Beruflich Dokumente
Kultur Dokumente
Tel-Aviv University
School of Social Studies
Department of Psychology
By
Michal Eisenstein
Directed by
November 1999
Table of Contents
2
Abstract 3
Introduction 4
Animal Studies 6
Human Studies 13
Methodology 22
Subjects 22
Tools 22
Design
25
Procedure 26
Results
28
Sleep data 28
Priming data 29
Discussion 33
References 36
3
Abstract
Introduction
as time passes. The term consolidation may help clarify this seemingly
years, and even decades. That is, some memories become more
1996). This is not the type of consolidation that this study addresses.
storage (Nadel & Moscovitch, 1997; for a review, see McGaugh, 1966).
extensively in rats, mice, and other even simpler organisms, and have
free to work through the experiences of the day, discarding the trivial
sleep, our brains are working hard to save the experiences that will
remain with us. The goal of this study was to shed more light upon the
Animal studies
Neuro-chemical evidence
There is more than one reason to believe that post-training REM sleep
(Corkin, 1981; Coyle, Price & DeLong, 1983; Deutsch, 1983), and REM
Hobson, 1987; Gillin & Sitaram, 1984; Jouvet, 1975; McGinty & Drucker-
to gradually increase over normal levels at the same time that REM
sleep increases were observed in the shuttle avoidance task (in which
Smith, Tenn & Annett, 1991). Interference with normal ACh metabolism
levels of ACh and AChE. These drugs were ineffective when applied
either before or after this specific time period of REM sleep increases
processing would be active during REM sleep. And second, that the
with wakefulness (Llinas & Paré, 1991; Steriade, 1989): both are brain
(SWS - for review, see Steriade, 1991). Despite this, the ability of the
Behavioral evidence
8
The first line of evidence comes from findings showing that REM sleep
Leconte, 1979; Hennevin & Leconte, 1971; McGrath & Cohen, 1978;
Smith, 1985; Smith, Kitahama, Valatx & Jouvet, 1974; Smith & Lapp,
1986; Smith & Wong, 1991; Smith, Young & Young, 1980). Thus, REM
to the other side of the cage after a sound that precedes an electric
shock. While rats were learning this task, a significant increase was
found in the quantity of their REM sleep recorded after each daily
performed the task almost perfectly, the quantity of their REM sleep
not exhibit learning did not show such an increase in REM sleep
for other types of learning, such as lever pressing and maze learning
In addition, when sleep was delayed for three hours after each
their studies: “It would appear that one of the essential elements of
baseline and for the duration of the training situation. In these studies,
the levels of REM sleep were observed to persist for times well beyond
the first three post-training hours (Fishbein & Gutwein, 1977; Smith,
Conway & Rose, 1993; Smith et al., 1974; Smith, Lowe & Smith, 1977;
Smith & Wong, 1991; Smith et al., 1980). Thus, all these studies
showed that the REM sleep increases following the end of training
persisted longer than three hours and often had a latency to onset
after the end of training of more than three hours (for a detailed
periods of several days after the end of acquisition (Smith & Lapp,
REM sleep increases are quite prolonged, and may persist for as much
as six days after training. As for the latency to onset and duration, they
appear to be a function of the strain and type of animal, the task, and
1979; Fishbein & Gutwein, 1977; Hennevin & Leconte, 1971; McGrath &
Cohen, 1978; Pearlman, 1979; Smith, 1985; Smith & Butler, 1982;
Smith & Kelly, 1988; Smith & Lapp, 1986; Smith & MacNeill, 1993). In
studies of this type, rats are trained in a specific task and are deprived
depriving animals of REM sleep is the “inverted plant pot” (also “water
down in a basin of water leaving only a very slim surface above water.
