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Cataplexy is a transient episode of muscle tone loss in which humans report that awareness of external stimuli is preserved, and presumably animals are likewise aware of their environment during cataplectic attacks. Neuromodulatory activity in cataplectic dogs is largely similar to that in REM sleep except that levels of histamine are high, much like during wakefulness[ ].

It thus seems that levels of histamine are correlated with our ability to incorporate sensory stimuli into conscious experience. It would be important to establish whether histamine is indeed necessary for such incorporation, and how it may do so. For instance, could it be that in wakefulness histaminergic tone facilitates transmission of feed-forward sensory inputs in cortical layer 4, at the expense of backward signal propagation?

To put the question in a modern context, do dreams start from activity in low-level sensory areas, which is then interpreted and synthesized by higher-order areas, as is presumably the case in waking perception?

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Or do they begin as wishes, abstract thoughts, and memories deep in the brain, which are then enriched with perceptual and sensory aspects, as in imagination? Of course, it is possible that such a dichotomy is misguided, and dreams may be best conceptualized as global attractors that emerge simultaneously over many brain areas. However, as we shall see, the available data do indeed suggest that there may be a privileged direction of dream generation. In the 19th century, sensory experience was often regarded as the source of dreams, which were considered to be an attempt of the mind to interpret somatic nerve-stimuli Supplementary Fig.

A similar notion was later adopted by Henri Beaunis, and recently championed by Allan Hobson Table 1 [ 4 , 11 , 47 ]. According to his AIM model, internally generated signals originating in the brainstem during REM sleep, such as PGO waves, excite visual cortex and are later processed and synthesized by higher-order areas.

High levels of acetylcholine in the absence of aminergic neuromodulation may enhance feed-forward transmission and suppress back-propagation[ 3 , ]. By contrast, Freud and some of his followers asserted that dreams originate from psychic motives that are later instantiated as sensory percepts, much like mental imagery[ 5 ].

Deciding between these alternative views will most likely require difficult experiments in which the direction of signal flow during dreaming sleep is evaluated and compared to that during waking perception and imagery[ ] Box 4. However, various lines of evidence already suggest that dreaming may be more closely related to imagination than to perception. From lesion studies Box 3 we know that dreaming requires an intact temporo-parieto-occipital junction[ 22 , 23 ] and lesions in this region also affect mental imagery in wakefulness[ ].

Cognitive studies indicate that the skill that maximally correlates with dream recall in adults is visuo-spatial imagery[ ]. In children, dream recall develops hand in hand with visuo-spatial imagery Box 2. Other evidence comes from lucid dreamers[ 25 ] who report that it is impossible to focus on fine-grain details of visual objects, as is the case in mental imagery[ ]. Perhaps top-down connections lack the anatomical specificity to support detailed representations. The rare occurrences of smells or pain in dreams may also be related to our difficulty in imagining them vividly when awake.

However, one important difference between dreaming and mental imagery is that while imagining we are aware that the images are internally generated preserved reflective thought. During wakefulness, sensory responses precede responses in higher-order areas by more than ms[ , ]. Does neural activity during dreaming sleep show a similar feed-forward progression as in perception? Or does neural activity propagate backwards, from higher to lower areas, as it is thought to do during imagery?

This issue, which is crucial to our understanding of dream generation, could be resolved by examining unit and field potential recordings from the same neuronal populations in wake and REM or late NREM sleep in both animals and humans[ ]. One can also apply directional measures of signal propagation e.

Granger causality to hd-EEG data, and check whether the main direction of signal flow inverts between wake and sleep. Finally, one could use TMS with concurrent hd-EEG during both wake and REM sleep, and examine whether there may be a preferential direction of the brain s response to perturbations depending on behavioral state[ 10 ]. So far, most regional studies of brain activity during sleep have employed PET.

While PET allows for quantification of cerebral blood flow and comparison across vigilance states, functional MRI fMRI offers superior spatial and temporal resolutions. Event-related fMRI has been already used to map brain activity associated with phasic events such as slow waves[ ] and eye movements[ , ]. Studies of functional and effective connectivity[ ] may be especially well suited to map the functional networks underlying dreaming.

