Chapter 9

Chapter 9
Wakefulness and Sleep
Rhythms of Waking and Sleep
• Animals generate endogenous 24 hour cycles
of wakefulness and sleep.
• 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.
Rhythms of Waking and Sleep
• All animals produce endogenous circadian
rhythms, internal mechanisms that operate on
an approximately 24 hour cycle.
– Regulates the sleep/ wake cycle.
– Also regulates the frequency of eating and
drinking, body temperature, secretion of
hormones, volume of urination, and
sensitivity to drugs.
Fig. 9-2, p. 267
Rhythms of Waking and Sleep
Circadian rhythms:
• Remains consistent despite lack of
environmental cues indicating the time of day
• Can differ between people and lead to
different patterns of wakefulness and
• Change as a function of age.
– Example: sleep patterns from childhood to
late adulthood.
Rhythms of Waking and Sleep
• Experiments designed to determine the
length of the circadian rhythm place subjects
in environments with no cues to time of day.
• Results depend upon the amount of light to
which subjects are artificially exposed.
– Rhythms run faster in bright light conditions
and subjects have trouble sleeping.
– In constant darkness, people have difficulty
Rhythms of Waking and Sleep
• Human circadian clock generates a rhythm
slightly longer than 24 hours when it has no
external cue to set it.
• Most people can adjust to 23- or 25- hour day
but not to a 22- or 28- hour day.
• Bright light late in the day can lengthen the
circadian rhythm.
Rhythms of Waking and Sleep
• Mechanisms of the circadian rhythms include
the following:
– The Suprachiasmatic nucleus.
– Genes that produce certain proteins.
– Melatonin levels.
Rhythms of Waking and Sleep
• The suprachiasmatic nucleus (SCN) is part of
the hypothalamus and the main control center
of the circadian rhythms of sleep and
– Located above the optic chiasm.
– Damage to the SCN results in less
consistent body rhythms that are no longer
synchronized to environmental patterns of
light and dark.
Fig. 9-4, p. 269
Rhythms of Waking and Sleep
• The SCN is genetically controlled and
independently generates the circadian
• Single cell extracted from the SCN and raised
in tissue culture continues to produce action
potential in a rhythmic pattern.
• Various cells communicate with each other to
sharpen the circadian rhythm.
Rhythms of Waking and Sleep
Two types of genes are responsible for
generating the circadian rhythm.
1. Period - produce proteins called Per.
2. 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.
Fig. 9-5, p. 270
Rhythms of Waking and Sleep
• The SCN regulates waking and sleeping by
controlling activity levels in other areas of the
• The SCN regulates the pineal gland, an
endocrine gland located posterior to the
• The pineal gland secretes melatonin, a
hormone that increases sleepiness.
Rhythms of Waking and Sleep
• Melatonin secretion usually begins 2 to 3
hours before bedtime.
• Melatonin feeds back to reset the biological
clock through its effects on receptors in the
• Melatonin taken in the afternoon can phaseadvance the internal clock and can be used
as a sleep aid.
Rhythms of Waking and Sleep
• The purpose of the circadian rhythm is to
keep our internal workings in phase with the
outside world.
• Light is critical for periodically resetting our
circadian rhythms.
• A zeitgeber is a term used to describe any
stimulus that resets the circadian rhythms.
• Exercise, noise, meals, and temperature are
others zeitgebers.
Rhythms of Waking and Sleep
• 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.
• Characterized by sleepiness during the day,
sleeplessness at night, and impaired
• Traveling west “phase-delays” our circadian
• Traveling east “phase-advances” our
circadian rhythms.
Fig. 9-6, p. 272
Rhythms of Waking and Sleep
• Light resets the SCN via a small branch of the
optic nerve known as the retinohypothalamic
– 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 cells respond directly to light and do
not require any input from the rods or
Stages of Sleep And Brain
• Sleep is a specialized state that serves a
variety of important functions including:
– conservation of energy.
– repair and restoration.
– learning and memory consolidation.
Stages of Sleep And Brain
• The electroencephalograph (EEG) allowed
researchers to discover that there are various
stages of sleep.
