It has been widely demonstrated that dreaming occurs throughout human sleep. However, we once again are facing new variants of the equation “REM sleep = Dreaming.” Nielsen proposes a model that assumes covert REM processes in NREM sleep. I argue against this possibility, because dream research has shown that REM sleep is not a necessary condition for dreaming to occur. [Nielson].

We hypothesize that rapid eye movement or REM sleep evolved, in part, to mediate sexual/reproductive behaviors and strategies. Because development of sexual and mating strategies depends crucially on early attachment experiences, we further hypothesize that REM functions to mediate attachment processes early in life. Evidence for these hypotheses comes from (1) the correlation of REM variables with both attachment and sexual/reproductive variables; (2) attachment-related and sex-related hormonal release during REM; (3) selective activation during REM of brain sites implicated in (...) attachment and sexual processes; (4) effects of maternal deprivation on REM; (5) effects of REM deprivation on sexual behaviors; and (6) the REM-associated sexual excitation. To explain why we find associations among REM sleep, attachment, and adult reproductive strategies, we rely on recent extensions of parent-offspring conflict theory. Using data from recent findings on genomic imprinting, Haig (2000) and others suggest that paternally expressed genes are selected to promote growth of the developing fetus/child at the expense of the mother, while maternally expressed genes counter these effects. Because developmental REM facilitates attachment-related outcomes in the child, developmental REM may be regulated by paternally expressed genes. In that case, REM may have evolved to support the “aims” of paternal genes at the expense of maternal genes. (shrink)

There are obvious flaws in REM sleep suppression paradigms that do not allow any conclusion to be drawn either pro or contra the REM sleep-memory hypothesis. However, less intrusive investigations of REM sleep suggest that this sleep stage or its adjunct neuroendocrine characteristics exert a facilitating influence on certain aspects of ongoing memory formation during sleep. [Nielsen; Vertes & Eastman].

In this commentary the “incredibly robust” evidence for the relationship between sleep and procedural memory is questioned; inconsistencies in the existing data are pointed out. In addition, some suggestions about extending research are made, for example, studying REM sleep augmentation or memory consolidation in patients with sleep disorders. Last, the possibility of a relationship between dreaming and memory processes is discussed.

Whereas many researchers see a heuristic potential in the covert REM sleep model for explaining NREM sleep mentation and associated phenomena, many others are unconvinced of its value. At present, there is much circumstantial support for the model, but validation is lacking on many points. Supportive findings from several additional studies are summarized with results from two new studies showing (1) NREM mentation is correlated with duration of prior REM sleep, and (2) REM sleep signs (eye (...) movements, phasic EMG) occur frequently in NREM sleep. The covert REM sleep model represents one class of explanatory models that combines the two assumptions of mind-body isomorphism and a 1-gen mentation generator; its future development will depend largely upon a more detailed understanding of sleep state interactions and their contribution to mind-body isomorphisms. (shrink)

We believe that this has been a constructive debate on the topic of memory consolidation and REM sleep. It was a lively and spirited exchange – the essence of science. A number of issues were discussed including: the pedestal technique, stress, and early REMD work in animals; REM windows; the processing of declarative versus procedural memory in REM in humans; a mnemonic function for theta rhythm in waking but not in REM sleep; the lack of cognitive deficits in (...) patients on antidepressant drugs that suppress or eliminate REM sleep; the disposition of conscious (dreams) and nonconscious material of REM sleep; and finally our theory of REM sleep. Although our position was strongly challenged, we still hold that REM sleep serves no role in the processing and consolidation of memory. (shrink)

The paradigmatic assumption that REM sleep is the physiological equivalent of dreaming is in need of fundamental revision. A mounting body of evidence suggests that dreaming and REM sleep are dissociable states, and that dreaming is controlled by forebrain mechanisms. Recent neuropsychological, radiological, and pharmacological findings suggest that the cholinergic brain stem mechanisms that control the REM state can only generate the psychological phenomena of dreaming through the mediation of a second, probably dopaminergic, forebrain mechanism. The latter mechanism (...) (and thus dreaming itself) can also be activated by a variety of nonREM triggers. Dreaming can be manipulated by dopamine agonists and antagonists with no concomitant change in REM frequency, duration, and density. Dreaming can also be induced by focal forebrain stimulation and by complex partial (forebrain) seizures during nonREM sleep, when the involvement of brainstem REM mechanisms is precluded. Likewise, dreaming is obliterated by focal lesions along a specific (probably dopaminergic) forebrain pathway, and these lesions do not have any appreciable effects on REM frequency, duration, and density. These findings suggest that the forebrain mechanism in question is the final common path to dreaming and that the brainstem oscillator that controls the REM state is just one of the many arousal triggers that can activate this forebrain mechanism. The “REM-on” mechanism (like its various NREM equivalents) therefore stands outside the dream process itself, which is mediated by an independent, forebrain “dream-on” mechanism. Key Words: acetylcholine; brainstem; dopamine; dreaming; forebrain; NREM; REM; sleep. (shrink)