Are such connectivity patterns also a hallmark of activity in the dreaming brain? What regional brain activity underlies dreaming in NREM sleep? How do functional networks of mental imagery and dreaming compare in the same subjects? Finally, hd-EEG may be particularly suited for sleep imaging since it a allows for relatively undisturbed sleep, b upon source modeling can provide a spatial resolution roughly comparable to PET, c offers high temporal resolution suitable for evaluating signal propagation, and d can be combined with TMS during sleep.

Progress in signal decoding may ultimately enable us to investigate the neural correlates not only of dream form — what is common to all dreams — but also of dream content — what is specific to a particular dream. At least initially, it may be worthwhile to consider some coarse properties of individual dreams, such as the frequency of occurrence of faces or places in a dream report, the amount of movement, or the dominant affective valence. In principle, it should be possible to predict not only the likelihood of a report upon awakening, but also the likelihood of specific features based on preceding brain activity.

An important step in this direction would be to identify the contents of internally generated mental imagery using the same approach[ ]. Furthermore, some patients with epilepsy or post-traumatic stress disorder who experience recurring dream contents[ , ] may provide a unique opportunity to relate specific dream content to its neural basis.

If the flow of brain activity during dreaming were shown to be largely backwards, as one would expect in imagery, rather than forwards, as in perception, many of the seemingly bizarre properties of dreams, such as blended characters and scene switches, would be easier to explain, as they are standard features of our imagination. Such a top-down mode may disrupt the encoding of new memories, and thus underlie dream amnesia.

In addition, top-down mental imagery could obstruct the processing of incoming stimuli and disconnect us from the environment. If this view is correct, waking consciousness is more like watching the news in real time, while dreaming is more like watching a movie created by an imaginative director[ 81 ]. As in some B-movies, the director is not particularly choosey and any actor, dress, means of transportation, or object that is readily available will do.

In summary, dream consciousness is remarkably similar to waking consciousness, though there are several intriguing differences in volition, self-awareness and reflection, affect, and memory, and there is great variability between individual dreams. The neurophysiology of REM sleep, and in particular recent insights into its regional activity patterns, offers a useful starting point for relating dream phenomenology to underlying brain activity.

However, the initial equation of REM sleep with dreaming has been shown to be inaccurate. Thus, it is time we moved beyond sleep stages when trying to link dream consciousness to neuronal events, and focused on more subtle features of brain activity in space and time. Our profound disconnection from the external environment when dreaming poses a central unsolved paradox, the answer to which may be instrumental for understanding dreams. Viewing dreams as a powerful form of imagination can help explain many of their unique features, such as sudden transitions, uncertainty about people and places, poor subsequent recall, disconnection from the environment, and offers testable predictions for future studies.

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National Center for Biotechnology Information , U. Author manuscript; available in PMC Feb 1. Author information Copyright and License information Disclaimer. The publisher's final edited version of this article is available at Trends Cogn Sci. See other articles in PMC that cite the published article. Associated Data Supplementary Materials Abstract Dreams are a most remarkable experiment in psychology and neuroscience, conducted every night in every sleeping person. Contemporary dream research Although dreams have fascinated us since the dawn of time, their rigorous, scientific study is a recent development[ 1 — 4 ] Supplementary Fig.

BOX 1Can reports be trusted to accurately convey internal experiences in sleep? Open in a separate window. Functional neuroanatomy of human REM sleep: Phenomenology of dreams and their relation to brain activity The level and nature of our conscious experience varies dramatically in sleep. Similarities between dreaming and waking In order to gain insight into the phenomenology and neural basis of dreams, it is useful to consider both similarities and differences between waking consciousness and dreaming consciousness, and to relate these differences to changes in brain activity and organization[ 11 ].