• Over the course of about 90 minutes:
– a sleeper goes through sleep stages 1, 2,
3, and 4
– then returns through the stages 3 and 2 to
a stage called REM.
Stages of Sleep And Brain
• Alpha waves are present when one begins a
state of relaxation.
• Stage 1 sleep is when sleep has just begun.
– the EEG is dominated by irregular, jagged,
low voltage waves.
– brain activity begins to decline.
Stages of Sleep And Brain
• 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-complexes - a sharp high-amplitude
negative wave followed by a smaller,
slower positive wave.
Stages of Sleep And Brain
• Stage 3 and stage 4 together constitute slow
wave sleep (SWS) and is characterized by:
– EEG recording of slow, large amplitude
– Slowing of heart rate, breathing rate, and
brain activity.
– Highly synchronized neuronal activity.
Stages of Sleep And Brain
• Rapid eye movement sleep (REM) are
periods characterized by rapid eye
movements during sleep.
• Also known as “paradoxical sleep” because it
is deep sleep in some ways, but light sleep in
other ways.
• EEG waves are irregular, low-voltage and
• Postural muscles of the body are more
relaxed than other stages.
Fig. 9-9, p. 276
Stages of Sleep And Brain
• Stages other than REM are referred to as
non-REM sleep (NREM).
• When one falls 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 than REM.
• The sequence repeats with each cycle lasting
approximately 90 minutes.
Stages of Sleep And Brain
• Stage 3 and 4 sleep predominate early in the
– The length of stages 3 and 4 decrease as
the night progresses.
• REM sleep is predominant later in the night.
– Length of the REM stages increases as the
night progresses.
• REM is strongly associated with dreaming,
but people also report dreaming in other
stages of sleep.
Fig. 9-10, p. 277
Stages of Sleep And Brain
• Various brain mechanisms are associated
with wakefulness and arousal.
• The reticular formation is a part of the
midbrain that extends from the medulla to the
forebrain and is responsible for arousal.
Table 9-1, p. 280
Stages of Sleep And Brain
• The pontomesencephalon is a part of the
midbrain that contributes to cortical arousal.
– Axons extend to the thalamus and basal
forebrain which release acetylcholine and
– produce excitatory effects to widespread
areas of the cortex.
• Stimulation of the pontomesencephalon
awakens sleeping individuals and increases
alertness in those already awake.
Stages of Sleep And Brain
• The locus coeruleus is small structure in the
pons whose axons release norepinephrine to
arouse various areas of the cortex and
increase wakefulness.
– Usually dormant while asleep.
Fig. 9-11, p. 279
Stages of Sleep And Brain
• The basal forebrain is an area anterior and
dorsal to the hypothalamus containing cells
that extend throughout the thalamus and
cerebral cortex.
• Cells of the basal forebrain release the
inhibitory neurotransmitter GABA.
• Inhibition provided by GABA is essential for
• Other axons from the basal forebrain release
acetylcholine which is excitatory and
increases arousal.
Fig. 9-12, p. 280
Stages of Sleep And Brain
• The hypothalamus contains neurons that
release “histamine” to produce widespread
excitatory effects throughout the brain.
– Anti-histamines produce sleepiness.
Stages of Sleep And Brain
• Orexin is a peptide neurotransmitter released
in a pathway from the lateral nucleus of the
hypothalamus highly responsible for the
ability to stay awake.
– Stimulates acetylcholine-releasing cells in
the forebrain and brain stem to increase
wakefulness and arousal.
Stages of Sleep And Brain
Decreased arousal required for sleep is
accomplished via the following ways:
1. Decreasing the temperature of the brain
and the body.
2. Decreasing stimulation by finding a quiet
3. Accumulation of adenosine in the brain to
inhibit the basal forebrain cells
responsible for arousal.
– Caffeine blocks adenosine receptors.
Stages of Sleep And Brain
4. Accumulation of prostaglandins that
accumulate in the body throughout the day
to induce sleep.
– Prostaglandins stimulate clusters of
neurons that inhibit the hypothalamic
cells responsible for increased arousal.