If reports given on experimental awakenings validly represent mental activity that was underway before the awakening, REM sleep is neither necessary nor sufficient for dreaming. Another intuitively attractive hypothesis for its function – that REM consolidates or otherwise modifies memory traces acquired while awake – is not supported by the preponderant evidence. There is growing acceptance of the possibility that REM functions to support sleep rather than waking brain processes. [Hobson et al.; Nielsen; Solms; Vertes & Eastman].

In this chapter, we aimed at further characterizing the functional neuroanatomy of the human rapid eye movement (REM) sleep at the population level. We carried out a meta-analysis of a large dataset of positron emission tomography (PET) scans acquired during wakefulness, slow wave sleep and REM sleep, and focused especially on the brain areas in which the activity diminishes during REM sleep. Results show that quiescent regions are confined to the inferior and middle frontal cortex and (...) to the inferior parietal lobule. Providing a plausible explanation for some of the features of dream reports, these findings may help in refining the concepts, which try to account for human cognition during REM sleep. In particular, we discuss the significance of these results to explain the alteration in executive processes, episodic memory retrieval and self representation during REM sleep dreaming as well as the incorporation of external stimuli into the dream narrative. (shrink)

The validity of dream recall is discussed. What is the relation between the actual dream and its later reflection? Nielsen proposes differential sleep mentation, which is probably determined by dream accessibility. Solms argues that REM sleep and dreaming are double dissociable states. Dreaming occurs outside REM sleep when cerebral activation is high enough. That various active sleep states correlate with vivid dream reports implies that REM sleep and dreaming are single dissociable states. Vertes & Eastman (...) reject that REM sleep is involved in memory consolidation. Considerable evidence for this was obtained by REM deprivation studies with the dubious water tank technique. [Nielsen; Solms; Vertes & Eastman]. (shrink)

To test whether mental activities collected from non-REM sleep are influenced by REM sleep, we suppressed REM sleep using clomipramine 50 mg or placebo in the evening, in a double blind cross-over design, in 11 healthy young men. Subjects were awakened every hour and asked about their mental activity. The marked REM-sleep suppression induced by clomipramine did not substantially affect any aspects of dream recall . Since long, complex and bizarre dreams persist even after suppressing REM (...) sleep either partially or totally, it suggests that the generation of mental activity during sleep is independent of sleep stage. (shrink)

The advent of functional imaging has reinforced the attempts to define dreaming as a sleep state-dependent phenomenon. PET scans revealed major differences between nonREM sleep and REM sleep. However, because dreaming occurs throughout sleep, the common features of the two sleep states, rather than the differences, could help define the prerequisite for the occurrence of dreams. [Hobson et al.; Nielsen; Solms; Revonsuo; Vertes & Eastman].

Although the emotional and motivational characteristics of dreaming have figured prominently in folk and psychoanalytic conceptions of dream production, emotions have rarely been systematically studied, and motivation, never. Because emotions during sleep lack the somatic components of waking emotions, and they change as the sleeper awakens, their properties are difficult to assess. Recent evidence of limbic system activation during REM sleep suggests a basis in brain architecture for the interaction of motivational and cognitive properties in dreaming. Motivational and (...) emotional content in REM and NREM laboratory mentation reports from 25 participants were compared. Motivational and emotional content was significantly greater in REM than NREM sleep, even after controlling for the greater word count of REM reports. (shrink)

Llewellyn has written a fascinating article about rapid eye movement dreams and how they promote the elaborative encoding of recent memories. The main message of her article is that hyperassociative and fluid cognitive processes during REM dreaming facilitate consolidation. We consider one potential implication of this analysis: the possibility that excessive or out-of-phase REM sleep fuels dissociative symptomatology. Further research is warranted to explore the psychopathological ramifications of Llewellyn's theory.

