Dreaming and Memory 1

Running head: DREAMING AND MEMORY

Dreaming of a Learning Task is Associated with Enhanced Memory Consolidation:

Replication in an Overnight Sleep Study

Erin J. Wamsley1 & Robert Stickgold2,3

1
Department of Psychology and Program in Neuroscience, Furman University, Greenville SC
2
Department of Psychiatry, Harvard Medical School, Boston MA
3
Department of Psychiatry, Beth Israel Deaconess Medical Center, Boston MA

Word Count: 4,212

References: 34

Corresponding Author:

Erin J. Wamsley
Assistant Professor of Psychology
Furman University
Johns Hall 206K
phone: 864-294-3218
fax: 864-294-2206
erin.wamsley@furman.edu
 Dreaming and Memory 2

Summary

Sleep following learning benefits memory. One model attributes this effect to the iterative “reactivation”

of memory traces in the sleeping brain, demonstrated in animal models. Although technical limitations prohibit

using the same methods to observe memory reactivation in the human brain, the study of mental activity during

sleep provides an alternative method of observing memory activation during sleep. In fact, the content of dream

experience may reflect the process of memory reactivation and consolidation in the sleeping brain. In line with

this hypothesis, we previously reported that dreaming about a spatial learning task during a nap strongly

predicts subsequent performance improvements. Here, we replicate this observation in an overnight sleep study,

for the first time demonstrating that pre-sleep training on a virtual maze navigation task is reflected in dreams

reported from all phases of sleep, with unambiguous representation of the task in dream content associated with

improved next-morning performance. These observations are consistent with reactivation-based models of

memory consolidation in sleep, confirming our earlier finding that the cognitive-level activation of recent

experience during sleep is associated with subsequent performance gains.

Keywords: Cognition, Learning, Sleep Mentation

 Dreaming and Memory 3

Introduction

Post-training sleep is beneficial for human memory (Diekelmann and Born, 2010; Stickgold, 2005).

Neurophysiological studies in rodents suggest that this is due to an iterative “reactivation” of memory networks

during sleep, strengthening synaptic connections and facilitating memory consolidation (Ji and Wilson, 2006;

Wilson and McNaughton, 1994). Human studies have also reported analogous observations, for example in

PET studies finding that patterns of neural activity present during encoding are reiterated during subsequent

sleep (Maquet et al., 2000; Peigneux et al., 2004). During sleep, the activity of memory networks is also evident

in the content of subjective experience, as recent waking experiences are often unambiguously represented in

the content of our nightly dreams (Stickgold et al., 2000; Wamsley and Stickgold, 2009). Indeed, the

observation that events from the previous day appear in dream content stretches back more than a hundred years

(Freud, 1913). Yet only recently has the effect of waking experience on dream reports been studied in the

laboratory using scientifically rigorous methods. Recently, we reported that dreaming of a learning task was

associated with improved memory for that information following sleep (Wamsley et al., 2010b). This

observation echoes earlier work in non-laboratory contexts, for example showing that participants enrolled in a

language immersion course show superior grade outcomes when their dreams incorporate the foreign language

(De Koninck et al., 1990).

The observation that patterns of neural activity representing recent experience are “reactivated” in the

sleeping brains of rodents (e.g. Wilson and McNaughton, 1994) suggests that sleep’s effect on human memory

might be attributed to a repeated “replay” of experience during sleep allowing cortical memory networks to be

strengthened and integrated. Indeed, recent studies demonstrate that hippocampal sharp-wave ripples (during

which memory reactivation preferentially occurs) increase following learning (Eschenko et al., 2008), and that

experimentally disrupting ripples impairs memory (Ego-Stengel and Wilson, 2010). Still, alternative accounts

attribute the memory benefits of sleep to other mechanisms. For example, a recent extension of the “synaptic

homeostasis hypothesis” suggests that a selective downscaling of synaptic strength during sleep could facilitate

memory by enhancing signal-to-noise ratio, without the need for invoking either memory reactivation or
 Dreaming and Memory 4

synaptic potentiation (Nere et al., 2013). In fact, despite the evidence described above, there has not yet been a

direct demonstration that the type of cellular-level memory reactivation observed in rodents even correlates with

sleep’s beneficial effect on memory in humans.

