Sleep Medicine Reviews 15 (2011) 179e186

Contents lists available at ScienceDirect

Sleep Medicine Reviews

journal homepage: www.elsevier.com/locate/smrv

CLINICAL REVIEW

Sleep, dreaming, and mental health: A review of historical and neurobiological

perspectives
Laura Palagini a, *, Nicholas Rosenlicht b
a
b

Department of Psychiatry, University of Pisa, Pisa, Italy

Department of Psychiatry, School of Medicine, University of California, San Francisco, CA, USA

a r t i c l e i n f o

s u m m a r y

Article history:
Received 26 February 2010
Received in revised form
14 July 2010
Accepted 14 July 2010
Available online 19 September 2010

Theories as to the function of sleep and dreaming and their relationship to emotions have been studied
since the beginning of recorded history. Earliest historical records show the predominant view to be that
dreams were considered divine in origin and only later did dream theory become linked with the
functioning of the brain, perhaps most famously in psychoanalytic theory. The development of sleep
laboratory techniques ushered in a new era of the dream study and their relationship to mental health. In
this review we outline the history of theories about the genesis and function of dreams and sleep and
their relationship to mental illness from ancient mythic and religious views to the rst tentative scientic
approaches to the ascendency of psychoanalysis and ultimately to the modern era of neuroscience.
! 2010 Elsevier Ltd. All rights reserved.

Keywords:
Sleep disorders
Dreams
Depression
Psychiatric disorders

Introduction
Sleep and dreaming have always aroused our curiosity and
theories as to their cause and function have been described since the
beginning of recorded history.1e4 This fascination is widely evidenced in the arts and literature.5,6 In earliest times artists, philosophers, poets, as well as clerics, considered dreams to be divine
manifestations.1,4,7e9 They have also been an important topic of
inquiry throughout the history of philosophy.10 At the end of 19th
century the interpretation of dream content and its relationship
with human emotions became the focus of psychoanalytic
theory.11,12 In the early decades of this psychoanalytic era dreaming
was regarded as the meaningful reection of unconscious mental
functioning. In the 1950s dream research had a central role in the
elds of psychiatry and psychology.13 The development of sleep
laboratory techniques ushered in a new era of sleep/dream study
and their relation to mental health. The discovery of REM (rapid eye
movement) sleep and other advances in sleep research caused a shift
in focus. After a half century as the dominant paradigm of dream
analysis, psychology was largely eclipsed by neurobiology as
dreaming became closely identied with the physiological events of
REM sleep14 as well as NREM (non-rapid eye movement) sleep.15,16
Abbreviations: CDFF, corollary discharge or feed-forward; EEG, electroencephalography; NREM, non-rapid eye movement; rCBF, regional cerebral blood ow;
REM, rapid eye movement.
* Corresponding author. Tel.: 39 050 992 658; fax: 39 050 992 965
E-mail address: [email protected] (L. Palagini).
1087-0792/$ e see front matter ! 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.smrv.2010.07.003

Studies of dreaming in psychiatric disorders were undertaken

from a more neurobiological perspective.17e22 Beginning in the
1970s, sleep research in mental health shifted its focus towards
studying sleep disorders and their relationship to psychiatric disorders.23e25 Only after 1990 did dream studies in psychiatric disorders experience a resurgence through advances in neuroscience and
the development of neuroimaging techniques that allow us to
characterize cerebral function throughout the sleepewake
cycle.26,27 But to date, the denitive functions of dreaming, REM
sleep, and even sleep itself and their relation to human emotions
continue to elude us and remain the subjects of considerable debate.
In this paper we review the history of the relation between
sleep, dreaming, and mental health from the ancient mythic and
religious views as they evolved to the rst tentative scientic
approaches, to the ascendency of psychoanalysis, and ultimately to
the modern era of neuroscientic based research and theory. We
conclude with considerations of future approaches to dream, sleep
and mental health.
Views of sleep and dreaming from ancient through medieval times
The current assumption that dreaming is a product of the mind
was not one generally shared by our ancient forebears. Authors from
the earliest recorded civilizations, including the Mesopotamians7,28
and the Egyptians,8 rmly held to the belief that dreams were divine
in origin, i.e., the product of the gods, and were the means by which
they communicated their wishes to mortal men. Dreaming,