The animal is placed on the surface for that period of time during
during waking and non-REM (NREM) sleep, yet as soon as they enter
the REM stage, they lose postural tone (due to the muscular paralysis
characteristic of REM sleep), and partially or fully slip from the pedestal
the end of the training session. It was usually assumed that REM sleep
training (for review, see Block et al., 1979; McGrath & Cohen, 1978;
In one such study, rats were trained and returned to their cages after
spending two to three hours on plant pots. When they were tested 24
who, after spending two hours undisturbed in their cages, were put on
the plant pots for the same period of time (Pearlman & Greenberg,
1973).
out on animals in which the post-training REM sleep changes had been
the REM sleep increases observed with the time after training that the
The REMW has been defined as a time after acquisition when there
are increases in REM sleep over normal levels. What's more, if REMD is
the REMW, the general pattern seems to be that, for a given task, the
more trials given in a single session, the shorter the latency to onset of
the first REMW (Smith, 1996). When a smaller number of trials per
REM sleep in the 24-h period just prior to an increase in actual correct
1980). These results suggest that the REM sleep changes can actually
In one study (Smith & Butler, 1982), during five days of shuttle
avoidance training, rats were allowed REM sleep only during the two
previously established REMW’s (9-12 h and 17-20 h after the end of the
last training trial; Smith & Butler, 1982; Smith et al., 1980), while REMD
controls.
impaired retention. With use of the inverted plant pot technique, stress
Human studies
conflicting results (Horne & McGrath, 1984; McGrath & Cohen, 1978).
14
increases in the amount of REM sleep (Allen, Oswald, Lewis & Tagney,
1972; Bowe-Anders, Herman & Roffwarg, 1974, Horne, 1976; Horne &
One study observed the sleep of students at times when they were
not taking any courses and again just after they had completed their
not taking exams, the test subjects exhibited increases in the number
after the end of their exams (Smith, 1993; Smith & Lapp, 1991).
Additional findings indicate that REM sleep does play a role in special
re-learning a large portion of the words they had forgotten, while the
followed by REMD and then subjects were tested for retention at some
West, 1971; Empson & Clarke, 1970; Lewin & Glaubman, 1975;
McGrath & Cohen, 1978; Smith, 1993). However, when the material to
Empson & Clarke, 1970; Grieser, greenberg & Harrison, 1972; McGrath
It has recently been reported that memory for a complex logic task
REMD, while memory for a paired associate task was not impaired
Smith, 1993; Smith & Whittaker, 1987). These data suggest that the
16
other words, the effects of REMD following the learning phase seem to
recall from previous events. These two types of memory are presumed
explicit and implicit tests of memory. The most striking of these, is that
possible that in humans, REM sleep is essential for only certain types of
17
learning and memory tasks, while not necessary for others, depending
next day. This improvement was also found in those participants who
were asked to recognize the embedded stimuli, and this was done
without reference to the prior learning episodes where this task had
be the important characteristic that defines its need for REM sleep to
and it is this aspect of the task that may be dependent on REM sleep
for consolidation1.
As for the times after acquisition when REM sleep is most important
resulted in memory loss when it occurred the same night or two nights
after acquisition (Smith, 1993; Smith & Whittaker, 1987). This result
REM sleep loss. However, there would not appear to be a 3-4-h REMW
suggest that on vulnerable nights, all of the REM periods of the night
necessary for all types of implicit memory tasks, or only for implicit
the role of REM sleep in an implicit memory task that does not test a
previously studied words than when not. For example, in the study
them and they remain unaware of the fact that they will subsequently
fragments of new, unstudied words, with the first word that comes to
some word fragments better than others. The same type of effect can
1993).
Research hypotheses
task was necessary to ensure that this is not the case, on condition
that performance on the control task would not differ following REM
always performed after the implicit memory task, when subjects would
Hence, it was hypothesized that: First, subjects’ sleep would not differ
between the two experimental nights, except for differences due to the
following the control night than the REMD night. And third, subjects’
performance on the control task would not differ between the control
sleep is critical not only for general skill learning, but also for item-
specific implicit memory. If, however, REM sleep does not affect
the implicit memory task would not differ between the two nights, or
control task would be worse following the REMD night, suggesting that
tests that are both implicit and procedural may prove to be the only
Methodology
Subjects
received credit for their participation in the experiment and were all
Peled & Lavie, 1985; see below). All participants spoke fluent Hebrew
Tools
using five channels: EEG, EOG and submental EMG. Data from all
Kales, 1968).
fragments (see appendix 1). The fragments were constructed such that
it would be possible to fill them correctly with more than one word, but
with the first word that came to mind, thus establishing baseline levels
target words were then randomly divided into two lists of equal length.
