, the memory consolidation
hypothesis provides a framework for moving forward with the study of subjective
experience during sleep. Far from being a meaningless distraction, dreams may reflect an
adaptive process by which new learning is stabilized and integrated with our existing
knowledge. As such, the content of these experiences could provide an important and unique
source of information about the functions of the sleeping mind and brain.

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Acknowledgments
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Conflict of Interest Erin J. Wamsley declares that the research described here was supported by National Institute
of Mental Health grant R01-MH48832 (principal investigator Robert Stickgold), National Institutes of Health T32
training grant HL07901-10 to the Harvard Division of Sleep Medicine, and a KL2 Medical Research Investigator
Training award (an appointed KL2 award) from Harvard Catalyst | The Harvard Clinical and Translational Science
Center (National Center for Research Resources and the National Center for Advancing Translational Sciences,
National Institutes of Health award 8KL2TR000168-05). She is also a principal investigator supported by an R21
grant from the National Institute of Mental Health (1R21MH098171-01A1) and has received grants from the BIAL
Foundation. She has also received compensation for teaching a course at Harvard University.

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