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Sigma Xi, The Scientific Research Society
Sleep to Remember: The brain needs sleep before and after learning new things, regardless of
the type of memory. Naps can help, but caffeine isn't an effective substitute
Author(s): Matthew P. Walker
Source: American Scientist, Vol. 94, No. 4 (JULY-AUGUST 2006), pp. 326-333
Published by: Sigma Xi, The Scientific Research Society
Stable URL: http://www.jstor.org/stable/27858801 .
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Matthew P.Walker
being a student? Faced cies that arise in complex fields of study, filing and retrieving information from
with a big test the next day, you
Remember thepreponderance of behavioral, neuro theneocortex, and italso seems to bind
have to learn something in a hurry?the anatomical, physiological, cellular and together different perceptual elements
General Prologue to Chaucer's Canter molecular evidence supports the idea of a single event ("Ah, yes. I put the
bury Tales, maybe,
or two-and-a-half oc that periods of the sleep cycle actively keys down tonab the last doughnut").
taves of a difficult scale on the clarinet. orchestrate changes in certain categories In contrast, nondeclarative memory
After long hours of practice you notice of memory. is the "know how" memory, rather
it's themiddle of the night. You haven't than the "know what" memory, and is
quite mastered the task. Should you Sleep, Memory manifest as an action or behavior. The
forgo sleep formore practice? The electrical signature of a sleeping category is divided further into proce
Depending on the stakes, most peo brain is different from that of an awake dural and implicitmemory. The former is
ple would say yes. But based on how brain, but equally big differences exist responsible formovements, habits and
our brains work, the answer is
prob during thevarious phases of sleep. Rapid skills (such as how to ride a bicycle);
ably no. In fact,psychologists have sus eye movement (REM) sleep, associated the latterencompasses less familiarphe
pected for some time that sleep relates with themost vivid dreaming, produces nomena such as classical
conditioning,
somehow to the development ofmem brain waves (measured with an electro habituation and priming (also known
ory, although the reasons were unclear. encephalograph) thatmost resemble the as the power of
suggestion). These dif
Behavioral tests showed that adequate ones found in awake subjects.Across the ferent forms of nondeclarative memory
sleep before and after a training session night, REM sleep alternateswith its an depend on somewhat differentbrain re
was essential for learning,whether the tithesis,non-REM (NREM) sleep, about gions. Although a map of these regions
taskwas tennis or algebra. every 90minutes inhumans. In primates must be overly simplistic, neuroscien
Scientists frommany disciplines have (including human beings), NREM sleep tistsconsider the core structures forpro
confirmed and elaborated those suspi has four substages. Psychologists refer cedural memory tobe the striatum,mo
cions over the past decade. In view of to the deepest and most electrically dis tor cortex and cerebellum; conditioning
the collective illumination ofmuch con tinctiveof these substages, NREM 3 and engages the cerebellum and, for emo
gruent data, most neuroscientists now NREM 4, as slow-wave sleep (SWS) be tional learning, the amygdala; priming
believe that sleep is integral to learning cause of the characteristic low-frequency, involves theneocortex; and habituation
and memory. However, some subtle
high-amplitude brain waves. isbased on reflexpathways in the spinal
ties lie beneath this blanket statement: Similar to sleep, memory exists in cord and brainstem.
Neither memory nor sleep is simple in several forms. The most popular clas Although these categories are con
terms of structure and function.What's sification scheme is based on the dis veniently separate on paper, real-life
more, the intersections between types tinction between those memories that memories are rarely so distinct. For
ofmemory and phases of sleep are also you can declare verbally and those that example, learning to speak a language
governed by sometimes cryptic vari you have to show throughperformance.