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L. Palagini, N. Rosenlicht / Sleep Medicine Reviews 15 (2011) 179e186

especially in religious contexts, was thought to be a supernatural

manifestation, and considered premonitory or prophetic.1e4,6
The signicance given to oneiric activity has been documented
since 4000 B.C., and the interpretation of dream content dates back
to at least the fourth century B.C. as described on clay tablets from
that period. The rst written record of dream interpretation came
from the Egyptians around 1275 B.C. A hieratic papyrus, called the
Dream Book,29 includes a list of dreams and symbols for interpretation. Considered to be messages from the gods, they were the
means by which deities communicated their wishes to mortals who
in turn consulted with special temple priests for help in their
interpretation. Dream oracles were common; sleep was induced on
supplicants in the hopes of producing prophetic dreams that could
then be interpreted by a professional diviner or priest. The belief in
a divine origin of dreaming was inherited by the ancient Greeks and
continued into Roman times.3,4,9
The Greeks built temples to Aesculapius, the god of medicine,
and utilized dream interpretation as a medical treatment.9,30 Their
belief in the prophetic power of dreams led to the publication of
dream interpretation manuals, culminating in the Oneirocritica.31
This ve-volume text on oneiromancy, which still survives, was
written by Artemidorus Daldianus, a professional diviner in the
mid-to-late second century A.D., and describes more than 30,000
different types of dreams. For Artemidorus and his contemporaries,
dreams were thought to be a pathway to the future and he was
committed to producing reliable principles for understanding
allegorical dreams. This sentiment is also evident in the epic literature, such as that of Homer and Virgil, in which dreams held
prophetic messages.3,32
The close relationship between sleep, dreaming and emotional
experience is seen widely in Greek mythology. According to their
theogony, Oneiros, the god of dreams, helped Hypnos, the god of
sleep, to reduce human suffering. Hypnos, the younger twin of
Thanatos (Death), was the fruit of the union between Nyx (Night)
and Erebus (Darkness) and used his two sons, known together as
the Oneiroi, to send sleep and dreams to mortals.33 These gods of
sleep and dreaming, who also included Morpheus (the dream
shaper), Ikelos (the god of realistic dreams), Phantasos (the god of
strange dreams), and Phobetor (the god of frightening dreams),
were the divine system that explained sleep and dreaming to the
ancient Greeks.33
Greek philosophers debated the meaning of sleep and
dreaming, and with advances in philosophy and natural science, we
begin to see rationalistic accounts interspersed with divine or
supernatural explanations of dreaming. Plato endorsed the possibility of divination through dreams,34 ascribing the site of dream
prophecy to the liver. He also supported the concept of a psychological component to dreams,35 and that in dream states one
expresses the bestial desires that are normally repressed during
wakefulness, presaging modern psychological theory.
The rst steps towards modern dream interpretation and their
relationship to emotions were taken in the 5th century B.C. when
the Greek philosopher Heraclitus suggested that a persons dream
world was created within his own mind.36 Aristotle provides us
with the most systematic study of sleep and dreaming in the
ancient world, elaborated in three full essays on the subject; On
Sleep and Waking (De Somno et Vigilia)37 On Dreams (De Insomniis)38
and On Divination Through Sleep (De Divination per Somnum).38 In
the latter studies Aristotle rmly rejects the traditional view of the
prophetic power of dreams, and explains away as mere coincidence
that dreams could predict future events. In On Dreams38 he posits
that dreams are the result of impressions traveling through the
bloodstream and thereby activating sensations in the heart. This
activity also occurs during wakefulness, but is more apparent in
sleep due to the suspension of normal perception and judgment.