BOX 2The development of dreams in children When do children start dreaming, and what kind of dreams do they have? BOX 3Lesion studies of dreaming The primary source on neuropsychology of dreaming is a study by Solms[ 22 ] who examined neurological patients and asked them in detail about their dreaming. Emotionality Some dreams are characterized by a high degree of emotional involvement, including joy, surprise, anger, fear, and anxiety[ 34 — 36 ].

Altered mnemonic processes Memory is drastically altered for the dream and within the dream. Table 1 Contemporary theories of dreaming.

Dreaming and the brain: from phenomenology to neurophysiology

Latent unconscious content is disguised via censorship creating the bizarre manifest dream content [ 5 ]. Our conscious state is determined by three factors: REM sleep and dreaming are characterized by high levels of activation, internal input, and cholinergic modulation [ 11 ].

Dreaming is what occurs when the mature brain is adequately activated, disconnected from external stimuli and without self-reflection. Once instigated, dreaming actively draws on memory schemas, general knowledge, and episodic information to produce simulations of the world [ 13 , 14 ].

Dream amnesia Since unconscious wishes are noxious to our consciousness, they are actively repressed via censorship processes [ 5 ]. Dream amnesia is anything but arbitrary: Dream amnesia largely stems from a state-change. Aminergic de-modulation and deactivation of dorsolateral prefrontal cortex in REM sleep create a brain state which is not favorable for subsequent memory [ 11 ].

This also explains why we forget moments of brief awakenings during sleep. Dream amnesia is primarily related to a cognitive state and lack of context. To remember, we need an external narrative to which internal events can be tied [ 14 , 21 ]. Dream amnesia cannot be explained by a state- change since dreaming can occur at any state NREM sleep and wake. PGO waves , later to be interpreted and synthesized by mnemonic and high-order modules [ 11 , 47 ].

Is REM sleep a good model for dreaming? REM sleep and dreaming can occur one without the other [ 23 , ], for example in neurological patients. Dream- like experiences are related to forebrain mechanisms rather than to REM sleep generators in brainstem [ 22 , 23 ]. Because REM sleep provides the most favorable brain conditions for dreaming, we can focus on its neurophysiology in our attempt to model the neuronal basis of dreaming [ 4 , 47 ].

Dream-like experiences can occur also in NREM sleep, sleep onset, and wakefulness [ 13 , ]. Children studies show that REM sleep may be an important condition for dreaming but not sufficient [ 13 , 21 ]. Is dreaming largely similar to waking consciousness? The apparent manifest aspect of dreams is bizarre and includes nonsensical changes in time and place, as well as incongruities of plot, character, and action [ 5 ]. This is because the true latent dream content is disguised by the censor [ 5 ].

Dreaming may be closely akin to mental illness [ 5 , ]. Dreaming is altogether comparable to delirium acute confusional state that can occur upon alcohol withdrawal [ 3 ]; REM sleep shares its physiological substrate with psycho-pathological conditions such as schizophrenia limbic hyper-activation and frontal hypo-activation [ , ]. Dreams are largely coherent and internally plausible narrative sequences rather than the stereotypical illogical sequences of bizarre images.

Content analysis indicate a strong continuity between dream content and waking life[ 13 ]. Evidence linking dreams to psychosis is limited[ ]: REM sleep deprivation does not alter schizophrenic pathology, aminergic agonists suppress REM sleep with no psychopathological effects. Primarily a cholinergic role for REM sleep and dreaming [ 4 , 11 ]. Administration of cholinergic agonists e. Dreaming is unlikely to be driven by a specific chemical or brain region. It is most likely related to a complex neurochemical mixture where serotonin, norepinephrine, and histamine are absent while both acetylcholine and dopamine are present [ 13 , ].

The function of dreaming According to Freud, dreams preserve sleep in the face of unconscious needs for excitement [ 5 ]. Dreams may serve a creative function by providing a virtual reality model protoconsciousness. The brain is preparing itself for integrative functions including learning and secondary consciousness [ 4 ]. Dreams probably have no function, but they do have coherence and meaning, which is often conflated with function[ 13 ]: This theory emphasizes dream content: This theory lacks in power with regards to explaining dreams shared by all people [ 5 ] e.

