Lecture 6 – Wakefulness and

Sleep

Credit: https://people.cornellcollege.edu/dsherman/illusions/# (Drawing of retina from Hubel DH Eye, Brain, and

Vision Scientific American Library p.38)
 Outline
1. Rhythms of Waking and Sleep
2. Stages of Sleep
3. Functions of Sleep REM
4. Brain Mechanisms of Sleep
5. Sleep Disorders
 Learning Objectives
1. Define and describe endogenous rhythms.
2. Explain the mechanisms that set and reset the biological
clock.
3. List and characterize the stages of sleep.
4. Describe the brain mechanisms of waking and sleeping.
5. Discuss several consequences of thinking of sleep as a
localized phenomenon.
6. List several sleep disorders with their causes.
7. Evaluate possible explanations of the functions of sleep.
8. Describe possible explanations of dreaming.
 Outline
1. Rhythms of Waking and Sleep
2. Stages of Sleep
3. Functions of Sleep REM
4. Brain Mechanisms of Sleep
5. Sleep Disorders
 Rhythms of Waking and Sleep
• Early psychologists (particularly the behaviorists)
believed that cycles of wakefulness and sleep were
dependent upon external stimuli
– Example: changes in light or temperature
• Curt Richter in 1922 proposed that the body generates
its own cycles of activity and inactivity
– The idea of self-generated rhythms was a major
step toward viewing animals as active producers of
behaviors.
 Endogenous Circannual Rhythms
• Some animals generate endogenous circannual
rhythms, internal mechanisms that operate on an
annual or yearly cycle
– Example: birds migratory patterns; animals storing
food for the winter
– Even if the birds are kept in a cage with no clues to
the season, they become restless in the spring, and
if they are released, they fly north (Gwinner, 1986).
 Endogenous Circadian Rhythms
• All animals produce endogenous circadian rhythms,
internal mechanisms that operate on an approximately
24-hour cycle
– Sleep cycle
– Frequency of eating and drinking
– Body temperature
– Secretion of hormones
– Urination
– Metabolism
– Mood
– Sensitivity to drugs
Daily Pattern of Body Temperature in Human

Figure 8.2 Mean rectal temperatures for nine adults

Body temperature reaches its low for the day about 2 hours after sleep onset; it
reaches its peak about 6 hours before sleep onset.
Adapted from “Sleep-Onset Insomniacs Have Delayed Temperature Rhythms,” by
M. Morris, L. Lack, and D. Dawson, Sleep, 1990, 13, pp. 1–14
Daily Pattern of Positive Moods
Figure 8.3 Reported positive mood
over time
During 30 hours in an unchanging
laboratory environment, the average
young adult reported most pleasant
mood in the late afternoon or early
evening, and the least pleasant mood
around 5 a.m. to 7 a.m. The pattern
was similar for those who started the
procedure in the morning (above) or
in the evening (below).
From “Nature’s clocks and human
mood: The circadian system
modulates reward motivation,” by G.
Murray, C. L. Nicholas, J. Kleiman, R.
Dwyer, M. J. Carrington, N. B. Allen, et
al., 2009, Emotion, 9, pp. 705–716
 Setting and Resetting the Biological Clock
• The purpose of the circadian rhythm is to keep our
internal workings in phase with the outside world
• The human circadian clock generates a rhythm slightly
longer than 24 hours when it has no external cue to
set it
 Setting and Resetting the Biological Clock
• Recall your sleep pattern during your summer
vacation!
• Do you realize that you are sleeping later and later and
wake up later and later day by day?
Summer Vacation Wake up Bed Time
Day 1 7 am (7 am) After 10 pm (10 pm)
Day 2 8 am 11 pm
Day 3 9 am Mid-night
…