Study Phase: During the study phase, half of the participants received
the first list of targets, and the other half the second list of targets. On
and only after the subject has responded the next trial began. A scale
were not told that they would be tested on these targets the following
morning.
Test Phase: During the test phase, all participants received all 90
order. Each fragment remained on screen until a key was pressed, but
participants were asked to fill in the fragments (on paper) with the first
appendix 2). The fragments were constructed such that it would not be
possible to fill them in with more than one country name. Country
adjacent letters), size 48 David font. They were centered on the same
screen until a key was pressed, but no longer than thirty seconds.
Design
25
sleep lab, going through both sleep conditions after one night of
served as his/her own control. The first subject went through the first
night on Dec 28th, 1998, the last on May 22nd, 1999. The first
experimental night was performed on Jan 30th, 1999, the last on Sep
7th, 1999. The mean interval between the adaptation night and the first
mean interval between the first experimental night and the second
first REM deprived and then NREM deprived, while the other half were
each time they entered the REM stage and were kept awake for at
least 5 min to assure that they will not go immediately back to REM
sleep. During this time, subjects were requested to report any mental
during NREM sleep. They were kept awake for the same period of time,
and also asked to report mental content. In the case of the NREM
condition being performed initially, the subject was awakened for the
26
deprivation first.
task. The same subject cannot perform both explicit and implicit tasks
since performance of the implicit task after the explicit task would yield
between subjects yet due to time and budget limitations this was not
possible.
Procedure
best sleepers (7 best men and 8 best women) were chosen (mean
score=18;
SD=4.07) and were assigned three nights in the sleep lab (The
Institute for Fatigue and Sleep Medicine, “Sheba” Medical Center, Tel-
Hashomer, Israel).
presented with a list of words and asked to rate the extent to which the
During the second and third nights, participants performed the study
phase upon arrival at the lab. During the study phase, each subject
went to sleep. During the night, the sleep condition manipulation was
For all three nights, subjects chose their own sleep onset and offset
times.
For one subject, the PSG device failed to record his first experimental
night. For the second subject, the memory test program threw her out
Results
Sleep data
were scored for sleep stages. That is, stages 1-4 and REM sleep.
sleep latency (time from lights out to falling asleep in minutes; scored
subjects was awake for more than one minute after sleep onset),
awake for less than one minute after sleep onset), TST (total sleep
sleep, % SWS (stages 3-4), % REM sleep, and REM latency (in minutes).
REM sleep while awakening subjects for about the same number of
times during the two experimental nights. This can be seen by the
29
experimental nights.
discussion section.
Priming data
The control task was the first to be examined. The score on this task
test between the two experimental nights, the dependent variable was
been correctly completed only when the subject filled it in with the
summed to give two correct scores for each subject, for each of the
that this pattern was displayed by all of the subjects on both sleep
conditions, meaning that learning took place following all of the study
a greater priming score following the NREMD night than following the
REMD night. A sign test was carried out on these scores and turned out
learning has occurred. If, on the other hand, learning did take place, a
studied score that is greater that the unstudied score should manifest
this, and the difference between the two represents the “amount” of
Discussion
This study attempted to shed light on the role played by REM sleep in
depriving it of REM sleep (see Horne & McGrath). Therefore, the control
and for the same duration of time as in the REMD condition. According
the subjects for about the same number of times on the two
difficult to control, such as total sleep time (TST) and sleep efficiency.
As subjects were their own controls and all good sleepers, it was
awakened for about the same number of times for approximately five
is more “difficult” or takes more time, after being awakened from REM
sleep than from SWS. The fact is that on the NREM deprivation night
subjects had more sleep, and not much could be done about it. The
two sleep conditions on the implicit memory task? We think not. If this
the NREM deprivation night than the NREM deprivation night on the
The control task was designed to test whether the subjects would
p<.13). Note that performance on the control task following the night
This makes it safe to say that the inferior performance on the implicit
memory task following REM deprivation was not due to greater fatigue
following REM deprivation was also higher than that following NREM
deprivation, thus supporting the claim that subjects were not more
between the two nights explain the lower priming scores following REM
the more SWS subjects had - the worse they performed on the implicit
task. Such a claim would mean that SWS disrupts learning, and does
to note that REM deprivation does not totally abolish this process of
and that what is processed during REM sleep can be transferred to the
that the memory task employed in this study was an implicit memory
task.
sleep can modulate newly acquired memories are still not understood,
the span of time during which REM sleep can play a role and the nature
References
35
Allen SR, Oswald L, Lewis SA, Tagney J (1972). The effects of distorted
visual input on sleep. Psychophysiology 9:498-504.