ables. Despite the inevitable discrepan Psychologists call these categories declar Figure 1. Sleep is critical to cognitive functions,
ative and nondeclarative particularly memory. Even during highly cho
memory, respec
P. Walker earned his Ph.D. from the
tively.The former is fact-based and in
Matthew reographed missions aboard the space shuttle,
Medical Research Council in London in 1999. Fol astronauts are instructed to sleep eight hours
cludes answers to questions like "What
lowing stints as a postdoctoral fellow and instructor is the value of Planck's constant?" and per night?a difficult feat given the excitement
in psychiatry and psychology atHarvard Medical and weightlessness. Recent studies demon
"Where did I put my keys?" The an
School, he became an assistant professor ofpsychol strate that learning requires physical changes
nals of neuropathology, brain imaging
ogy in 2004. He is currently director of the Sleep in the brain, at least some of which occur dur
and Neuroimaging Laboratory. Address: Sleep and
and modern computer models agree of the sleep cycle. Here,
ing specific phases
thatdeclarative memory requires a part Mission Guion Bluford
Neuroimaging Laboratory, Department ofPsychia Specialist (left) and
try,Beth Israel Deaconess Medical Center, Feldberg,
of the brain called the hippocampus, Commander Richard "Dick" Truly (right) doze
FD 862, Harvard Medical School, Boston, MA which lieswithin themedial temporal while floating on themiddeck of the Challeng
02115. Internet: mwalker@hms.harvard.edu lobe. This little structure is a nexus for er in 1983.
(Photograph courtesy of NASA.)
nonREMI
MM
?
Figure 2. During sleep, a person cycles through periods of electrically distinct brain activity. Rapid eye movement (REM) and non-REM (NREM)
sleep alternate about every 90 minutes, the ratio of NREM to REM sleep shifts as the progresses (as shown here for a person falling
although night
asleep atmidnight). NREM stages 3 and 4,which are characterized by high-amplitude, low-frequency waves, predominate during the firsthalf of the
but stage 2 NREM and REM are more common later.
night, sleep
requires several memory modes, rang on the passage from the firstmental If that encoded memory is destined for
ing from nondeclarative memory for glimmer to a permanent record. The long-term storage, itwill go through
how tomove themouth and tongue, steps occur on a continuum, although successive
stages
to become more sta
tomemory of grammatical rules and the exact timing is variable by task, ble in a process known as consolida
structure (a mix of the conscious and strength of the initialmemory, circum tion. In classical psychology, a memory
unconscious), todeclarative memory for stance and individual. Thus, making is consolidated when, in the absence
vocabulary.
someone's
acquaintance forms an of mental rehearsal, it becomes stur
But regardless of type, all memory ephemeral representation of that per dy enough to resist disruption from
appears to go through similar stages son's name and facewithin the brain. competing new learning, perceptions,
thoughts or actions.
Recent findings show that consolida
declarative nondeclarative
tion goes beyond simply stabilizing or
memory memory
fixatingmemories?it enhances them
as well. The two processes seem to be
distinct: Although stabilization appears
to occur over time regardless of brain
state, enhancement occurs primarily, if
not exclusively, during sleep. This "of
fline" effect can restore previously lost
memories or produce additional learn
ing, both without the need for further
practice. In other words, the enhance
ment phase of memory consolidation
is an active process, not merely one of
simple maintenance; the brain contin
ues to learn even though ithas stopped
practicing.
I've chosen in this article to focus
primarily on the influence of sleep on
encoding and consolidation, but the
later stages of memory processing are
also important. In them, new patterns
Figure 3. Human memory can be classified several ways. Most schemes distinguish between de of information are integratedwith past
clarative and nondeclarative memory. The former is consciously accessible and fact-based (know
experiences and knowledge. At about
ing what), and includes general knowledge (semantic) and autobiographical memory (episodic). the same time,memories can be reor
In the brain, these categories require the diencephalon and medial temporal lobe, including the
Nondeclarative memory to our conscious mind and includes proce
is inaccessible ganized and moved to new anatomical
hippocampus.
sites in a process called translocation.
dural memory how), as well as implicit forms of learning,
for actions, habits and skills (knowing
on various other parts of the brain. For For declarative memories, thismeans
which depend example, learning to swing a golf club
that thememory trace is no longer ex
requires the striatum, motor cortex and cerebellum; nonassociative learning involves reflex path
ways in the spinal cord and elsewhere; priming engages the neocortex; and associative learning
clusive to the hippocampus, but be
requires the amygdala or cerebellum, depending on whether the cued behavior has an emotional comes more distributed through por
or a physical component. (Functional anatomy after Squire and Knowlton 1994.) tions of the cortex.