Demonstrating the reactivation of specific memory networks during human sleep has been challenging due

to technical limitations – the large scale single-cell recordings enabling this work in rodents are not available in

human subjects, and the spatial and temporal resolution of noninvasive neuroimaging methods are inadequate to

the task. Our own work has focused on mental activity during sleep as information about the activation of

memory networks. Our laboratory and others have demonstrated that memories formed during engaging,

interactive visuospatial learning tasks are clearly incorporated into dream experience during post-training sleep

(Kusse et al., 2011; Solomonova et al., 2015; Stickgold et al., 2000; Wamsley et al., 2010a). These dream

reports are not only “symbolically” or weakly related to prior experience. For example, after playing the video

game “Tetris”, 63% of participants reported unambiguous imagery of the game during the first minutes of sleep,

with one participant reporting:

…just seeing Tetris shapes floating around in my head like they would in the game, falling

down, sort of putting them together in my mind (Stickgold et al., 2000).

In another study, after playing the downhill skiing arcade game Alpine Racer II, 23% of reports contained

unambiguous reference to the game, at times including imagery of specific locations on the virtual slope. For

example, one participant in these studies reported that:

I get like flashes of that . . . game in my head, virtual reality skiing game . . . downhill umm

race, in my head. Umm, there’s this one particular corner that I haven’t quite been able to

master, and every time I get flashes of it, it’s like that corner that umm I keep crashing into

(Wamsley et al., 2010a).

Although these reports were collected from the first few minutes of sleep (the “sleep onset” period), we later

found incorporation of a virtual maze navigation task during N2 sleep further into the sleep cycle of a daytime

nap (Wamsley et al., 2010b). Together with other evidence (De Koninck et al., 1990; Kusse et al., 2011; Nielsen
 Dreaming and Memory 5

and Stenstrom, 2005; Tauber et al., 1968), these observations provide a clear demonstration that memory

networks encoding recent learning experiences are activated during sleep.

This evidence raises the question of whether the activation of memory networks underlying dream content is

related to sleep-dependent memory consolidation and to the effects of sleep on human memory. Importantly,

this is not necessarily the case—the appearance of recent experience in dreams could be driven by neural

processes entirely independent of those which support the consolidation of memory. However, we have

previously reported that participants who dreamed about a virtual maze navigation task during a daytime nap

showed a tenfold greater performance gain at subsequent test than those whose dream reports were unrelated to

the task, supporting this hypothesis (Wamsley et al., 2010b). This observation suggests that dreaming of recent

experience reflects the reactivation and consolidation of memory in the sleeping brain, which in turn leads to

post-sleep performance improvements.

But rigorous science demands the replication of surprising new observations before accepting them as fact.

The generalizability of our original study could be criticized on several grounds. First, these observations were

based on only a small number of participants who incorporated the maze task into dream content, and dreams

were collected only from NREM sleep during a short daytime nap. Second, several subsequent studies testing

the same hypothesis using either a different learning task (Schredl and Erlacher, 2010), or a modified version of

the VMT (Stamm et al., 2014; Wamsley et al., 2016), failed to find any association between incorporation of the

task into dreaming and subsequent memory.

In the current study, we sought (1) to replicate our prior observation that dreaming of the VMT learning task

is associated with memory enhancement, (2) to extend this observation to reports from all sleep stages across a

full night of sleep.

 Dreaming and Memory 6

Methods

Participants

N=39 healthy college students (20±2SD yrs; 33% male) completed the study. Participants reported a

mean habitual sleep time of 7.0±1.1SD hrs, and 66.7% of the sample reported habitual napping on an average of

3.6±1.6SD days/week. No participants reported diagnosis of a sleep disorder or mental disorder. One participant

was under treatment with an antibiotic, and no others reported current medication use. Frequency of 3-D style

video game play averaged 2.1 on a 5-point ordinal scale, with the majority of participants reporting game play

at least once per month (35%) or once per year (30%), with a minority of participants reporting game play once

a week (22.5%) or every day (5%), or less than once a year (7.5%). Participants were randomly assigned to

either a “Sleep” group (n=21) or an “Immediate Test” control group (n=18). Two additional control participants

enrolled in the study but failed to complete testing due to “cybersickness” induced by the Virtual Maze Task

(VMT).

