Slow wave sleep reinforces the consolidation of declarative memory

The latest trend of napping stations create the awareness of how important
sleep can improve each individual performance. For university students, it is
important to boost the academic performance and to achieve the aim of declarative
memory consolidation that need to be significantly improved. The hypothesis for this
paper is that slow wave sleep reinforces the consolidation of declarative memory. A
type of long term memory that based on facts and events which can consciously
recalled is defined as declarative memory. The encoding and short-term retrieval of it
depends primarily on the hippocampus. Overtime, retrieval of declarative memories
becomes independent of the hippocampus due to an allocation of data to neocortical
networks. Memory consolidation is the route that reinforces the new encoded
memory, which is normally fragile and likely to deteriorate, and integrates it into the
complex of long term memories. A connection between the neocortex and
hippocampus is significant during this consolidation and appears to be predominantly
regulated by 1 Hz measured using electroencephalographic (EEG). The wave in this
stage is called slow-wave sleep (SWS) which normally known as the stage three and
four of non-rapid eye movement sleep (NREM) and it can be reached after 35
minutes of sleep. Sharp-wave ripples are known to complement a memory replay of
consolidated information in the hippocampus during slow-wave sleep which excites
the transfer of this memory-related information to the neocortex. This process is
understood to be substantial to the long-term storage of particular memories within
neocortical networks.
To support the hypothesis numerous articles have been analyzed carefully.
The search engine that being used was Mahidol University database which can
legally access to countless journal database such as PubMed and Sciencedirect.

To narrow down and find the topic of interest, the search words that were being used
were slow wave sleep, declarative memory, long-term memory, consolidation of
memory, napping, and performance. To support the hypothesis the first article which
was written by Plihal and Born in 1997 is chosen because the goal of the paper was
to distinguish between the role of slow wave sleep (light sleep) and rapid eye
movement sleep (deep sleep) on the memory consolidation. To further see the role
of SWS, the second article published by Neurobiology of Learning and Memory was
selected in order to distinguish the role between the short nap (10 minutes) which
contain no SWS and the longer nap (60 minutes). These two papers are essential for
comparing the percentage of SWS sleep and comparing the result, however they fail
to take into account that the delay between learning and sleeping might contains
some potential effect on the percentage of slow wave sleep. In addition, to recognize
the suitable delayed time to sleep after learning, the last article had been chosen.
In the article of Plihal and Born, the participants acquired to complete the
paired-associate word lists design. The researchers recruited 10 healthy men for this
study which were the psychology students from the University of Bamberg. The day
prior to the study the participants spent one night at the facility and were awakened
at 0700 h. The task consists of 24 pairs of German noun words, one is the stimulus
word and other is the response word. During the testing period the participants
required to answer the correct response word after the stimulus word had
displayed. On night A, the participants started learning the task at 2215 h to 2300 h
then they were tested at the end of learning period by completing the response word
that match with stimulus. After 3 hours of sleep, they were awakened and had 15
minutes to adapt to the surrounding which later on they were given the second test.
On night B, they had a similar procedure, however, they started learning the task

after had a 3 hour of sleep at the same time as the night A (Figure 1). The outcome
being measured is the percent improvement between the learning and recall
conditions. The improvement was measured by the number of trials until the
correctness reach 60% and the amount of correct words during the last trials. From
the polysomnography sleep data, it indicated that during the early nocturnal sleep
the SWS% was significantly higher compared to the late nocturnal sleep (Table 1).
Additionally, the recall of declarative paired-associate word lists was also significantly
greater after a period of SWS-rich period (Table 2).
Figure 1. Study design. On night A, the paired-associate list task was presented
between 2215 and 2300 h (learning). After learning, a 3-h retention interval followed.
At 0215 h, cued recall of the list and performance on the mirror-tracing task were
tested. On night B, all subjects slept from 2300 to 0200 h. fifteen min after
awakening, the learning tasks was presented. The retention interval lasted from
0300 to 0600 h. Fifteen min later, cued recall of the word list was tested between
0615 and 0700 h.

Table 1. Sleep parameters. Values are means SEM. Sleep onset (with reference to
2300 h), sleep time (i.e., time from sleep onset until the end of the respective
retention interval), percentage of awake time (Wake), stage 1 sleep (S1), stage 2
sleep (S2), slow wave sleep (SWS), and rapid eye movement sleep (REM).

Table 2. Performance on paired associate lists. Values are means SEM. In the
learning condition, presentation of a 24 paired associated was immediately followed
by a cued recall. This procedure was repeated until the subject reached the criterion
of at least 15 correctly recalled associates (criterion trail). The recall condition began
with a cued recalled in the learning and the recall condition. Percentage of
improvement refers to improvement with recall on the criterion trail of the learning
condition set to 100%.

