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YourMemoriesNeedTheirSleep

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Your Memories Need Their Sleep

by Jim Schnabel
August 7, 2012

BRIEFING PAPER
Ann Whitman
(212) 223-4040
awhitman@dana.org

The big test is tomorrowshould you stay up late and study, or cut short the cram session and get a good
nights sleep? Most if not all students face this dilemma at some point in their lives. Until very recently,
their choice might have seemed obvious: stay up and study, to commit as much information to memory as
possible. But research now indicates that missing sleep in order to study may well be self-defeating. A
good nights sleep helps greatlyand is essential in some casesto making just-learned information
consolidate or stick in memory.
This issue doesnt affect only students or business people whose performance depends heavily on
memorization. Older peoples lives are often complicated by a where did I put my car keys? impairment
in memory function, and it now seems that aging-related sleep problems may cause some of this cognitive
decline.
The good news is that researchers may soon find ways to counteract this process, by helping people to
sleep better and by developing techniques to strengthen specific memories during sleep. We now know
that its possible to cue, during sleep, the reactivation of certain memories that were just learned or trained,
and this has the effect of enhancing the ability to recall those memories on subsequent days, says Jan
Born, Ph.D., a professor of medical psychology at the University of Tbingen in Germany.
A recent awakening
There have long been tantalizing hints of the relationship between sleep and memory. One of these was
the discovery in the 1950s of dream-rich rapid eye movement (REM) sleep, which eventually led to the
hypothesis that REM dreaming is the brains way of strengtheningby semi-consciously re-experiencing
new memories. People thought that it was a rehearsal of daytime experience, says J. Allan Hobson, M.D.,
professor of psychiatry emeritus at Harvard Medical School, and a member of the Dana Alliance for Brain
Initiatives.
But testing the relationship between sleep and memory was and still is tricky. A researcher cant easily
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disentangle the direct effect of a good nights sleep on memory consolidation from the indirect effect of
improved alertness and recall performance. There are also different types of memory, such as procedural
memory, which underlies physical skills, and declarative memory of events, dates, and places. They are
stored in different brain regions, and are not consolidated in the same way during sleep.
Sleep itself is not a uniform process. Brain activity in REM sleep differs from that within non-REM sleep,
which in its deepest stages is called slow-wave sleep (SWS). The initial scientific emphasis on REM sleep as
the principal window for memory consolidation produced inconsistent results and controversy.[1] The
recent focus on SWS has led to a more consistent story about memory consolidation, says Ken Paller,
Ph.D., a professor of psychology at Northwestern University.
The scientific shift towards SWS as the major memory-consolidation phase began in the mid-1990s with
experiments in animals, the results of which were later confirmed in human subjects. For example, Borns
laboratory reported in 2000 that people who had practiced an image discrimination taska visual skill
requiring procedural memoryimproved after a period of mostly SWS sleep, but not after the same
duration of mostly REM sleep. The experiment was one of the first to control for some of the confounding
effects of sleep loss.[2]
In a different study, Robert Stickgold, Ph.D., a postdoctoral researcher in Hobsons laboratory at Harvard
Medical School, found that subjects given a similar image discrimination challenge did not improve with
repeated testing unless they slept on the first night after their initial testing session. Stickgold also was able
to control for lost alertness by comparing subjects who slept for three nights after the first test with those
who stayed awake the first night but slept on the next two nights and recovered to normal alertness.[3]
Subsequent experiments in the mid and late 2000s indicated that SWS strengthens not only procedural
memories but also place memorieslearned routes within a townas well as declarative verbal memories,
such as homework-type memories of historical facts. Overall, the degree of memory consolidation seems
to depend on factors such as the duration and type of sleep (REM sleep having little or no effect), the type
of memory (declarative memories being helped most consistently) and the importance of the memory in
the life of the person.[4] The memory benefits of sleep also depend on age, because the amount of SWS
in a typical nights sleep declines, on average, after people reach their thirties.[5] By age 50 your memory
consolidation capacity is probably significantly worse than when you are 30, says Born.
Age-related sleep problems may even contribute to the normal cognitive decline seen among the
elderly. Theres a substantial amount of evidence that fragmented or inadequate sleep in elderly people
makes their cognitive function worse, says Dana Alliance member Clifford B. Saper, M.D., Ph.D., a
professor of neurology and neuroscience at Harvard Medical School, head of the neurology department at
Beth Israel Deaconess Medical Center.[6] Saper adds that sleep apnea, a breathing-interruption condition
which disturbs sleep patterns and is found more commonly among elderly and obese people, is the
leading treatable cause of dementia in his clinic. If we cure their sleep apnea, their memory usually comes
back and they feel better than they have in years, he says.
How does memory consolidation work?
An event happens, and you witness it. You hear it; or see it; or read it; or taste it; or feel it; or perhaps
engage all these senses at once. These seemingly simple acts of perception are brought about by millions
of your neurons firing in highly coordinated ways: neurons that correspond to the sensory features of the
things you perceive, the semantic meanings of these things, the place and time, even how you feel.