Following criteria established in our earlier work (Wamsley et al., 2010c), participants were excluded

from VMT performance analyses if they reported being excessively sleepy at the time of maze training

(Stanford Sleepiness Scale (SSS) ≥5) or playing 3D style video games less than once per year (Wamsley et al.,

2010c). Following exclusions, n=17 Sleep and n=16 Immediate Test participants were included in analyses of

VMT performance. Based on the very large effect size reported in our previous paper (d=2.02 (Wamsley et al.,

2010b)), this sample size was adequate to test our primary hypothesis with power >0.9. This research was

approved by Beth Israel Deaconess Medical Center’s IRB.

[FIG1 HERE]

Procedures

Procedures closely followed those in our prior study of dreaming during daytime naps (Wamsley et al.,

2010b), with the primary exception being that here we tested participants across a full night of sleep (Fig1).

Participants in the Sleep group reported to the laboratory at approximately 9PM, signed written informed
 Dreaming and Memory 7

consent, and completed questionnaires including a demographics information form, 3-day retrospective sleep

log, Epworth Sleepiness Scale (ESS), and Beck’s Anxiety Inventory. Participants were then wired for

polysomnographic recording, with 58 EEG electrodes placed according to the international 10-10 system of

electrode placement, bipolar chin electrodes for recording electromyography, and electrooculography electrodes

placed at the left and right outer canthus. At 11PM, participants completed the SSS as a measure of state

sleepiness before reclining with their eyes closed for a 10min baseline EEG recording of resting wakefulness.

Participants were then trained on the Virtual Maze Navigation Task (VMT, described below) before undergoing

a second 10min resting EEG recording and completing visual analog scales describing subjective task

engagement and difficulty. Lights were then turned off and participants began an 8.5hr sleep opportunity,

during which subjective experience was sampled as described below. Sleep participants were awoken at 8am

the following morning and were retested on their ability to solve the same maze approximately 30min later.

Sleep participants were prompted to provide open-ended verbal reports on their mental experience

during both the waking rest periods and nocturnal sleep. During each 10min rest period, participants were

prompted to provide 2 reports. During the first hour of sleep, participants were awakened to provide up to 10

sleep onset reports, after 30sec, 60sec, or 90sec of PSG-defined sleep (order of sleep latency awakenings

pseudorandomized). During the remainder of the night, participants were awoken three times from N2 NREM

sleep (on each occasion following at least 10min of continuous N2) and provided one final report upon morning

awakening, regardless of sleep stage. Subsequent analysis showed that, amongst the full sample of participants,

morning awakenings were from N2 sleep in 10 participants, and were from REM sleep in 10 participants. For

one participant, sleep stage upon morning awakening could not be determined due to excessive artifact in the

EEG record. For each report, participants were contacted by calling their name over an intercom, at which point

they were prompted to provide an open-ended verbal report describing “everything that was going through your

mind” just prior to awakening.

Participants in the Immediate Test control group trained and were tested on the maze without

intervening sleep. Immediate Test subjects reported to the laboratory and completed the same initial paperwork
 Dreaming and Memory 8

as Sleep group participants prior to training on the VMT, as described as below. Following training,

participants took a 15min break during which they were allowed to stand up and move around the laboratory,

after which they were immediately tested on the maze. No mentation reports were collected from Immediate

Test participants.

Virtual Maze Task

The Virtual Maze Learning Task (VMT) has been described in several of our prior papers (Nguyen et

al., 2013; Stamm et al., 2014; Wamsley et al., 2010b; Wamsley et al., 2010c). Constructed using the Unreal

Tournament 3 Editor (Epic Games), the task is implemented on a PC computer and projected onto a screen in a

darkened testing room. Participants navigate through the virtual environment using a USB number pad with

four arrow keys. The training session begins with a 5min exploration period during which participants are

placed at the maze exit and instructed to explore as much of the maze as possible and to remember it as well as

possible, as they will subsequently be tested on their ability to locate the exit. This exploration period is

followed by a series of three training trials in which participants attempt to find the exit as quickly as possible.