The variable turbulence of blood in sleep also accounts for why

dream appearances may at times be coherent and akin to reality,
while at other times are incomplete and distorted. He further
postulated, in On Sleep and Waking37 that sleep and waking are
diametrically opposed phenomena characterized, respectively, by
the absence or presence of perception. With Aristotle dreams
moved from being viewed as externally driven phenomenon to
being the result of residual perceptions.39
With the advent of Christianity came a return of the idea that
dreams could have prophetic properties. The Old Testament has an
abundance of dream stories, and a more modern concept of dream
interpretation.4,40 The most famous interpreter of dreams in the
Bible was Joseph, and probably the most famous dream interpreted
was that of the king of Egypt in the Book of Genesis.40
From medieval times through the 19th century
During medieval times theologians practiced a more careful,
and to some extent more scientic, study of sleep and dream
phenomena. However, their interpretations were still often constrained by superstition and witchcraft.2,4 During the Renaissance,
dream interpretation went through a different process. Girolamo
Cardano (1501e76),41 an Italian mathematician and physician
wrote Synesiorum somniorum omnis generis insomnia explicantes
libri quatuor (1562) resurrecting interest in the analysis of dreams,
which were seen as complex phenomena strictly related to sleep
and emotions. Cardano produced four books about sleep disorders
and the relationship between sleep, dreams, and health, with the
rst 15 chapters describing all possible dreams. He identied four
major types of dreams based on causes. The causes he identied,
which were already present in the dreamer, depended on the
persons psychic condition. Based on the Hippocratic Theory of
Humors in which bodily uids, called humors, were responsible
for the health and mental stability of individuals. These humors
were of four types; blood, black bile, yellow bile, and
phlegm. Cardano believed that each of the humors would
correspond to a different dream through vapours, which then
would create the dream. For example, people with black bile
usually dreamt of .. darkness, earthquakes, lightning and
thunder, jails, mourning. He thus made the rst known correlation between pathological emotions, sleep and dream content.
The ancients viewed dreams as passive processes caused by
external stimulation such as vapours. But in many cases dreams
were seen as the manifestation of a divinity and therefore were
interpreted within a prophetic framework.9,10,42 Beginning with
Leibniz and continuing through modern philosophy dreaming was
seen as a product of the mind.10,42 Leibniz hypothesized that the
brain, like other internal organs, was capable of autonomous action,
and ideas could be the product of both thought and physiological
activity.43 He permitted his successors to explain dreaming as an
endogenous product of the brain in both philosophy44 and in the
rst physiologic studies of sleep.45
Towards the end of the 1800s dream interpretation centered on
the new psychological approach of psychoanalysis in which the
content of a dream was analyzed to reveal its underlying or latent
meaning about the dreamers psyche. One of the seminal works on
the subject is The Interpretation of Dreams by Sigmund Freud,11
probably the best known of relatively modern dream investigators. The rst edition was rst published in German in November
1899 as Die Traumdeutung and describes Freuds process for dream
analysis. According to Freuds point of view, dream activity reects
mans emotive side. Dreams were forms of wish-fulllment e
attempts by the unconscious to resolve a conict of some sort,
whether recent or from the recesses of the past. With Freud, the
study of dream phenomena revived the ancient practice of dream