Stages of Sleep And Brain
• During REM sleep:
– Activity increases in the pons (triggers the
onset of REM sleep), limbic system,
parietal cortex and temporal cortex.
– Activity decreases in the primary visual
cortex, the motor cortex, and the
dorsolateral prefrontal cortex.
Stages of Sleep And Brain
• REM sleep is also associated with a
distinctive pattern of high-amplitude electrical
potentials known as PGO waves.
• Waves of neural activity are detected first in
the pons and then in the lateral geniculate of
the hypothalamus, and then the occipital
• REM deprivation results in high density of
PGO waves when allowed to sleep normally.
Fig. 9-13, p. 281
Stages of Sleep And Brain
• Cells in the pons send messages to the
spinal cord which inhibit motor neurons that
control the body’s large muscles.
– Prevents motor movement during REM
• REM is also regulated by serotonin and
– Drugs that stimulate Ach receptors quickly
move people to REM.
– Serotonin interrupts or shortens REM.
Stages of Sleep And Brain
• Insomnia is a sleep disorder associated with
inability to fall asleep or stay asleep.
– Results in inadequate sleep.
– Caused by a number of factors including
noise, stress, pain medication.
– Can also be the result of disorders such as
epilepsy, Parkinson’s disease, depression,
anxiety or other psychiatric conditions.
– Dependence on sleeping pills and shifts in
the circadian rhythms can also result in
Fig. 9-15, p. 282
Stages of Sleep And Brain
• Sleep apnea is a sleep disorder characterized
by the inability to breathe while sleeping for a
prolonged period of time.
• Consequences include sleepiness during the
day, impaired attention, depression, and
sometimes heart problems.
• Cognitive impairment can result from loss of
neurons due to insufficient oxygen levels.
• Causes include, genetics, hormones, old age,
and deterioration of the brain mechanisms
that control breathing and obesity.
Stages of Sleep And Brain
• Narcolepsy is a sleep disorder characterized
by frequent periods of sleepiness.
• Four main symptoms include:
– Gradual or sudden attack of sleepiness.
– Occasional cataplexy - muscle weakness
triggered by strong emotions.
– Sleep paralysis- inability to move while
asleep or waking up.
– Hypnagogic hallucinations- dreamlike
experiences the person has difficulty
distinguishing from reality.
Stages of Sleep And Brain
(Insomnia cont’d)
• Seems to run in families although no gene
has been identified.
• Caused by lack of hypothalamic cells that
produce and release orexin.
• Primary treatment is with stimulant drugs
which increase wakefulness by enhancing
dopamine and norepinephrine activity.
Stages of Sleep And Brain
• Periodic limb movement disorder is the
repeated involuntary movement of the legs
and arms while sleeping.
– Legs kick once every 20 to 30 seconds for
periods of minutes to hours.
– Usually occurs during NREM sleep.
Stages of Sleep And Brain
• REM behavior disorder is associated with
vigorous movement during REM sleep.
– Usually associated with acting out dreams.
– Occurs mostly in the elderly and in older
men with brain diseases such as
– Associated with damage to the pons
(inhibit the spinal neurons that control large
muscle movements).
Stages of Sleep And Brain
• “Night terrors” are experiences of intense
anxiety from which a person awakens
screaming in terror.
– Usually occurs in NREM sleep.
• “Sleep talking” occurs during both REM and
NREM sleep.
• “Sleepwalking” runs in families, mostly occurs
in young children, and occurs mostly in stage
3 or 4 sleep.
Why Sleep? Why REM? Why Dreams?
• Functions of sleep include:
– Energy conservation.
– Restoration of the brain and body.
– Memory consolidation.
Why Sleep? Why REM? Why Dreams?
• The original function of sleep was to probably
conserve energy.
• Conservation of energy is accomplished via:
– Decrease in body temperature of about 1-2
Celsius degrees in mammals.
– Decrease in muscle activity.
Why Sleep? Why REM? Why Dreams?
• Animals also increase their sleep time during
food shortages.
– sleep is analogous to the hibernation of
• Animals sleep habits and are influenced by
particular aspects of their life including:
– how many hours they spend each day
devoted to looking for food.