Neuroscientists widely assume that a memory of such an event is a set of connections among all these
neuronsconnections that bind them into a functional group, so that the reactivation of one can lead to
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the reactivation of the whole group, and thus a subjective re-experience of the event.
Although perception occurs within the very large and lately evolved structure known as the neocortex, the
initial neuronal connections that make a memory are formed chiefly elsewhere. For a declarative, whowhat-where memoryalso called an episodic memorythe initial connections are thought to be made in
the hippocampus, a seahorse-shaped appendage that is wired extensively into the neocortex.
The hippocampus works a bit like an old-fashioned telephone switchboard, making quick but temporary
connections among the millions of nerve channels that enter it from the cortex and other regions. In
animals and people, destruction of the hippocampus removes the ability to form new declarative
memories. We think that the hippocampus essentially links the separate features of an episode together,
says Born.
During SWS, these hippocampal links come alive again, apparently to consolidate the new memory.
Experiments on rats who learn their way through mazes indicate that the same hippocampal neurons that
are active during the learning of a maze are reactivated during the next period of SWS. With a similar study
in humans, Pierre Maquet, D.M., Ph.D., and colleagues at the University of Lige in Belgium have shown
that the degree of this hippocampal reactivation during SWS correlates with improvements in the
subsequent recall of the newly learned place memories.[7]
One likely effect of this hippocampus-initiated memory reactivation is to strengthen the new hippocampal
connections. But its more important goal seems to be the stimulation of slower-growing connections
among memory-associated neurons outside the hippocampus, mainly in the neocortex. These slowergrowing connections are more direct and permanent, and correspond to what we usually think of as long
term memory. During a period of weeks to months they form and strengthen, until the temporary
connections in the hippocampus are no longer needed. The longer-term neocortical connections are less
prone to interference, compared to hippocampal connections, and we suspect that they are better
connected to prefrontal structures that regulate memory retrieval, says Born.
What determines whether a new memory is consolidated during SWS? There is evidence, Born says, that
the highly evolved prefrontal cortex and an evolutionarily older, emotion-related structure called the
amygdala do this by chemically tagging new memory-associated links in the hippocampus, based on
their apparent logical or emotional importance.
Why does this memory consolidation process occur principally during SWS, rather than in REM sleep or
wakefulness? Born points to a number of key brain chemistry changes that occur during SWS, including a
sharp drop in the arousal-related neurotransmitter acetylcholine. This has the effect of switching the
normal direction of communication between the neocortex and hippocampus, so that now the
hippocampus can send information to the neocortex, he says. REM sleeps function here is still a matter of
debate, but Born suspects that it further exercises and strengthens newly formed neocortical connections,
albeit locally, without input from the hippocampus. [8]
Finally, what about procedural memories of sensory-and-motor skills, such as how to ride a bicycle? Less is
known about the ways in which these are initially stored and consolidated, but the muscle-controlling
structure known as the striatum appears to play a major role in their long-term storage. A related brain
structure callled the cerebellum could have a short-term storage role for procedural memories, analogous
to that of the hippocampus for declarative memories, says Born, but for now that is very speculative.
Enhancing consolidation
Pharmaceutical companies now are developing drugs to help enforce more normal, youthful sleep patterns
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in older people. Such drugs may have a significant effect in reversing age-related declines in memory
function. But neuroscientistsmost visibly Borns and Pallers laboratorieshave also investigated non-drug
methods to enhance sleep-based memory consolidation.
In 2007, Born and his colleagues reported an experiment in human subjects in which they strengthened
the recall of new declarative, hippocampus-based memories by cueing the reactivation of the memories
during SWS. The cueing was done with a floral odor that had been associated with the memory during
learning. [9] Pallers group recently reported similar effects for a largely procedural memorya keyboardplaying task requiring visual-motor coordinationusing the learned melody as the cue during sleep.[10]
Turning these simple experimental protocols into methods that ordinary people can use to enhance
memorization may not be easy. Some cueing methods may disturb peoples sleep, thereby spoiling the
effect. Others, such as odors, may not be useful for cueing highly specific and complex memories, says
Born. There is some evidence, too, he adds, that consolidation-enhancement with cues during sleep
involves a tradeoff, in which specifically cued memories are strengthened but other newly learned
memories are weakened, possibly reflecting a strict overall budget for memory consolidation every night.
Perhaps most importantly, no one has yet reported the enhancement of memory consolidation for the
verbal memories that dominate school and business learning, and that might require specific verbal cues
during SWS. Thats one of the next questions we have to ask: can you improve your verbal score on the
SAT, for example? says Paller.
He is cautiously optimistic. And certainly the demand for memory enhancement is there. I think its just a
matter of time before people are using such methods, Born saysand maybe also misusing them, since
people when they sleep cant entirely control the cues to which their brains are exposed.
Published August 2012