REM sleep behaviour disorder is a REM sleep-related parasomnia which may be considered a “dissociated state of wakefulness and sleep”, given that conflicting elements of REM sleep and of wakefulness coexist during the episodes, leading to motor and behavioural manifestations reminiscent of an enacted dream. RBD has been reported in association with α-synucleinopathies: around a third of patients with Parkinson’s disease have full-blown RBD.Recent data indicate that PD patients with RBD are more prone to hallucinations than (...) PD patients without this parasomnia. However it is still not clear why RBD in PD is associated with an increased prevalence of VHs.Data exist which suggest that visual hallucinations in PD may be the result of untimely intrusions of REM visual imagery into wakefulness. RBD, which is characterised by a REM sleep dissociation pattern, might be a condition that particularly favours such intrusions. However, other hypotheses may be advanced. In fact, deficits in attentional, executive, visuoperceptual and visuospatial abilities have been documented in RBD and found to occur far more frequently in PD with RBD than in PD without RBD. Neuropsychological deficits involving visual perception and attentional processes are thought to play an important role in the pathophysiology of VHs. On this basis, RBD in PD could be viewed as a contributory risk factor for VHs. (shrink)

Although the dream narrative, of (bio)logical necessity, originates with the dreamer, s/he typically does not know this. For the dreamer, the dream world is the real world. In this article I argue that this nightly misattribution is best explained in terms of the concept of mental ownership (e.g., Albahari, 2006; Klein, 2015a; Lane, 2012). Specifically, the exogenous nature of the dream narrative is the result of an individual assuming perspectival, but not personal, ownership of content s/he authored (i.e., “The content (...) in my head is not mine. Therefore it must be peripherally perceived”). Situating explanation within a theoretical space designed to address questions pertaining to the experienced origins of conscious content has a number of salutatory consequences. For example, it promotes predictive fecundity by bringing to light empirical generalizations whose presence otherwise might have gone unnoticed (e.g., the severely limited role of mental time travel within the dream narrative). -/- . (shrink)

We present evidence disputing the hypothesis that memories are processed or consolidated in REM sleep. A review of REM deprivation (REMD) studies in animals shows these reports to be about equally divided in showing that REMD does, or does not, disrupt learning/memory. The studies supporting a relationship between REM sleep and memory have been strongly criticized for the confounding effects of very stressful REM deprivation techniques. The three major classes of antidepressant drugs, monoamine oxidase inhibitors (MAOIs), tricyclic antidepressants (...) (TCAs), and selective serotonin reuptake inhibitors (SSRIs), profoundly suppress REM sleep. The MAOIs virtually abolish REM sleep, and the TCAs and SSRIs have been shown to produce immediate (40–85%) and sustained (30–50%) reductions in REM sleep. Despite marked suppression of REM sleep, these classes of antidepressants on the whole do not disrupt learning/memory. There have been a few reports of patients who have survived bilateral lesions of the pons with few lingering complications. Although these lesions essentially abolished REM sleep, the patients reportedly led normal lives. Recent functional imaging studies in humans have revealed patterns of brain activity in REM sleep that are consistent with dream processes but not with memory consolidation. We propose that the primary function of REM sleep is to provide periodic endogenous stimulation to the brain which serves to maintain requisite levels of central nervous system (CNS) activity throughout sleep. REM is the mechanism used by the brain to promote recovery from sleep. We believe that the cumulative evidence indicates that REM sleep serves no role in the processing or consolidation of memory. Key Words: antidepressant drugs; brain stem lesions; dreams; functional imaging; memory consolidation; REM deprivation; REM sleep; theta rhythm. (shrink)