L. Palagini, N. Rosenlicht / Sleep Medicine Reviews 15 (2011) 179e186

interpretation and established its place in the forefront of human

psychological study. Freud also proposed a relationship between
psychic functioning and the neurobiology of the brain. In his
Project for a Scientic Psychology,12 he hypothesized a model of the
mind/brain that included consideration of the neurobiological
aspects of psychological functions. Meanwhile a group of eager
European scientists were systematically studying the phenomenon
of dreaming, which became the topic of discussion at scientic
meetings. In 1861 Alfred Maury, a French physician, studied over
3000 different dreams.46 He felt external stimuli were the catalyst
of all dreams, and he questioned whether dreams were truly
recollections of mental processes occurring during sleep, or were
manufactured during the awakening process. In 1886 Goblot47 also
proposed we dream while we are awakening, and that the dreams
we report on awakening may be developed during the waking
process. Mary Calkins (1893)48 was another who undertook
a statistical analysis of dream reports.
In the early part of the 20th century, with the advent of electroencephalography, incidental observations were made of the
occurrence of dream reports by investigators studying brain waves
during sleep. Natural observations on the state of the brain during
sleep were the seeds of an experimental approach to dreaming
which had to wait for more than a century to develop. These
researchers reported that dreams may occur in association with
any type of sleep potential pattern, and no specic brain wave
activity seemed to be related to dreaming.
The discovery of REM sleep and the era of physiologic-based studies
of sleep and dreaming and their relation to mental illness
In 1953 Aserinsky and Kleitmans discovery of REM sleep14
ushered in an exciting and important epoch of research on this
newly identied state of the brain. The resulting explosion in sleep
experimentation generated an extensive literature on the nature of
REM sleep in humans and much information about the characteristics and content of REM dreams. Many of these studies originally
sought to demonstrate a link between REM sleep physiology and
specic dream content. After half a century of studying the
psychology of dreams based primarily on Freuds theories they had,
it seemed, nally found in REM sleep a unique physiologic correlate
of dreaming. In 1957 Dement and Kleitman49 published a report
which supported Aserinsky and Kleitmans previous assumptions.
In this study the authors reported dream recall rates of about 80%
after awakenings from REM sleep and only 7% from awakenings
from NREM sleep. They also observed that patterns of eye movements seemed to relate to the dream content. This nding became
known as the scanning hypothesis. A third major observation was
that the subjective dream duration seemed consistent with the
duration of the REM period preceding the awakening.50,51
The discovery of REM sleep also gave impetus to the study of
dreaming in psychiatric disorders from a more neurobiological
prospective. As a result of the belief in the inseparability of REM
sleep and dreaming, studies were pursued looking at dreaming in
mental disorders such as schizophrenia,17 manic episodes18,21
depression.19e21 Beginning in the 1970s a number of reports
described alterations of REM and NREM sleep correlating with various
psychiatric disorders.23e25 Specic REM sleep alterations were
considered for years to be biological markers for depressive disease.24,25,52e56 Studies in the literature suggest that altered sleep
patterns may precede the expression of mood disorders57,58 and vice
versa,59 thus suggesting a very close relationship with these psychiatric disorders. Renements in dream research, often based on the
assumption of a REM dreaming isomorphism, and better characterization of dream length, content and recall were employed, and
seem to indicate a relationship between dreaming and psychiatric

181

illness. This ushered in a new era of dream studies in mental disorders

and suggested that they had diagnostic as well prognostic importance
for therapeutic assessment, especially in the mood disorders.
Several studies found that depressed individuals recall fewer or
less detailed dreams than healthy controls,60,61 and in bipolar
disorder a shift to depression is associated with a decrease in
overall number of dreams reported,62 and perhaps give an indication on the duration of illness.61 Overall dream length was also
found to be shorter in depressed individuals than healthy
controls.60 Dream content seemed to reect the prevalent mood
state. Barrett et al.,60 for example, found that depressed mood in the
morning may be related to negative dream content. In mania
dreams tend to feature bizarre and improbable themes as
compared to the negative content of depressed patients, and those
with a neutral mood.62 A change in dream content was found to
closely precede an upward shift in mood, especially dreams of
death or body injury, which were more frequently reported in
developing mania.62,63 Reductions in the dreamlike quality of REM
dreams,64 or the presence of nightmares and frightening dreams
may affect and predict suicidal tendency64e66 especially if they are
repetitive65 or persistent.67 Dream characteristics may also predict
remission from depression,68 or reect response to antidepressant
treatment.69 Dreaming has been further hypothesized to function
in mood regulation,68,70 prevalent mood state and as an indicator of
suicidality,64e67, 71 resistance to treatment,68 as well as effectiveness of treatment.69 Naturally, dreaming as a model of psychosis
has been proposed and studied.72,73
Dream activity in anxiety disorders has also studied, especially in
post-traumatic stress disorder where dreaming has both diagnostic
and prognostic implications; the occurrence and content of posttraumatic dreams constitute part of the phenomenology of the
disorder and may relate to its severity.74 Recall has been found to be
reduced in these patients and may represent a coping mechanism.74
Specic dream content studied indicates the centrality of dream
imagery intensity to the underlying emotional state,75,76 and its
perturbation may serve as a potential guide in clinical care in PTSD76
as well as a better understanding of the function of dreams.74
Diagnostic and clinical implications have also been suggested in
studies of dreaming in other psychiatric disorders. Anorexic
patients have been found to have less frequent dream recall, fewer
pleasurable themes than normal controls and generally saw
themselves in their dreams as having a distorted body and experience of food.77 REM dreams of schizophrenics have also been
found to be shorter than those of controls, and dream narratives
less bizarre, suggesting these characteristics may reect an
impairment in neurocognitive processes, including emotional
processing.78 As such, data from dream studies in psychiatric
disorders suggest important clinical and prognostic implications as
dream recall, length and dream content may reect the psychopathologic evolution of these diseases.
In 1977, Hobson and McCarley79 suggested that dreaming is
fundamentally physiological and not psychologically driven. Their
activation-synthesis hypothesis proposed that the brain is periodically activated, while sensory input and motor output are
blocked e except for the oculomotor pathway. After being activated by random stimulation from pontine brain stem structures,
the brain compares this input with stored sensorimotor data and
synthesizes dream content. The authors further postulated that
REM sleep is a unique physiological state and that it alone, by
providing the requisite conditions to support dreaming, produces
bizarre and hallucinoid dreams. According to this theory the relationship between dreams and emotions is not necessarily important to ones psychology, thus ushering in a viewpoint in which
dreaming was thought to be more neurophysiological than
psychological. This extremely inuential hypothesis helped push