School day 7 am (but you can

barely wake up!)
 Setting and Resetting the Biological Clock
• The purpose of the circadian rhythm is to keep our
internal workings in phase with the outside world
• The human circadian clock generates a rhythm slightly
longer than 24 hours when it has no external cue to
set it
– On weekends, when most of us are freer to set our
own schedules, we expose ourselves to lights,
noises, and activity at night and then awaken late
the next morning. By Monday morning, when the
clock on your table indicates 7 am, your biological
clock may say 5 am
 Setting and Resetting the Biological Clock
• Resetting our circadian rhythms is sometimes
necessary or we would gradually drift away from the
correct time
• Zeitgeber: German meaning “time giver”; refers to the
stimulus that resets the circadian rhythm
– Examples: sunlight, tides, exercise, meals, arousal of
any kind, meals, temperature of environment, etc.
– Social stimuli (i.e., the effects of other people) are
ineffective as zeitgebers, unless they induce
exercise or other vigorous activity
– Depression, irritability, and impaired job
performance may result during the adjustment
 Setting and Resetting the Biological Clock
• What happen to blind people?
• Some set their circadian rhythms by noise,
temperature, meals, and activity.
• When their cycles are in phase with the clock, all is
well, but when they drift out of phase, they experience
insomnia at night and sleepiness during the day
• More than half of all blind people report frequent
sleep problems
 Jet Lag
• Refers to the disruption of the circadian rhythms due
to crossing time zones
– Stems from a mismatch of the internal circadian
clock and external time
– Sleepiness during the day, sleeplessness at night,
and impaired concentration
 Jet Lag
• Traveling west “phase-delays” our circadian rhythms
→ Most people find it easier to adjust
→ We stay awake later at night and then awaken late
the next morning,
• Traveling east “phase-advances” our circadian rhythms
→ Most people find it difficult to adjust
→ Most people find it difficult to sleep before their
body's usual time and difficult to wake up early the
next day
• Adjusting to jet lag is often stressful → Increase in
cortisol → decrease in hippocampal volume →
memory impairment
 https://images.app.goo.gl/f1HuDWfpWuCVJpvL8

Britannica, T. Editors of Encyclopaedia (2015, January

23). Jet lag. Encyclopedia Britannica.
https://www.britannica.com/science/jet-lag
 Shift Work
• Sleep duration depends on when one goes to sleep
• Working at night does not reliably change the
circadian rhythm
– Even after long periods of working at night, people
can still feel sleepy, sleep poorly during the day, and
body temperature peaks while sleeping instead of
while working
– People who have done shift work for years tend to
perform worse than average on cognitive tests, and
have more accidents than day-shift workers
• People adjust best to night work if they sleep in a very
dark room during the day and work under very bright
lights at night
 Morning People and Evening People
• Cycles can differ between people and lead to different
patterns of wakefulness and alertness
• “Morning people”: awaken early, reach their peak of
productivity early, and become less alert later in the
day.
• “Evening people”: warm up more slowly, both literally
and figuratively, reaching their peak in the late
afternoon or evening
– They tolerate staying up all night better than
morning people do
 Morning People and Evening People
• Change as a function of age:
– Young children are morning people
– Adolescents are often night people
→ Observed in other species; result from increased
levels of sex hormones
– Starting about age 20, people gradually reverses
• For adults: change as a function of genetics and
several environmental factors, including artificial light
– Example: People who live in a big city, surrounded
by bright lights, are more likely to stay up late than
are people in rural areas.
 Morning People and Evening People
Behavioral differences between morning and evening
people:
• Morning people reported being happier than evening
people, on average
– Possibly because their biological rhythms are more
in tune with their 9-to-5 work schedule
• Morning type people tend to be more moral and
honest in the morning, whereas evening type people
tend to be more moral and honest in the evening
 Mechanisms of the Biological Clock
• Mechanisms of the circadian rhythms
– The suprachiasmatic nucleus (SCN)
– Genes that produce certain proteins
– Melatonin levels
 The Suprachiasmatic Nucleus (SCN)
• The main control center of the circadian rhythms of
sleep and temperature
– Located above the optic chiasm and part of the
hypothalamus
• Damage to the SCN results
in less consistent body
rhythms that are no longer
synchronized to
environmental patterns of
light and dark

Figure 8.7 The suprachiasmatic nucleus (SCN) of humans

 The Suprachiasmatic Nucleus (SCN) and
the Circadian Rhythm
• Generates circadian rhythms in a genetically
controlled, unlearned manner
• If SCN neurons are disconnected from the rest of the
brain or removed from the body and maintained in
tissue culture, they continue to produce a circadian
rhythm of action potentials
• Various cells communicate with each other to sharpen
the circadian rhythm
 The Suprachiasmatic Nucleus (SCN) and
the Retinohypothalamic Path
• Light resets the SCN via a small branch of the optic
nerve called the retinohypothalamic path
– Travels directly from the retina to the SCN
• The retinohypothalamic path comes from a special
population of ganglion cells that have their own
photopigment called melanopsin
 The Path of Visual Input