For each trial, participants began at a different starting point, each equidistant from the exit, with the order of

starting positions counterbalanced across participants. In this version of the task, participants were incentivized

with a performance-based payment (Stamm et al., 2014). A monetary reward value was displayed on screen at

the beginning of each trial, and counted down as the participant played. Participants were told that the

remaining money was theirs to keep at the end of the trial. Trials ended when subjects reached the exit of the

maze or after a maximum of 10min. Retesting consisted of three additional trials using the same starting-point

order as during training, but without the 5min exploration period.

The primary dependent measure of “Completion Time” is the number of seconds required to reach the

maze exit on each trial. To facilitate further analysis of movement behavior during the task, the maze space is

divided into a 20x20 grid. “Distance Traveled” to the exit is defined as the total number of grid boundaries

traversed in each trial. “Backtracking” (1 – [number of unique positions / Distance Traveled]) measures the

extent to which participants retrace their steps while navigating towards the maze exit. For all performance
 Dreaming and Memory 9

measures, overnight improvement was calculated as the change from the last training trial to the mean

performance on the 3 retest trials (Wamsley et al., 2010b).

Data Analysis

Report transcripts were scored for content related to the learning task by two raters blind to subject,

experimental condition, sleep stage, and maze performance. Following procedures established in our earlier

work (Wamsley et al., 2010a), reports were scored first according to whether the participant recalled any mental

content, and then according to whether the report directly mentioned the maze task (making explicit reference to

the maze task from the experiment or another maze, see Table 1) and/or described content indirectly related to

the maze (incorporating sensations, objects, locations, actions or themes similar to those contained in the maze).

To test whether maze-related content was reported at greater-than-chance levels, raters additionally

scored N=100 control dream reports collected in a similar sleep study (Rowley et al., 1998), in which

participants were awakened to provide mentation reports without having trained on a learning task prior to

sleep. Control reports were randomly intermixed with the experimental reports given to raters for scoring.

Inter-rater agreement was high for the presence of mental content (Cohen’s Kappa=0.75), and for direct

incorporation of the learning task into dreaming (Kappa=0.89). Disagreements were resolved by a third rater,

also blind to subject, condition, sleep stage, and performance. However, agreement on whether reports

indirectly incorporated the learning task was very poor (Kappa=0.07). Scores of indirectly related content were

also not correlated with any performance improvement metric. Data on indirect incorporation are therefore not

further discussed in this paper.

Results

Incorporation of the Maze Task into Dreaming and Waking Cognition

347 mentation reports were elicited across states of sleep and wake, 310 (89%) of which contained at

least some mental content (see Tables 1&2). All told, 39 reports contributed by 12 of the 17 Sleep participants

included in analysis (71%) contained content related to the maze task, and these occurred across all conscious
 Dreaming and Memory 10

states (Table 2). In contrast, no Control reports (Rowley et al., 1998) were judged to contain maze-related

content. In all states except for N2, task incorporation was significantly greater than the 0% rate established in

the Control set (chi-square tests of independence, Table 2).

Waking rest reports contained incorporation of the learning task even before participants were first

exposed to the maze (Table 2). The content of these pre-exposure reports reflected anticipation of the upcoming

learning task, which participants had been informed would be occurring shortly. During post-training rest,

incorporation of the learning task increased significantly above pre-training levels (52% of reports vs. 11%,

Wilcoxon rank sign test, Z=2.60, p=0.009; Fig2).

Both Dreaming and Waking Cognition Predict Next-Morning Memory Performance

Sleep Reports. Participants that directly incorporated the maze task into at least one dream report

(“Incorporators”, n=8) showed greater overnight improvement in both Completion Time (172 vs. 2 sec,

t15=2.38, p=0.03, d=1.16; Fig3, left) and Distance Traveled (150 vs. -2 grid units, t15=2.27, p=0.04, d=1.10;

Fig3, right), than those without direct task incorporations (“Non-Incorporators”, n=9). Incorporators also

showed a trend toward decreased Backtracking relative to Non-Incorporators (14% vs. -3%, t15=2.04, p=0.06).

Waking Rest Reports. Participants whose post-training rest reports directly incorporated the maze task

also showed non-significantly greater improvement in Completion Time (112 vs. -17 sec, t14=1.49, p=0.16) and

Distance Traveled (71 vs. -17 grid units, t14=1.53, p=0.15), and a trend toward greater improvement in

Backtracking (9% vs. -7%, t14=1.83, p=0.09), than those without direct incorporations. The failure of these

comparisons to reach significance, however, appears to reflect the difference in improvement between subjects

who incorporated the task into one, as opposed to two, reports (Fig2).