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L. Palagini, N. Rosenlicht / Sleep Medicine Reviews 15 (2011) 179e186

sleep and dream research away from investigating psychological

factors and towards examining physiological and cognitive factors.
Hobson and colleagues later proposed an AIM model for
dreaming and hallucinations. In this model Activation (information
processing capacity, i.e., cerebral activation or electroencephalography (EEG) amplitude), Input (gating function, the capacity to
exchange information with outside world), and Modulation (aminergic vs. cholinergic balance) interact to produce different types of
mental experience. When aminergic modulation is low (more
cholinergic), hallucinations and dreaming can occur. As sensory
input and motor output is reduced, as in REM sleep, internal
sources of stimulation predominate, resulting in more vivid
dreaming.80
A major complicating factor in much dream research since the
discovery of REM, particularly in older studies, was the widespread
assumption that REM sleep was equivalent to dream sleep. More
recently, evidence has accumulated that shows the REM dream
isomorphism assumption to be false. Over time, a large body of
evidence demonstrated that mental activity persists throughout all
stages of sleep15,16 and one by one the claims defending the
integrity of the REM dreaming equation began to fall. Still, some
argued that REM dreams were qualitatively different from NREM
reports. It was argued that reports from NREM tended to have less
organismic involvement, less elaboration, and tended to show
a closer relation to waking life e which may explain why they were
less likely to be labeled as dreams, while reports elicited from REM
sleep were more vivid and bizarre, containing more visual imagery,
participation, and affect.
Later ndings also disproved the relationship between eye
movements and the visual imagery in dreaming.49,81 By using
mechanical rather than electrical measures, the existence of rapid
eye movements in congenitally blind subjects was demonstrated82
and visual imagery was described in the absence of eye movements
in both REM83 and NREM sleep.15 Also, we were able to show using
experimental awakenings, that if one controlled for time of night
REM sleep duration did not correlate with length of dream report,
as formerly thought.84 A number of investigators including Antrobus,85 Foulkes and Schmidt,16 Fein,86 Cavallero and Foulkes87 have
also shown that when length of dream report is controlled, there is
little or no difference between REM and NREM reports in terms of
content or affect. Koulak and Goodenough88 suggested an arousalretrieval model, which hypothesized that mental activity persists
throughout all stages of sleep, but that the recall and description of
this mentation depends on arousal level, which increases as sleep
progresses.84,89 As such, stage differences in dream recall appear
more closely related to level of mnemonic activation and to access
to memory traces than to any special dream production mechanism
unique to one stage of sleep.90,91 Dream sources, as well as content,
appear similar in REM and NREM mentation.87 In general, these
results suggest that the same cognitive systems produce mental
activity irrespective of EEG sleep stage, as Foulkes and Schmidt
proposed in 1983.16 Moreover when Cicogna et al.90 compared
memory traces from daydreaming and sleep onset dreaming, they
found a similarity, suggesting that cognitive processes involved in
the creation of original narrative sequences may be similar in sleep
and waking. Ultimately, since stage REM is neither necessary nor
sufcient for dreaming, one must exercise caution in interpreting
studies where the two are assumed to be synonymous.
Studies examining sleep architecture and psychiatric illness
Sleep architecture and its relationship to mental illness has also
been extensively studied. Beginning around 1970, polysomnographic
studies suggested that, in addition to disturbances of sleep continuity,
sleep in depression is characterized by a reduction of slow wave sleep,