Figure 5.16 Major connections in the (b) Axons from the retina maintain their
visual system relationship to one another—what we call their
(a) Part of the visual input goes to theretinotopic organization—throughout their
thalamus and from there to the visualjourney from the retina to the lateral geniculate
cortex. Another part goes to the and then from the lateral geniculate to the
superior colliculus. cortex. © Cengage Learning
 The Suprachiasmatic Nucleus (SCN) and
the Retinohypothalamic Path
• These special ganglion cells…
– Located mainly near the nose
– Respond directly to light and do not require any
input from the rods or cones
– Respond to the overall average amount of light
– Respond mainly to short-wavelength (blue) light

Do you know why it is more difficult to fall asleep after

watching a long period of television, video games,
computers?
 The Suprachiasmatic Nucleus (SCN) and
the Retinohypothalamic Path
• These special ganglion cells…
– Located mainly near the nose
– Respond directly to light and do not require any
input from the rods or cones
– Respond to the overall average amount of light
– Respond mainly to short-wavelength (blue) light
→Exposure to television, video games, computers,
etc., all of which emit mostly short-wavelength
light, tends to phase delay the circadian rhythm
and make it difficult to fall asleep at the usual
time
 The Biochemistry of the Circadian
Rhythm
• Two types of genes are responsible for generating the
circadian rhythm
– Period: produce proteins called PER
– Timeless: produce proteins called TIM
• PER and TIM proteins increase the activity of certain
kinds of neurons in the SCN that regulate sleep and
waking
• Mutations in the PER gene result in odd circadian
rhythms or decreased alertness if deprived of a good
night’s sleep
 Interaction of mRNA with PER and TIM
Proteins in Fruit Flies

Figure 8.9 Feedback between proteins and genes to control sleepiness

In fruit flies (Drosophila), the concentrations of the mRNA levels for PER and
TIM oscillate over a day, and so do the proteins that they produce.
© Cengage Learning
Mammals have three versions of the PER protein and several proteins closely
related to TIM and the others found in flies. So, findings from the fruit flies may
tell us something about human beings.
 Melatonin
• The SCN regulates waking and sleeping by controlling
activity levels in other areas of the brain
• The SCN regulates the pineal gland, an endocrine gland
located posterior to the thalamus
• The pineal gland secretes melatonin,
a hormone that increases sleepiness
• Melatonin is a widespread chemical,
found in nearly all animals, as well as
in plants and bacteria
• Melatonin also helps control the onset of puberty and
bodily adjustments to changes of season (e.g.,
hibernation).
 Melatonin
• In diurnal animals, melatonin secretion usually begins
two to three hours before bedtime at night
• In diurnal animals like humans, it increases sleepiness.
• In nocturnal animals, it increases wakefulness
• Melatonin feeds back to reset the biological clock
through its effects on receptors in the SCN
• Melatonin taken in the afternoon can phase-advance
the internal clock and can be used as a sleep aid
– People start to become sleepy earlier than usual
 Outline
1. Rhythms of Waking and Sleep
2. Stages of Sleep
3. Functions of Sleep REM
4. Brain Mechanisms of Sleep
5. Sleep Disorders
 Sleep
• Sleep is a state that the brain actively produces
– Characterized by a moderate decrease in brain
activity and decreased response to stimuli
• Sleep differs from the following states:
– Coma, vegetative state, minimally conscious state,
and brain death
 Other Interruptions of Consciousness
• Coma: extended period of unconsciousness
characterized by low brain activity that remains fairly
steady
– Person shows little response to stimuli
• Vegetative state: person alternates between periods of
sleep and moderate arousal but no awareness of
surrounding
– Some autonomic arousal to painful stimulus
– No purposeful activity/response to speech
 Other Interruptions of Consciousness
• Minimally conscious state: one stage higher than a
vegetative state marked by occasional brief periods of
purposeful action and limited speech comprehension
• Brain death: no sign of brain activity and no response
to any stimulus
– Physicians usually wait until someone has shown no
sign of brain activity for 24 hours before
pronouncing brain death, at which point most
people believe it is ethical to remove life support
What’s our brain doing when we are sleeping?
 Stages of Sleep—EEG
• The electroencephalograph (EEG) allowed researchers
to discover that there are various stages of sleep
– Allows researchers to compare brain activity at
different times during sleep
• A polysomnograph is a combination of EEG and eye-
movement records