Methods Young healthy heterosexual couples underwent sleep-lab-based polysomnography of two sleeping arrangements: individual sleep and co-sleep. Individual and dyadic sleep parameters (i.e., synchronization of sleep stages) were collected. The latter were assessed using cross-recurrence quantification analysis. Additionally, subjective sleep quality, relationship characteristics, and chronotype were monitored. Data were analyzed comparing co-sleep vs. individual sleep. Interaction effects of the sleeping arrangement with gender, chronotype, or relationship characteristics were moreover tested. Results As compared to (...) sleeping individually, co-sleeping was associated with about 10% more REM sleep, less fragmented REM sleep (p = 0.008), longer undisturbed REM fragments (p = 0.0006), and more limb movements (p = 0.007). None of the other sleep stages was significantly altered. Social support interacted with sleeping arrangement in a way that individuals with suboptimal social support showed the biggest impact of the sleeping arrangement on REM sleep. Sleep architectures were more synchronized between partners during co-sleep (p = 0.005) even if wake phases were excluded (p = 0.022). Moreover, sleep architectures are significantly coupled across a lag of ± 5min. Depth of relationship represented an additional significant main effect regarding synchronization, reflecting a positive association between the two. Neither REM sleep nor synchronization was influenced by gender, chronotype, or other relationship characteristics. Conclusion Depending on the sleeping arrangement, couple's sleep architecture and synchronization show alterations that are modified by relationship characteristics. We discuss that these alterations could be part of a self-enhancing feedback loop of REM sleep and sociality and a mechanism through which sociality prevents mental illness. (shrink)

Numerous studies have replicated the finding of mentation in both rapid eye movement (REM) and nonrapid eye movement (NREM) sleep. However, two different theoretical models have been proposed to account for this finding: (1) a one-generator model, in which mentation is generated by a single set of processes regardless of physiological differences between REM and NREM sleep; and (2) a two-generator model, in which qualitatively different generators produce cognitive activity in the two states. First, research is reviewed demonstrating (...) conclusively that mentation can occur in NREM sleep; global estimates show an average mentation recall rate of about 50% from NREM sleep – a value that has increased substantially over the years. Second, nine different types of research on REM and NREM cognitive activity are examined for evidence supporting or refuting the two models. The evidence largely, but not completely, favors the two-generator model. Finally, in a preliminary attempt to reconcile the two models, an alternative model is proposed that assumes the existence of covert REM sleep processes during NREM sleep. Such covert activity may be responsible for much of the dreamlike cognitive activity occurring in NREM sleep. Key Words: cognition in sleep; dreaming; NREM sleep; REM sleep; sleep mentation. (shrink)

Sleep is mandatory in most animals that have the nervous system and is universally observed in model organisms ranging from the nematodes, zebrafish, to mammals. However, it is unclear whether different sleep states fulfill common functions and are driven by shared mechanisms in these different animal species. Mammals and birds exhibit two obviously distinct states of sleep, i.e., non-rapid eye movement sleep and rapid eye movement sleep, but it is unknown why sleep should be (...) so segregated. Studying sleep in other animal models might give us clues that help solve this puzzle. Recent studies suggest that REM sleep, or ancestral forms of REM sleep might be found in non-mammalian or -avian species such as reptiles. These observations suggest that REM sleep and NREM sleep evolved earlier than previously thought. In this review, we discuss the evolutionary origin of the distinct REM/NREM sleep states to gain insight into the mechanistic and functional reason for these two different types of sleep. (shrink)

Hallucinatory states are experienced not only in connection with drugs and psychopathologies but occur naturally and spontaneously across the human circadian cycle: Our nightly dreams bring multimodal experiences in the absence of adequate external stimuli. The current study proposes a new, tighter measure of these hallucinatory states: Sleep onset, REM sleep, and non-REM sleep are shown to differ with regard to motor imagery indicating interactions with a rich imaginative world, and cognitive agency that could enable sleepers to (...) recognize their hallucinatory state. Mentation reports from the different states were analysed quantitatively with regard to two grammatical–semantic constructs, motor agency and cognitive agency. The present results support earlier physiological and psychological evidence in revealing a decline in cognitive functions and an increase in simulated interactions with a hallucinatory world, en route to normal REM sleep. This leads us to introduce the hypothesis that REM sleep, which exhibits remarkably high levels of sensorimotor processes, may have evolved to serve as a virtual laboratory for the development and rehearsal of embodied cognition. The new measure of hallucinatory states presented here may also hold implications for the study of executive functions and cognitions, which might be interesting, for example, for the investigation of lucid dreaming. (shrink)