Dreaming is an attempt to best interpret activating signals in a coherent manner, and contents of individual dreams are nearly random. Nevertheless, the process of interpretation may carry some psychological meaning[ 11 ]. This theory emphasizes dream form: Are dreams directly related to previous experience? Familiar settings and people are sometimes incorporated into dreams but dreams are not a recollection of everyday life [ 14 ].

Neurophysiology of wake and sleep states A comparison of cortical activity upper panel and neuromodulator activity bottom panel in wake, early NREM when sleep pressure is high and dream reports are rare , late NREM when sleep pressure dissipates, and dream reports are more frequent , and REM sleep when dreams are most common. What determines the level of consciousness during sleep? Why is the dreamer disconnected from the environment? Are dreams more like perception or imagination? Box 4Future directions 1.

Signal propagation in dreams During wakefulness, sensory responses precede responses in higher-order areas by more than ms[ , ]. Functional networks underlying dreaming So far, most regional studies of brain activity during sleep have employed PET. Initial steps towards studying dream content Progress in signal decoding may ultimately enable us to investigate the neural correlates not only of dream form — what is common to all dreams — but also of dream content — what is specific to a particular dream. Concluding remarks In summary, dream consciousness is remarkably similar to waking consciousness, though there are several intriguing differences in volition, self-awareness and reflection, affect, and memory, and there is great variability between individual dreams.

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Acknowledgments We apologize to those whose work was not cited because of space constraints. Glossary Slow waves Oscillations of cortical origin that have frequencies below 4Hz Spindles Waxing and waning oscillations of thalamic origin that have frequencies in the sigma band 12—15Hz Ponto-Geniculo-Occipital PGO waves Phasic field potentials occurring immediately before and during REM sleep originally discovered in cats. Arkin AM, et al. The mind in sleep: Lawrence Erlbaum Associates; REM sleep and dreaming: The Interpretation of Dreams.

Foulkes D, Rechtschaffen A. Presleep Determinants of Dream Content: Effect of Two Films. Arkin A, Antrobus JS. The effects of external stimuli applied prior to and during sleep on sleep experience. Arkin A, et al. The Mind in Sleep. Rechtschaffen A, Foulkes D. Effect of Visual Stimuli on Dream Content.

The Psychophysiology of Thinking: Studies of Covert Processes. Massimini M, et al. Breakdown of cortical effective connectivity during sleep.

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Hobson JA, et al. Dreaming and the brain: Hall C, Van de Castle R. The content analysis of dreams.

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The Scientific Study of Dreams: American Psychological Association; Maquet P, et al. Functional neuroanatomy of human rapid-eye-movement sleep and dreaming. Braun AR, et al. Regional cerebral blood flow throughout the sleep-wake cycle. Nofzinger EA, et al. Forebrain activation in REM sleep: Dissociated pattern of activity in visual cortices and their projections during human rapid eye movement sleep. Experience-dependent changes in cerebral activation during human REM sleep.



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Functional neuroimaging of normal human sleep by positron emission tomography. Harvard University Press; The neuropsychology of dreaming: Dreaming and REM sleep are controlled by different brain mechanisms. Finding meaning in dreams: Behavioral and Brain Sciences. Goldberg I, et al. Desmurget M, et al. Movement intention after parietal cortex stimulation in humans.

The single-mindedness and isolation of dreams. The neuropsychology of REM sleep dreaming. Schwartz S, Maquet P.

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Sleep imaging and the neuro-psychological assessment of dreams. Goldberg II, et al. When the brain loses its self: Wolman RN, Kozmova M. Last night I had the strangest dream: Varieties of rational thought processes in dream reports. Kahn D, Hobson A. Theory of Mind in Dreaming: Awareness of Feelings and Thoughts of Others in Dreams.

Foulkes D, et al. Appropriateness of dream feelings to dreamed situations. Strauch I, et al. In search of dreams: State University of New York Press; Fosse R, et al.

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The mind in REM sleep: Nielsen TA, et al. Emotions in dream and waking event reports.

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