Figure 8.10 Sleeping person

with electrodes in place on the
scalp for recording brain activity
Hank Morgan/Rainbow/RGB
Ventures LLC dba
SuperStock/Alamy
 Our Brain Is Active During Sleep!

FIGURE 4.14
Brain Activity During Sleep
Credit: https://people.cornellcollege.edu/dsherman/illusions/# (Drawing of retina from Hubel DH Eye, Brain, and
Vision Scientific American Library p.38)
Polysomnograph Records

Figure 8.11 Polysomnograph

records from a college
student
For each of these records, the
top line is the EEG from one
electrode on the scalp. The
middle line is a record of eye
movements. The bottom line
is a time marker, indicating 1-
second units. Note the
abundance of slow waves in
stages 3 and 4.
Source: Records provided by
T. E. LeVere
Wakefulness and Relaxation
• Beta waves are present when
one is alert and fully awake
• Alpha waves are present
when one begins a state of
relaxation
– Alpha waves also occur
when people really focus
their attention on
something or when they
close their eyes and relax
Stage 1 Sleep
• Stage 1 sleep is when sleep
has just begun
– The EEG is dominated by
irregular, jagged, and low
voltage waves
– Brain activity begins to
decline
– Theta waves
Stage 1 Sleep
– Non-REM sleep
– Can be easily aroused
from Stage 1 sleep; if
awakened, you will
probably deny that you
were sleeping
– Might see fantastical
images or geometric
shapes; might have the
sensation of falling or that
your limbs are jerking
Stage 2 Sleep
• Stage 2 sleep is characterized
by the presence of:
– Sleep spindles: 12- to 14-
Hz waves during a burst
that lasts at least half a
second
– K-complex: a sharp wave
associated with temporary
inhibition of neuronal
firing
Stage 2 Sleep - Sleep spindles
• Result from oscillating
interactions between cells in
the thalamus and the cortex.
• Sleep spindles represent
activity related to the
consolidation of memory
• The amount of spindle
activity correlates more than
0.7 with nonverbal tests of IQ
Stage 2 Sleep
• Stage 2 sleep is characterized
by the presence of:
– Non-REM sleep
– Really asleep!
– Breathing becomes more
regular
– Become less sensitive to
external stimulation
– As people age and sleep
more lightly, their EEGs
show fewer sleep spindles
Slow Wave Sleep—Stage 3 and Stage 4
• Stage 3 and stage 4 together
constitute slow wave sleep
(SWS) and is characterized
by:
– EEG recording of slow,
large amplitude wave
– Slowing of heart rate,
breathing rate, and brain
activity
– Highly synchronized
neuronal activity
– Delta waves
Slow Wave Sleep—Stage 3 and Stage 4
– Deep sleep stage
– Very hard to wake up;
often very disoriented
when they do wake up
– Can still process some
relevant information of
the environment,
especially those signals for
potential danger (e.g.,
parents wake up to
children’s cries but not to
traffic noise)
Paradoxical or REM Sleep
• Rapid eye movement sleep
(REM) describes periods
characterized by rapid eye
movements during sleep
– Also know as paradoxical
sleep: deep sleep in some
ways but light sleep in
other ways
• REM sleep for human
• Paradoxical sleep for species
that lack eye movements
 Paradoxical or REM Sleep
– EEG waves are irregular,
low-voltage, and fast
– Beta waves
– REM sleep is light, similar
to Stage 1 except for its eye
movements
– Postural muscles of the
body are more relaxed than
other stages
→ Most of the body
muscles are paralyzed
REM sleep combines aspects of deep sleep, light sleep, and
features that are difficult to classify as deep or light
Paradoxical or REM Sleep
– Occurs when the sleep cycles
reverse (after about 90
minutes of sleep and
returning to stage 1)
→ We cycle through REM five
times per night
– Most dreaming occur (80%-
90% of the time when people
are awakened during REM
sleep reported dreaming
compared to less than half of
the time during non-REM
sleep)
 NREM and REM Cycles
• Stages other than REM are referred to as non-REM
sleep (NREM)
• When people fall asleep, they progress through stages
1, 2, 3, and 4 in sequential order
– After about an hour, the person begins to cycle back
through the stages from stage 4 to stages 3 and 2
and then REM
– The sequence repeats with each cycle lasting
approximately 90 minutes
 Stages of sleep in a night