We present one of the first quantitative studies on auditory verbal experiences and auditory verbal agency voices or characters”) in healthy participants across states of consciousness. Tools of quantitative linguistic analysis were used to measure participants’ implicit knowledge of auditory verbal experiences and auditory verbal agencies, displayed in mentation reports from four different states. Analysis was conducted on a total of 569 mentation reports from rapid eye movement sleep, non-REM sleep, sleep onset, and waking. Physiology was controlled (...) with the nightcap sleep–wake mentation monitoring system. Sleep-onset hallucinations, traditionally at the focus of scientific attention on auditory verbal hallucinations, showed the lowest degree of VE and VA, whereas REM sleep showed the highest degrees. Degrees of different linguistic-pragmatic aspects of VE and VA likewise depend on the physiological states. The quantity and pragmatics of VE and VA are a function of the physiologically distinct state of consciousness in which they are conceived. (shrink)

As reported by Vertes & Eastman, convincing evidence rules out any role for REM sleep in memory consolidation. However, they do not provide convincing evidence for their claim that sleep in generaI – as opposed to REM sleep per se – has no influence on memory consolidation. Recent correlational data suggest that the number of NREM/REM cycles is associated with performance on a verbal recall task. [Vertes & Eastman].

An ongoing assumption made by sleep researchers is that since dreams are more often recalled on awakening from rapid eye movement sleep, dreams must occur more often during this stage of sleep. An alternative hypothesis is that cognition occurs throughout sleep, but the recall of this mentation differs on awakening. When a dream is not reported on awakening, there is no method of establishing whether it did not happen or was forgotten. The aim of the present (...) study was to investigate this issue using an eye movement signal verification technique. Participants were instructed to produce an EM signal whenever they heard a tone. Tones were presented at increasing volume during Stage 2 and REM sleep until EM signal verification was observed. Ninety seconds after signal verification, participants were awakened and asked if they remembered hearing the tone or responding with the EM signal. Such recollection of signal verified tone presentations was significantly less after Stage 2 sleep compared to REM sleep presentations. Furthermore, SVT recall was significantly correlated with reported dream recall frequency, suggesting the same processes involved in recalling SVTs might also underlie dream recall. (shrink)

We present one of the first quantitative studies on auditory verbal experiences (“hearing voices”) and auditory verbal agency (inner speech, and specifically “talking to (imaginary) voices or characters”) in healthy participants across states of consciousness. Tools of quantitative linguistic analysis were used to measure participants’ implicit knowledge of auditory verbal experiences (VE) and auditory verbal agencies (VA), displayed in mentation reports from four different states. Analysis was conducted on a total of 569 mentation reports from rapid eye movement (REM) (...) class='Hi'>sleep, non‐REM sleep, sleep onset, and waking. Physiology was controlled with the nightcap sleep–wake mentation monitoring system. Sleep‐onset hallucinations, traditionally at the focus of scientific attention on auditory verbal hallucinations, showed the lowest degree of VE and VA, whereas REM sleep showed the highest degrees. Degrees of different linguistic‐pragmatic aspects of VE and VA likewise depend on the physiological states. The quantity and pragmatics of VE and VA are a function of the physiologically distinct state of consciousness in which they are conceived. (shrink)

Primary and secondary processes are the foundational axes of the Freudian mental apparatus: one horizontally as a tendency to associate, the primary process, and one vertically as the ability for perspective taking, the secondary process. Primary process mentation is not only supposed to be dominant in the unconscious but also, for example, in dreams. The present study tests the hypothesis that the mental activity during REM-sleep has more characteristics of the primary process, while during non-REM-sleep more secondary process (...) operations take place. Because the solving of a rebus requires the ability to non-contexually condensate the literal reading of single stimuli into a new one, rebus solving is a primary process operation by excellence. In a replication of the dream-rebus study of Shevrin and Fisher (1967), a rebus, which consisted of an image of a comb (German: “Kamm”) and an image of a raft (German: “Floß”), resulting in the German rebus word “kampflos” (Engl.: without a struggle), was flashed subliminally (at 1 ms) to 20 participants before going to sleep. Upon consecutive awakenings participants were asked for a dream report, free associations and an image description. Based on objective association norms, there were significantly more conceptual associations referring to Kamm and Floß indexing secondary process mentation when subjects were awakened from non-REM sleep as compared to REM-awakenings. There were not significantly more rebus associations referring to kampflos indexing primary process mentation when awakened from REM-sleep as compared to non-REM awakenings. However, when the associations were scored on the basis of each subject’s individual norms, there was a rebus effect with more idiosyncratic rebus associations in awakenings after REM than after non-REM-sleep. Our results support the general idea that REM-sleep is characterized by primary process thinking, while non-REM-sleep mentation follows the rules of the secondary process. (shrink)