The second article compared between the 10-minute nap group (Stage 1 and
2), the 60-minute nap group (SWS), and awake group which is the control group.
There were 36 participants in total which after learning the task the participants
assigned to each of the three group equally. A bimodal paired-associates task
contained of 36 two syllable nouns pairing with the 36 two second sound clips.
During the testing period, the two seconds sound clip would be presented and the
participants needed to correctly respond to the correct word pairs. The
measurements were collected from the baseline performance immediately after
training, after a nap or no nap for short term retention, after the inference task, and
one week after for the long term retention (Figure 2). The SWS recorded by the

electroencephalogram (EEG) show that during 10-minute nap there was no SWS
sleep compared to the 60-minute nap (Table 3). Even though every groups learned
the tasks similarly, both nap groups performed better than the wake group when
comparing the scores on the short-term retention, 1.5 hour after training. However,
performance benefits seen in the 10-minute nap group showed to be impermanent.
Performance after the interference task shown significantly higher memory retention
in the 60-minute nap group, with interference disrupting the memory trace far less
than both the wake and 10-minute nap groups. After a weeklong retention time,
sleeps benefit to memory continued in the 60-minute nap group, but not the other
two groups (Table 4).
Figure 2. Summary of experimental protocol: Two experimental groups and the
wake control group were all trained at 12 pm on the declarative bimodal pairedassociates task to a criterion level of 75%. After the training phase, the nap groups
immediately slept, either for a short 10-min nap or a longer 60-min nap, while the
control group remained awake. At 2:30, all groups were tested by cued recall and
then immediately trained on the interference task. 10 min after training, subjects
were tested by cued recall on both original and interfering pairs. One week after
initial learning, subjects were again tested on remaining learned pairs.

Table 3. Summary of sleep parameters across nap groups (mean SEM, in

minutes).

Table 4. Summary of sleep parameters across nap groups (mean SEM, in

minutes).

The third article examined the correlation between the delay of nap time and
the consolidation of declarative memory. There were 36 participants from City
College of the City University of New York. Firstly, the participants learned the tasks
by involving 150 neutral pictures. Following the picture presentation, the participants
were test on the ability to recognize whether the pictures were old (150 pictures) or
the new one. Then the participants were divided equally into one of four groups; 90minute nap immediately, 2 hours, 4 hours after learning, or stayed awake. After initial
learning (6 hours), participants were again tested on the ability to distinguish

between the old and new pictures (Figure 3). The result illustrated a significant
different between every group performance from test to retest. However, the 4-hour
delay group performed expressively better than those who slept immediately and
those who remained awake (Table 5). Analysis of sleep data discovered a positive
correlation between amounts of slow wave sleep (SWS) attained and time of the
delay before sleep onset (Table 6).

Figure 3. Description of Experimental Protocol. Experimental Protocol: Three

experimental groups and the wake control group were all trained at 11am on the
declarative memory recognition task, followed immediately by a testing session on a
portion of the previously viewed stimuli intermixed with new pictures. After the test
phase, sleep groups napped at staggered intervals, either immediately after testing
at 12 noon, at 2pm, or at 4pm, while the control group remained awake. All groups
were then retested at 6pm on the remaining stimuli, again intermixed with new
pictures.

Table 5. Percentage of correctly recognized previously viewed pictures at Test and

Retest (mean SEM).

Table 6. Summary of sleep parameters across nap groups (mean SEM, in

minutes).

All of these articles support the hypothesis that the higher percentage of SWS
during sleeping period is positively correlate with the consolidation of declarative
memory by observing the correctively recall tasks. From the first article, the data
from polysomnographic shown that the SWS percentage of early retention interval
was about five times higher comparing to late retention interval (Table 1) and the
percent improvement of the recall on early retention was about 22 percent higher
(Table 2). Thus, there is potential relationship between the SWS percentage and the
percent improvement. Alternatively, the percent improvement of paired-associate
word lists might address a disruptive influence of REM sleep on memory. Consistent
with the distribution of REM sleep perceived under typical undisturbed sleeping
circumstances, time spent in REM sleep was more than thrice during the late sleep
retention interval than during the early one (Table 1). Hence, the disruptive effect of
REM sleep can also be potential cause for improvement of memory consolidation.
However, from the data comparing between the 10-min nap, 60-min nap, and control
wake group it eliminated the ambiguity about REM disruptive influence because the

data shown that in the 10-min nap there was no time spent in either the REM or
SWS period (Table 3). It is crystal clear from the Table 4 data that the performance of
participants of 60-minute nap shown a significant improvement in every test session:
after the sleep period or control wake period, after the interference task, and after a
week of learning. The result also suggest that the latency stage and stage two alone
was insufficient for protection from interference and long-term retention. Still, the
reasons for temporary preservation of 10-min nap group it remained unclear. It might
due to the fact that the environmental influence during the 10-mins was omitted, the
more relax stage during learning the interference task, or because the process in the
late sleep period (SWS) is not yet active.
Slow wave sleep help declarative memory retention compared to an
equivalent time spent awake, although it is clear from the results that the act of
sleeping, alone, is not enough to account for the differences in retention. From Table
5 superior performance on the recognition task was seen in the longer participants
remained awake before nap onset (4-hour). In addition this nap group also contain
more time spent in SWS period. The authors of this paper explain the potential
causes for this surprising data that it occurs by the effect of circadian rhythm and the
homeostasis need for SWS.
This field of research is difficult to eradicate all the limitations. Since it
associates with human brain functions, there are differences in the ability to
memorize among each participant. Even though all of these research try to minimize
by having the first test after learning which used as a baseline to determine the
improvement after the sleeping period, there can be potential difference of memory
ability between each group of participants which might manipulate the result. The
next thing to be considered is the sample size of the researches which is relatively

small compared to other field of research. The mainly reason is due to the fact that
every participant need to wear the polysomnography or electroencephalogram
(EEG) during the research which will be enormously expensive to conducting the
research. Noted that the second and third articles were written by the same
researchers and conducted in the same facilities, so the procedural had a minimal
fallacy as they already know their limitation and try to obligate it. There might be
other factors that alter the SWS percentage rather than the delay time after the
learning period, which havent been discuss in this paper.
In conclusion, every research support the hypothesis that SWS improves the
consolidation of declarative memory.