In Praise ofNaps
One twist in this story is thatdespite the
clear importance of nightly sleep for full
memory enhancement, short daytime
naps yield surprisingly largebenefits.We
used thefinger-tapping task to compare
theperformance gains of two groups of
subjects:One had a 60- to 90-minutemid
Figure 5. Sleep enhances memory. The author and his colleagues trained subjects on a finger
day nap afteramorning training session,
tapping task (a form of procedural memory) in the morning. All participants improved with and the other did not. Later the same
practice. Later that day, the author tested half the subjects and found that their performance day, the group thathad napped was sig
had not changed (a, green bar at center). However, the other half, which the author tested after
nificantly (about 16 percent) better at per
normal sleep, improved (b, purple bar at center). A night of sleep subsequently these sequences than theyhad
significantly
enhanced thememory of the first group (a, purple bar), but the second group did not continue to forming
been thatmorning. In contrast, and as
get better following another 12 hours awake (b, purple bar at right). Asterisks indicate significant
we noted in theearlier study,people who
differences between initial and later testing; error bars represent standard error.
did not nap and remained awake across
Avi Kami, Dov Sagi and coworkers provements the next day. Subsequently, theday did not improve.
at theWeizmann Institute of Science Robert Stickgold,my frequent collabora Interestingly, when we tested both
in Rehovot, Israel, demonstrated that tor atHarvard, showed that the degree of these groups (subjects who did or
subjects who learned todistinguish spe of improvement correlated positively did not nap) again the next day follow
cific details in a patterned image (a so with the time spent in REM sleep and ing a full night of sleep, those subjects
called visual-skill task) also improved SWS. SteffenGais and his colleagues in who had napped showed only a 7 per
their performance after sleep (but not JanBorn's research team at theUniver cent additional increase inperformance
after an equivalent period of wakeful sity of L?beck inGermany suggest that speed, resulting in a summed total im
ness). In that task, the enhancement ap SWS initiates consolidation, but subse provement of 23 percent. However, sub
peared to depend on REM sleep, since quent REM sleep promotes additional jectswho had not napped were nearly
subjects who were deprived of this type enhancement. Stickgold has since dem 24 percent fasterafter sleeping thenight.
of sleep across a night showed no im onstrated that if subjects are deprived Therefore, by the following day, both
groups had improved by approximately
- -
before awake interval after the same total amount. These data sug
gest theremay be a limit to how much
absolute improvement sleep can trigger
across a 24-hour period. Thus, amidday
nap changes the time course of when
that offline motor memory improve
ment occurs, but ultimately not the total
benefit, as the two groups improved by
the same amount after24 hours.
Daytime naps also appear to improve
the learning of a visual skill, although
key-press transition key-press transition the effectsare subtly different.Stickgold,
Sara C. Mednick and their colleagues at
Figure 6. Sleep selectively enhances themost difficult portions of a complex motor task. Subjects Harvard have shown thatwhen people
learned to tap quickly a sequence of five numbers on a keyboard (for example, 4-1-3-2-4), which
rehearse the visual-perceptual task (de
contained four unique key-press transitions: (A) 4 to 1, (B) 1 to 3, (C) 3 to 2, and (D) 2 to 4. Not all
scribed earlier) several times across the
transitions are equally hard: Some are particularly slow, as reflected in greater reaction times, and
therefore difficult (labeled as problem points). After training, some subjects slept while others day, theybegin to getworse rather than
remained awake. This figure shows representative performance data from separate individuals,
better. However, if the subjects take a
either before and after sleep (a) or before and after an equivalent of wakefulness (b). Fol 30- to 60-minute nap among these tests,
period
lowing sleep (a, purple line), subjects performed the task more quickly, but the improvement was
then thedeterioration halts. If they sleep
specific to the slowest, problem-point transition. By contrast, eight hours awake did not improve longer?60 to 90 minutes?then per
performance, and the transition profile remained unaltered (b, yellow line). formance not only stops declining, but
W??? daytime
napgroup W??? controlgroup (no nap)
ffiilP
liflli
Figure 8. Daytime naps can fillmuch the same role inmemory enhancement as a full
nights sleep, although adding
a nap to a
night's sleep doesn't
offer any greater benefit than a night of sleep alone. In this experiment, subjects practiced a motor skill task in themorning and either napped (60
to 90 minutes) or remained awake until retested later the same day, subjects who had napped
atmidday evening. When (purple) were significantly
faster (16 percent), but the performance of controls (red) was unchanged. After a full night of sleep, the performance speed of subjects in the nap
group only increased by an additional 7 percent, but the control group sped up by 24 percent. Therefore, within 24 hours, both groups averaged the
same total amount of
delayed learning. Several famously creative thinkers throughout history have been dedicated nappers, including Leonardo
Da Vinci, Salvador Dali, Buckminster Fuller and Thomas Edison (shown here asleep on a laboratory bench in 1911). Asterisks indicate significant
performance differences between training and testing. (Photograph courtesy of the National Park Service, Edison Historical Site.)