References:
[1] The REM Sleep-Memory Consolidation Hypothesis, by Jerome M. Siegel, Science 2001 (vol. 294,
November): 1058-1063.
[2] Early sleep triggers memory for early visual discrimination skills, by Steffen Gais et al, Nature
Neuroscience 2000 (vol. 3, December): 1335-1339.
[3] Visual discrimination learning requires sleep after training, by Robert Stickgold, LaTanya James and J.
Allan Hobson, Nature Neuroscience 2000 (vol. 3, December): 1237-1238.
[4] The whats and whens of sleep-dependent memory consolidation, by Susanne Diekelmann, Ines
Wilhelm and Jan Born, Sleep Med. Rev. 2009 (October): 309-321.
[5] Midlife decline in declarative memory consolidation is correlated with a decline in slow wave sleep,
by Jutta Backhaus et al, Learning and Memory 2007 (vol. 14):336-341.
[6] Increased fragmentation of rest-activity patterns is associated with a characteristic pattern of cognitive
impairment in older individuals, by Andrew S. Lim et al, Sleep 2012 (May 1): 633-640.
[7] Are Spatial Memories Strengthened in the Human Hippocampus during Slow Wave Sleep? by
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Philippe Peigneux et al, Neuron 2004 (vol. 44, Oct 28): 535545.
[8] The memory function of sleep, Nature Reviews Neuroscience 2010 (vol. 11, Feb): 114-26.
[9] Odor cues during slow-wave sleep prompt declarative memory consolidation, by Bjrn Raschet al,
Science 2007 (vol. 315, Mar 9):1426-9.
[10] Cued memory reactivation during sleep influences skill learning, by James W. Antony et al, Nature
Neuroscience 2012 (pub. online 24 June).

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