– Safety from predators while they sleep
• Examples: Sleep patterns of dolphins,
migratory birds, and swifts.
Fig. 9-17, p. 287
Why Sleep? Why REM? Why Dreams?
• Sleep enables restorative processes in the
brain to occur.
– Proteins are rebuilt.
– Energy supplies are replenished.
• Moderate sleep deprivation results in
impaired concentration, irritability,
hallucinations, tremors, unpleasent mood,
and decreased responses of the immune
Why Sleep? Why REM? Why Dreams?
• People vary in their need for sleep.
– Most sleep about 8 hours.
• Prolonged sleep deprivation in laboratory
animals results in:
– Increased metabolic rate, appetite and
body temperature.
– Immune system failure and decrease in
brain activity.
Why Sleep? Why REM? Why Dreams?
• Sleep also plays an important role in
enhancing learning and strengthening
– Performance on a newly learned task is
often better the next day if adequate sleep
is achieved during the night.
• Increased brain activity occurs in the area of
the brain activated by a newly learned task
while one is asleep.
– Activity also correlates with improvement in
activity seen the following day.
Why Sleep? Why REM? Why Dreams?
• Humans spend one-third of their life asleep.
• One-fifth of sleep time is spent in REM.
• Species vary in amount of sleep time spent in
– Percentage of REM sleep is positively
correlated with the total amount of sleep in
most animals.
• Among humans, those who get the most
sleep have the highest percentage of REM.
Fig. 9-18, p. 289
Why Sleep? Why REM? Why Dreams?
• REM deprivation results in the following:
– Increased attempts of the brain/ body for
REM sleep throughout the night.
– Increased time spent in REM when no
longer REM deprived.
• Subjects deprived of REM for 4 to 7
nights increased REM by 50% when no
longer REM deprived.
Why Sleep? Why REM? Why Dreams?
• 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.
Why Sleep? Why REM? Why Dreams?
Biological research on dreaming is
complicated by the fact that subjects can not
often accurately remember what was
• Two biological theories of dreaming include:
1. The activation-synthesis hypothesis.
2. The clinico-anatomical hypothesis.
Why Sleep? Why REM? Why Dreams?
• The activation-synthesis hypothesis suggests
dreams begin with spontaneous activity in the
pons which activates many parts of the
– The cortex synthesizes a story from the
pattern of activation.
– Normal sensory information cannot
compete with the self-generated
stimulation and hallucinations result.
Why Sleep? Why REM? Why Dreams?
• Input from the pons activates the amygdala
giving the dream an emotional content.
• Because much of the prefrontal cortex is
inactive during PGO waves, memory of
dreams is weak.
– Also explains sudden scene changes that
occur in dreams.
Why Sleep? Why REM? Why Dreams?
• The clinico-anatomical hypothesis places less
emphasis on the pons, PGO waves, or even
REM sleep.
– Suggests that dreams are similar to
thinking, just under unusual circumstances.
• Similar to the activation synthesis hypothesis
in that dreams begin with arousing stimuli that
are generated within the brain.
– Stimulation is combined with recent
memories and any information the brain is
receiving from the senses.
Why Sleep? Why REM? Why Dreams?
• Since the brain is getting little information
from the sense organs, images are generated
without constraints or interference.
• Arousal can not lead to action as the primary
motor cortex and the motor neurons of the
spinal cord are suppressed.
• Activity in the prefrontal cortex is suppressed
which impairs working memory during
Why Sleep? Why REM? Why Dreams?
• Activity is high in the inferior part of the
parietal cortex, an area important for visualspatial perception.
– Patients with damage report problems with
binding body sensations with vision and
have no dreams.
– Activity is also high in areas outside of V1,
accounting for the visual imagery of
Why Sleep? Why REM? Why Dreams?
• Activity is high in the hypothalamus and
amygdala which accounts for the emotional
and motivational content of dreams.
• Either internal or external stimulation
activates parts of the parietal, occipital, and
temporal cortex.
• Lack of sensory input from V1 and no
criticism from the prefrontal cortex creates the
hallucinatory perceptions.

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