and a disinhibition of REM sleep e with a shortening of REM latency,

a prolongation of the rst REM period, and an increase in REM
density.24,25 Shortened REM latency was, in fact, considered by many
to be a biological marker for depressive disease.52e55,92 More recently
specic components of sleep, total delta activity, delta ratio, and REM
activity, have been considered episode-related biological features
while other components such as slowest delta activity, have been
hypothesized to represent vulnerability factors for recurrence.56
Polysomnographic studies found not only a 49% incidence psychiatric illness in rst-degree relatives of early-onset depressed patients
with reduced REM latency, but also a 70% concordance rate for
shortened REM latency, regardless of psychiatric history.93 The relative risk for unipolar depression among relatives with reduced REM
latency was almost 2 times93 and in other studies94 3 times greater
than for relatives without reduced REM latency. Moreover 1/5 of
healthy subjects with a high genetic load for psychiatric disorders
showed a conspicuous depressed-like sleep pattern.95 Short REM
latency has been associated with increased risk of major depression
beyond the familial risk associated with a depressed proband indicating that polysomnographic abnormalities may precede the clinical
expression of depression and may be useful in identifying those at
highest risk for the illness.93 In addition, alterations of REM and NREM
sleep have also been associated with schizophrenia,23 psychosis and
schizoaffective disorders.96 Recent literature suggests that the clinical
expression of a sleep disorder may precede the expression of a mood
disorder episode and mostly supports the hypothesis that sleep
disorders, especially insomnia, may represent a risk factor for the
future development of psychiatric illness, especially depression and
anxiety disorders.97e99 Further, sleep disturbance prior to trauma has
been found to predict the development of subsequent psychiatric
disorders.100
Several theoretical models originating from basic research have
tried to explain sleep abnormalities in depression. Manipulation of
the sleepewake cycle, using methods such as sleep deprivation
have been studied, showing that sleep deprivation appears to have
transient antidepressant properties.101 Thus, the relationship
between sleep, dreaming and psychiatric disorders appears to be
very complicated and far from fully understood. Further, the relationship appears two-way: psychiatric illness may lead to sleep
disturbance, and vice versa.102
Recent advances in neurobiology and investigations into the
function of sleep
The neurobiology of sleep is undoubtedly important to the
development and maintenance of healthy brain function, but its
exact role remains elusive. One proposed function of sleep is the
maintenance of synaptic homeostasis. According to this model,
several cortical circuits undergo a synaptic potentiation during
wakefulness as a product of waking cognitive experience. Sleep,
particularly NREM delta sleep, may be important in synaptic
remodeling and downscaling, which, in turn, is tied to the benecial
effects of sleep on performance.103,104 Sleep deprivation or disturbed
sleep may compromise synaptic changes and brain plasticity.105,106
Research over the last few decades has rmly established that new
neurons are generated in selected areas of the adult mammalian
brain, particularly of the hippocampal formation.105,107 The function
of adult-born neurons is still a matter of debate.108 In the case of the
hippocampus, integration of new cells into existing neuronal
circuitry may be involved in memory processes and the regulation of
emotionality.106,109 In recent years, various studies have examined
how the production of new cells and their development into neurons
is affected by sleep and sleep loss.105,106,110e115 While disruption of
sleep for a period shorter than one day appears to have little effect on
the basal rate of cell proliferation,105,106 restriction or disruption of