curred) and, interestingly,cripples their severely compromised in remembering spindle density. Journal of Neuroscience
22:6830-6834.
insight intohow well they actually per positive and, to a lesser extent, neutral
formed the task, according to a study by stimuli (almost a 60 percent decrement Kami, A., D. Tanne, B. S. Rubenstein, J. J.Aske
nasy and D. Sagi. 1994. Dependence on REM
Yvonne Harrison and JamesA. Home at forpositive associations). Furthermore, a
sleep of overnight improvement of percep
Loughborough University in theUnited negative emotional memories were tual skill. Science 265:679-682.
Kingdom. This bodes ill for the success somewhat more resistant to the effects Maquet, P., et al 2000. Experience-dependent
of people who need to acquire and ana of sleep deprivation; the groups were changes in cerebral activation during human
REM sleep. Nature Neuroscience 3:831-836.
lyze new information while sleep de not significantly different in their recol
S. C., K. Nakayama, J.L. Cantero, M.
prived, such as physicians or soldiers. lection of negative word pairs. Mednick,
Atienza, A. A. Levin, N. Pathak and R. Stick
They may think that they're doing just The take-home message from these on
gold. 2002. The restorative effect of naps
fine on little sleep, but because their studies is that sleep before learning is
perceptual deterioration. Nature Neuroscience
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Cognitive Neurosciences, ed. M. S. Gazzaniga.
while increasing general alertness, did matically, resulting inmemories thatdo
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Stickgold, R., D. Whidbee, B. Schirmer, V. Patel
However, not all forms of declarative more, the impact of inadequate sleep and J.A. Hobson. 2000. Visual discrimina
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deprivation. My lab recently looked at information, specifically positive forms occurring during sleep. Journal of Cognitive
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memory consolidation and reconsolidation.
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thathad positive (happy, love, sunlight), Dreams Becoming Reality Walker, M. P., T. Brakefield, A. Morgan, J.A.
negative (cancer,grief, jail) or neutral as The publication rate in the field of sleep Hobson and R. Stickgold. 2002. Practice with
sociations, the participants slept as they and memory has doubled in each of the sleep makes perfect: sleep dependent motor
skill learning. Neuron 35:205-211.
wished fortwo nights and then returned; past two decades?a rate that eclipses
we surprised them at thatpoint with an the growth of research in either sleep
unexpected testofword recognition. or
memory alone. These reports, from
The data showed that subjects who cellular and molecular studies in ani
were sleep-deprived before learn mals to behavioral studies in humans, For relevant Web links, consult this
issue of American Scientist Online:
ing remembered 40 percent less than provide converging evidence that pre
controls?a striking impairment?but training sleep prepares the brain for
this deficit was not the same in each learning, and posttraining sleep trig http://www.americanscientist.org/
emotional category. Rested subjects gers memory consolidation through lssueTOC/issue/861
had better recall of positive and nega neural plasticity, leading to enhanced
tive stimuli than of neutral stimuli, a recall thenext day.