[FIG2 HERE]

[FIG3 HERE]

Because incorporation rates were quite high during post-training rest, with a full 50% of post-training

rest reports directly related to the maze (Table 2), we were also able to examine how the amount of maze-

related content predicted subsequent overnight performance improvements. The number of post-training rest
 Dreaming and Memory 11

reports related to the maze predicted overnight improvements in navigation performance (Spearman’s rank

order correlations: Completion Time ρ=0.50, p=0.051; Distance Traveled ρ=0.53, p=0.04; Backtracking ρ=0.55,

p=0.03; Fig2). The n=3 individuals who incorporated direct references to the maze task into pre-training rest

reports did not improve significantly more than other participants (p-values for all three measures >0.17).

VMT Performance

Overall, performance improved substantially across trials (linear polynomial contrast for the decrease in

Completion Time across trials: F1,32=16.45, p=0.0003). Following training, performance change across a night

of sleep did not differ significantly from change across 15min in the Immediate Test group (Completion Time:

t31=1.62, p=0.11; Distance Traveled: t 31=1.25,p=0.22; Backtracking: t 31=0.58, p=0.57).

Baseline Performance Predicts Task-Related Dreaming

As in our prior study (Wamsley et al., 2010b), participants who incorporated the learning task into

dreaming performed more poorly during training than those who did not. In the current study, “Incorporators”

performed more poorly than “Non-Incorporators” on final training trial for Completion Time (439 vs. 212 sec,

t15=2.39, p=0.03), Distance Traveled (286 vs. 147 grid units, t15=2.27, p=0.04), and Backtracking (42% vs. 21%,

t15=2.10, p=0.05). In contrast, this baseline performance was not correlated with incorporation of the task into

post-training rest reports (Completion Time: p=0.59, Distance Traveled: p=0.55, Backtracking: p=0.63).

Because Incorporators showed both poor baseline performance and enhanced overnight improvement (which

were themselves strongly correlated: r17=0.92, p<0.001), we conducted an ANCOVA testing the effect of task-

related dreaming on overnight Completion Time improvement while controlling for baseline Completion Time

as a covariate. In this model, the effect of dream incorporation on overnight improvement was no longer

statistically significant (p=0.64).

Sleep Architecture
 Dreaming and Memory 12

On the experimental night, Incorporators slept an average of 51±19 (SEM) min longer than Non-

Incorporators (t14 =2.74 p=0.02), resulting from similar reductions in all sleep stages (Table 3). No significant

correlations were found between TST, N2, N3, or REM sleep amounts and post-sleep performance gains

(Completion Time: TST: r16=0.18, p=0.51; N2: r16=0.32, p=0.23; SWS: r16=-0.23, p=0.39; REM: r16=-0.24,

p=0.37). However, there was a significant correlation between overnight performance improvement and time

spent in N1 sleep (Completion Time: r16=0.67, p=0.005), which survives Bonferroni multiple-comparisons

correction. We therefore conducted an ANCOVA testing the effect of task-related dreaming on improvement in

overnight Completion Time while controlling for the amount of N1 sleep as a covariate. In this model, the

effect of dream incorporation on overnight improvement remained statistically significant (F1,14=6.36, p=0.03).

Discussion

In support of the hypothesis that dreaming reflects memory processing in the sleeping brain, here we

report that participants who incorporate a recent learning experience into their overnight dream content show

enhanced overnight performance improvement. Importantly, this observation exactly replicates the findings of

our previously published nap study (Wamsley et al., 2010b), using the same criteria for direct task

incorporation, the same learning task, and the same methods of assessing performance improvement. We also

confirm, for the first time, that dream reports can incorporate an experimental learning task in all phases of

sleep—sleep onset, NREM and REM.