FIGURE 4.15 Stages of Sleep

During a typical night, a sleeper goes through this sequence of EEG
stages 5 times. Notice that sleep is deepest during the first part of the
night and shallower later on, when REM sleep (highlighted in green)
becomes more prominent.
 NREM and REM Cycles
• Stages 3 and 4 sleep predominate early in the night
– Length of stage decreases as the night progresses
• REM sleep is predominant later at night
– Length increases as the night progresses
• The amount of REM depends on time of day more
than how long you have been asleep
• REM is strongly associated with dreaming (80%-90%),
but people also report dreaming in other stages of
sleep
 Pattern of Sleep as a Function of Age,
Health, and Others
• Older adult has less slow-wave sleep and many more
awakenings during the night
• The results vary from one older adult to another,
depending on health and other factors
• The frequency of awakenings correlates with loss of cells
in the hypothalamus, and with a tendency toward
cognitive decline
 Amounts of REM Sleep
• Humans spend one-third of their life asleep; about
one-fifth is spent in REM
• Species vary in amount of sleep time spent in REM
– Percentage of REM sleep is positively correlated
with the total amount of sleep in most animals
– Cats spend up to 16 hours a day sleeping with lots
of that time in REM sleep
– Among humans, those who get the most sleep have
the highest percentage of REM
 The Relationship Between Age and REM
Sleep for Humans

Figure 8.20 Sleep patterns for people of various ages

REM sleep occupies about 8 hours a day in newborns but less than 2 hours in
most adults. The sleep of infants is not quite like that of adults, however, and
the criteria for identifying REM sleep are not the same.
Source: From “Ontogenetic Development of Human Sleep- Dream Cycle,” by H. P. Roffwarg, J. N. Muzio, and W. C.
Dement, 1966, Science, 152, pp. 604–609. Copyright 1966 AAAS. Reprinted by permission.
 Outline
1. Rhythms of Waking and Sleep
2. Stages of Sleep
3. Functions of Sleep and REM
4. Brain Mechanisms of Sleep
5. Sleep Disorders
 Functions of Sleep
• Some of the many functions of sleep include:
– Resting muscles
– Decreasing metabolism
– Performing cellular maintenance in neurons
– Reorganizing synapses
– Strengthening memories
 Sleep and Energy Conservation
• The original function of sleep was probably to
conserve energy
• Sleep conserves energy during the inefficient times,
when activity would be wasteful and possibly
dangerous.
• Conservation of energy is accomplished via:
– Decrease in body temperature of about 1–2 Celsius
degrees in mammals
– Decrease in muscle activity
 Sleep and Memory
• Sleep also plays an important role in enhancing
learning and strengthening memory
– Performance on a newly learned task is often better
the next day if adequate sleep is achieved during
the night
– → increase in sleep spindles
• Increased brain activity occurs in the area of the brain
activated by a newly learned task while one is asleep
 Sleep and Memory
• Patterns of activity in the hippocampus during learning
were similar to those shown during sleep
– Suggests that the brain replays its daily experiences
during sleep → forms new dendritic branches →
strengthens memory
• The brain strengthens some synapses and weakens
others during sleep
• Sleep spindles increase in number after new learning
 Functions of REM Sleep
• Research is inconclusive regarding the exact functions
of REM
• During REM:
– The brain may discard useless connections
– Learned motor skills may be consolidated
• Maurice (1998) suggests the function of REM is simply
to shake the eyeballs back and forth to provide
sufficient oxygen to the corneas