Baghdoyan HA, Rodrigo-Angulo ML, Assens F, McCarley RW, Hobson JA
(1987). A neuroanatomical gradient in the pontine tegmentum for the
cholinoceptive induction of desynchronized sleep signs. Brain
Research 414:245-261.
Bloch V, Hennevin E, Leconte P (1979). Relationship between
paradoxical sleep and memory processes. In M. A. Brazier (Ed): Brain
Mechanisms in Memory and Learning: From the Single Neuron to
Man, vol. 4 (pp. 329-343). New-York: Raven.
Bowe-Anders C, Herman J, Roffwarg H (1974). Effects of goggle-altered
color perception on sleep. Perceptual and Motor Skills 38:191-198.
Cartwright RD, Lloyd S, Butters E, Weiner L, McCarthy L, Hancock J
(1975). Effects of REM time on what is recalled. Psychophysiology
12:561-568.
Castaldo V (1969). Down’s syndrome: a study of sleep patterns related
to level of mental retardation. American Journal of Mental Deficiency
74(2):187-190.
Cataldo V, Krynicki V, Goldstein J (1974). Sleep stages and verbal
memory. Perceptual and Motor Skills 39:1023-1030.
Chernik DA (1972). Effect of REM sleep deprivation on learning and
recall by humans. Perceptual and Motor Skills 34:283-294.
Cohen NJ, Eichenbaum H (1993). Memory, Amnesia, and the
Hippocampal System. MIT Press: Cambridge, MA.
Conway J, Smith C (1994). REM sleep and learning in humans: a
sensitivity to specific types of learning tasks. Journal of Sleep
Research 3(Suppl.1):48.
Corkin S (1981). Acetylcholine, aging and Alzheimer’s disease:
implications for treatment. Trends Neurosci. 4:287-290.
Coyle J, Price DL, DeLong M (1983). Alzheimer’s disease: a disorder of
cortical cholinergic inervation. Science 219:1184-1190.
36
Schacter DL (1996). Searching for Memory: The Brain, the Mind and
the Past (pp. 81-86). Basic Books: New York, NY.
Schacter DL, Tulving E (1994). Memory Systems 1994. MIT Press:
Cambridge, MA.
Seligman MEP (1970). On the generality of the laws of learning.
Psychological Reviews 77:406-418.
Sitaram N, Weingartner H, Gillin JC (1978). Human serial learning:
enhancement with arecholine and choline and impairment with
scopolamine. Science 201:274-276.
Smith CT (1985). Sleep states and learning: a review of the animal
literature. Neuroscience & Biobehavioral Reviews 9:157-168.
Smith C (1993). REM sleep and learning: some recent findings. In
Moffitt A, Kramer M, and Hoffmann R (Eds), The Functions of
Dreaming, (pp. 341-361). New York: SUNY.
Smith C (1996). Sleep states, memory processes and synaptic
plasticity. Behavioural Brain Research 78:49-56.
Smith C, Butler S (1982). Paradoxical sleep at selective times following
training is necessary for learning. Physiology and Behavior 29:469-
473.
Smith C, Conway J, Rose G (1993). Evidence of a paradoxical sleep
window for spatial memory. Sleep Research 22.
Smith C, Kelly G (1988). Paradoxical sleep deprivation applied two days
after the end of training retards learning. Physiology and Behavior
43:213-216.
Smith CT, Kitahama K, Valatx JL, Jouvet M (1974). Increased
paradoxical sleep in mice during acquisition of a shock avoidance
task. Brain Research 77:221-230.
Smith C, Lapp L (1986). Prolonged increases in both PS and number of
REMS following a shuttle avoidance task. Physiology and Behavior
36:1053-1057.
Smith C, Lapp L (1991). Increases in number of REMs and REM density
in humans following and intensive learning period. Sleep 14:325-330.
40