These task-related dreams could reflect neural “replay” of the learned task. But even these “direct”

incorporations rarely resembled an exact replay or rehearsal of the maze, similar to what one might perform

when awake. Instead, participants described their dreams as being about or taking place in a maze, with the

details of the report differing from the original experience in almost every detail. Thus, it is difficult to interpret

these experiences as anything resembling a “rehearsal” of learned information, and it thus is unlikely that the

increased performance improvement is a result of conscious rehearsal of the route taken to reach the goal.

Instead, we suggest that maze-related dreams predict subsequent task performance because they are indicators
 Dreaming and Memory 13

that performance-relevant components of task memory are being reactivated in the sleeping brain, albeit outside

of the dream experience. It has been argued that because they fail to veridically reiterate prior experience,

dreams are unlikely to support a memory function (Hartmann, 2010; Vertes, 2004). However, even the

“reactivation” of memory networks described in the rodent literature is far from a faithful reiteration of

experience and can “replay” never-before-experienced paths in a learned spatial environment (Gupta et al.,

2010). Thus, the dream reports described here could function to link specific recent experience of the maze task

with subjects’ preexisting knowledge structures about the hedge mazes, coliseums, or rooms described in the

reports. Indeed, it has become increasingly clear that sleep can transform memory traces over time and brain

regions, allowing for the extraction of generalizations, and integration of new information with past experience

(Lewis and Durrant, 2011; Payne et al., 2009).

We also found that thinking of the task during post-training quiet rest predicted performance improvement

following the subsequent night of sleep. This seemingly contradicts our prior study, in which waking thoughts

of the maze were unrelated to performance following a 5hr waking delay in a control group (Wamsley et al.,

2010b). This could be attributed to the fact that in our previous study Wake subjects did not sleep before being

retested. Indeed, prior evidence suggests that task-related brain activity in humans during quiet rest may predict

task improvement after subsequent sleep, but not wake (Gregory et al., 2014; Murphy et al., 2018).

Although our model proposes that dreams are a reflection of memory processing in the sleeping brain,

alternative explanations of these data are possible. One alternative account is suggested by the fact that

participants who dreamed about the maze performed especially poorly on the task at baseline. This poor

baseline performance could point to a third variable driving both the incorporation of the task into dreaming and

enhanced overnight improvement. For example, the brains of participants who struggle with completing the task

prior to sleep may prioritize this task for consolidation during the night, in line with prior evidence that baseline

performance level mediates selection of memories for sleep-dependent consolidation (Tucker and Fishbein,

2008; Wilhelm et al., 2012). Meanwhile, participants who found the task challenging may have been more

likely to dream about the experience for other reasons, perhaps increased anxiety over their relatively poor
 Dreaming and Memory 14

initial performance. But the same explanation cannot hold true for incorporation of the task into waking

cognition; although the appearance of the learning task in quiet rest reports was also associated with enhanced

overnight improvement, poor baseline performance was not predictive of these incorporations.

It is not entirely clear why several other recent investigations have failed to detect an association

between task-related dreams and post-sleep memory improvement (Schredl and Erlacher, 2010; Stamm et al.,

2014; Wamsley et al., 2016). But in some of these studies the rate at which the learning task was incorporated

into dreaming was very low, which could have prohibited a confident test of the hypothesis that these

incorporations relate to subsequent memory performance (Schredl and Erlacher, 2010; Stamm et al., 2014;

Wamsley et al., 2016). Indeed, in one of these studies, only one subject reported a dream that directly

incorporated the learning task (Schredl and Erlacher, 2010), and in a second, only 7 task-related reports were

obtained (Wamsley et al., 2016). The characteristics of learning paradigms and participant samples that might

cause incorporation rates to vary so widely across studies remain unknown. At the same time, it is also likely

that the true effect size of the association between dreaming and memory performance is smaller than we

originally reported. Although the effect size seen here (d=1.16 for Completion Time) is still large, it is

considerably smaller than the value of d=2.2 reported in (Wamsley et al., 2010b), raising the possibility that

these relatively small sample-size studies were underpowered to detect this effect.