L. Palagini, N. Rosenlicht / Sleep Medicine Reviews 15 (2011) 179e186

sleep for 24 h,110 48 h,111 or 72e96 h,112 reduces cell proliferation in

the dentate gyrus and may have cumulative effects leading to
a major decrease in hippocampal cell proliferation, cell survival and
neurogenesis.105,106 Importantly, while short sleep deprivation may
not affect the basal rate of cell proliferation, one study in rats shows
that even mild sleep restriction may interfere with the increase in
neurogenesis113 that normally occurs with hippocampus-dependent learning.113 Since sleep deprivation also disturbs memory
formation,116,117 these data suggest that promoting survival, maturation and integration of new cells may be an unexplored mechanism by which sleep supports learning and memory processes.
The structural integrity and function of the hippocampus appears
to depend in part on sleep. Imaging studies in humans have found
the importance of sleep to hippocampal function. The formation of
memory118,119 and cognitive performance is disrupted in patients
suffering from chronic insomnia,120 in whom in vivo imaging
revealed a signicant reduction in hippocampal volume.121 The
functional consequences and cognitive disturbances associated with
insomnia, independent of the reduction in volume, may be related to
reductions in neurogenesis. These effects of sleep loss may endanger
hippocampal integrity, thereby leading to cognitive dysfunction and
contributing to the development of mood disorders.105,106,114,115,122
These ndings open new frontiers in the search for sleeps relationship to psychiatric illness, particularly mood disorders.
Advancements in neuroimaging and investigations in sleep and
dreaming
Advancements in neuroimaging techniques have made it
possible to investigate cerebral function throughout the
sleepewake cycle in normal human subjects. This approach has
allowed sleep physiological theories developed from animal data to
be conrmed, but has also introduced original concepts about the
neurobiological mechanisms of sleep, dreams and memory in
humans.27 NREM sleep, when compared to wakefulness or REM
sleep, is characterized by a global decrease in cerebral blood ow,
and a decrease in regional cerebral blood ow (rCBF) in the dorsal
pons, mesencephalon, thalamus, basal ganglia, basal forebrain and
anterior hypothalamus, prefrontal cortex, anterior cingulate cortex
and precuneus.12, 123e125 In contrast to NREM sleep, REM sleep is
characterized by high cerebral blood ow126 and regional activations during REM sleep were found in the pontine tegmentum,
thalamus, amygdala, hippocampus, anterior cingulate cortex, temporo-occipital areas, basal forebrain, cerebellum and caudate
nucleus; conversely, regional deactivations were located in the
dorso-lateral prefrontal cortex, posterior cingulate gyrus, the precuneus and inferior parietal cortex.127
Although these changes are usually interpreted in relation to
physiological and cellular mechanisms, the specic regional distribution of brain activity during REM sleep has been suggested to
relate to specic dream features.27 For example, the perceptual
aspects of dreams could be related to the activation of posterior
cortices. Accordingly, patients with occipito-temporal lesions may
report a cessation of visual dream imagery.26 Secondly, emotional
features in dreams could similarly be expected to be related to
activation of amygdalar complexes, the orbito-frontal cortex and the
anterior cingulate cortex.26 Thirdly, the relative hypoactivation of the
prefrontal cortex could explain the alteration in logical reasoning,
working memory, episodic memory and executive functioning that
appear in dream reports from REM sleep awakenings.26,27,127 These
assumptions are of great theoretical interest to the understanding of
dream physiology but remain highly speculative because combined
dreaming and functional imaging data are still very sparse and little
is known about the physiology of NREM sleep dreaming. We must be
cautious not to repeat the earlier mistake of equating REM sleep with