In summary, we have replicated the observation that dreaming of a recent learning experience is

associated with enhanced next-morning performance. These subjective reports provide unequivocal evidence

that the memory of the maze learning task was reactivated during post-training sleep. This observation, and

especially the correlation of these dreams with next-morning improvement, is consistent with reactivation-based

models of memory consolidation during sleep. Together with other evidence, these findings offer added

support for the claim that the activation of recent memory networks during sleep is associated with subsequent

performance improvement.
 Dreaming and Memory 15

Acknowledgements

We thank Stephanie Thompson and Andrew Olsen for assistance with data collection, and Nam Nguyen for

programming the Virtual Maze Task. Supported by NIH grants MH048832 and MH098171, and a Tom Slick

research award from the Mind Science Foundation. E.W. and R.S. designed the research, supervised data

collection, and wrote the manuscript. E.W. conducted the statistical analyses. Figures were prepared with the

assistance of Daniel’s XL Toolbox v7.2.13.

Disclosure Statement

Financial Disclosures: None.

Non-Financial Disclosures: None.

 Dreaming and Memory 16

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Figure Legends

Figure 1 – Experimental timeline.

Figure 2 – Quiet rest reports. Left: Task-related reports per subject during Pre-Training and Post-Training rest

periods. Right: Improvement in completion time (left axis, open circles) and in distance traveled (right axis,

solid circles) as a function of the number of task-related reports subjects gave during post-training quiet rest.

Error bars: +/- SEM.

Figure 3 – Task improvement and task-related dream content. Improvement in completion time (left) and

distance travelled (right) for subjects who did (“Incorporators”) or did not incorporate (“Non-Incorporators”)

the task into at least one dream report. Error bars: +/-SEM.
 Dreaming and Memory 19

Table 1 – Example Maze Task Incorporations

Task Incorporation Examples

Sleep Onset “. . . I can see the maze, and I think if we could swim above it, we could see everything.”

REM Sleep “. . . we were just like walking through it but it was like a formal maze . . . one of those outdoor like ones made out of like
hedges and bushes.”

Stage 2 Sleep “. . . dreamed I was like standing in the middle of a maze, waiting for my friend to find me. And I was waiting and waiting and
she just kept going around and around, calling my name . . .”

Wake “I was trying to map out the design of the maze and . . . I was thinking about, how much money I’m not going to get, which is
about ten, fifteen bucks, and that was about it.”
 Dreaming and Memory 20

Table 2 – Subjective Reports by Conscious State

N Elicited N Content N Task-Related

# % p
Pre-Training Rest 36 36 (100%) 4 11% 0.001*
Post-Training Rest 40 40 (100%) 20 50% 2 x 10-14*

Sleep Onset 188 170 (90%) 12 7% 0.01*

N2 Sleep 62 48 (77%) 1 2% 0.2
Morning N2 and REM‡ 21 16 (76%) 2 13% 0.002*
Control Reports 100 64 (64%) 0 0% -

Notes. P-values derived from chi-square tests comparing task incorporation in each state to the 0% incorporation
observed in the control report set. *: Incorporation of the learning task was significantly greater than in the Control
report set.; ‡50% of Morning reports were elicited upon awakening from N2, and the remaining 50% upon
awakening from REM. Of the 2 task-related morning reports, one was recalled from N2, and one was recalled
from REM sleep.
 Dreaming and Memory 21

Table 3 – Sleep and Demographic Characteristics of Incorporators and Non-Incorporators.

Incorporators Non-Incorporators I:NI p-value

mean SD mean SD %
Age (yrs) 19.3 1.4 20.0 1.9 104% .37
Sex (% male) 25% 22% .89
Baseline Completion Time 439 244 212 140 48% .03*

TST (min) 411.1 29.6 360.1 43.5 88% .02*

N1 (min) 36.5 13.0 26.3 10.5 72% .10
N2 (min) 228.6 31.5 205.7 28.2 90% .15
N3 (min) 69.2 26.0 62.1 19.8 90% .55
REM 76.8 17.1 66.1 23.5 86% .32
ESS 7.5 4.1 7.4 3.5 99% .98
SSS 3.0 0.5 2.7 0.5 90% .20

Notes. Sleep Log TST – mean total sleep time from pre-study sleep log; ESS – Epworth
Sleepiness Scale; SSS – Stanford Sleepiness Scale at training; Baseline Completion Time –
mean time to solve the maze (sec) in the final training trial. I:NI – mean value for Non-
Incorporators as percent of value for Incorporators. p-values are derived from independent
samples t-tests, with the exception of sex, which was tested using a Chi-Square test of
independence.