183

dreaming, and examine why many REM and NREM dreams are
indistinguishable.15,16,85e91 As yet, relatively few neuroimaging
studies have been dedicated to human sleep disorders, dreaming
and their relationship to mental illness. Future studies could attempt
to link dreaming experiences with patterns of regional cerebral
activity in psychiatric disorders. Thus far, precise neural correlates of
dreaming remain elusive and the denitive functions of dreaming
and of sleep itself e as well as their relationship to normal and
pathological emotions in humans e continue to elude us and remain
the subject of considerable debate.
One of the most intriguing similarities of dreams and psychosis
is that, in both states, neural activity in an individuals brain is
experienced as produced by the external environment. It has been
hypothesized that such experiences can be produced when the
corollary discharge or feed-forward (CDFF) mechanisms of the
mind are disabled. Such mechanisms were rst identied in
sensorimotor systems where they operate to distinguish sensory
experiences produced by self-initiated action (neural commands)
from sensory experiences produced by the environment. Feinberg128 hypothesized that similar systems exist for mental activity,
which Hughlings Jackson129 proposed is simply the highest level of
motor activity. In psychoses such as schizophrenia, neural activity
produced in the patients brain is experienced as produced from
activity coming from outside, often in the form of voices. It is true,
of course, that in the dreams of normal individuals, the hallucinatory experiences are more often visual than auditory. Nevertheless,
in both brain states e dreaming and psychosis e this disorder of
agency can be understood as a failure of CDFF systems, as Feinberg and Guazzelli130 pointed out. This eld is now ripe for
sophisticated experimentation with new imaging techniques.
Conclusion
In this review we have traced the evolution of our knowledge
and beliefs about sleep and dreaming, their function, and their
connection to mental illness. The various theories attempting to
explain these phenomena have shifted dramatically through
history; for centuries dreams were a means to manage worries
about the future, rst as messengers from gods, later as
a messenger from the unconscious. With new and improved
techniques for assessing brain function in vivo, attention has shifted away from studying the subjective characteristics of dreaming
to examining specic neurophysiology. Still, the integration of sleep
and dream study methodology with recent developments in neuroimaging and functional brain exploration provides an exciting
and promising area of inquiry that may one day help answer the
many remaining questions about the nature and function of sleep
and dreams, and their relationship to psychiatric illness.
Research agenda
1. The integration of sleep and dream study methodology
with the study of functional brain exploration in vivo in
psychiatric illness is an exciting and promising area of
inquiry that may one day help answer the many
remaining questions about the nature and function of
sleep and dreams in humans, and the pathophysiology
of the major psychiatric disorders.
2. Some potentially useful findings from earlier studies on
dreaming in patients with mood or anxiety disorders
may guide clinical care, as well as help guide future
research. Studies on dream content and recall in larger
samples of specific psychiatric disorders in future
research can increase our knowledge in this field.

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Practice points
1. Since earliest times dream content has been studied in
relationship with human health and function.
2. The development of sleep laboratory techniques
ushered in a new era of the study of dreams and mental
health. Over time, this focus has become more neurobiological. Studies on dreaming have focused especially on dreaming mechanisms, functions and
relationship with cognitive human functions and sleep
research within a more neuroscientific perspective, and
shifted from dreaming to the neurobiological activity of
sleep and its relationship with mental health.
3. Perturbations of sleep have then been studied as
related to pathological emotional states: sleep disorders are thus commonly considered risk factors in the
development of psychiatric disorders, particularly
mood disorders. Pathological emotional states may
lead to sleep disturbance, and vice versa. The neurobiology of sleep may be moreover important in the
development and maintenance of brain function and
may have a possible role in the maintenance of
synaptic homeostasis. Prolonged restriction or
disruption of sleep may have cumulative effects leading
to a major decrease in hippocampal cell proliferation,
cell survival and neurogenesis.
4. Neuroscience and the development of neuroimaging
techniques have made possible the characterization
of many aspects of cerebral function throughout the
sleepewake cycle, but to date we do not yet fully
understand the significance of sleep and dreaming. The
various theories attempting to explain these
phenomena have shifted dramatically through history.

Acknowledgment
Thanks to Aldo Palagini for his important support.
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