Research in the neurosciences continues to provide evidence that sleep plays a role in the processes of learning and memory. There is less of a consensus, however, regarding the precise stages of memory development during which sleep is considered a requirement, simply favorable, or not important. This article begins with an overview of recent studies regarding sleep and learning, predominantly in the procedural memory domain, and is measured against our current understanding of the mechanisms that govern memory formation. Based on (...) these considerations, I offer a new neurocognitive framework of procedural learning, consisting first of acquisition, followed by two specific stages of consolidation, one involving a process of stabilization, the other involving enhancement, whereby delayed learning occurs. Psychophysiological evidence indicates that initial acquisition does not rely fundamentally on sleep. This also appears to be true for the stabilization phase of consolidation, with durable representations, resistant to interference, clearly developing in a successful manner during time awake (or just time, per se). In contrast, the consolidation stage, resulting in additional/enhanced learning in the absence of further rehearsal, does appear to rely on the process of sleep, with evidence for specific sleep-stage dependencies across the procedural domain. Evaluations at a molecular, cellular, and systems level currently offer several sleep specific candidates that could play a role in sleep-dependent learning. These include the upregulation of select plasticity-associated genes, increased protein synthesis, changes in neurotransmitter concentration, and specific electrical events in neuronal networks that modulate synaptic potentiation. Key Words: consolidation; enhancement; learning; memory; plasticity; sleep; stabilization. (shrink)

Information about positions, from which differences in position are computed (as proposed in the vector-integration-to-endpoint model), provides a more plausible perceptual basis for the control of goal-directed arm movements than information about distance (as proposed in the kinematic model).

Neural Darwinism (ND) is a large scale selectionist theory of brain development and function that has been hypothesized to relate to consciousness. According to ND, consciousness is entailed by reentrant interactions among neuronal populations in the thalamocortical system (the ‘dynamic core’). These interactions, which permit high-order discriminations among possible core states, confer selective advantages on organisms possessing them by linking current perceptual events to a past history of value-dependent learning. Here, we assess the consistency of ND with 16 widely recognized (...) properties of consciousness, both physiological (for example, consciousness is associated with widespread, relatively fast, low amplitude interactions in the thalamocortical system), and phenomenal (for example, consciousness involves the existence of a private flow of events available only to the experiencing subject). While no theory accounts fully for all of these properties at present, we find that ND and its recent extensions fare well. (shrink)

The standard behavioral index for human consciousness is the ability to report events with accuracy. While this method is routinely used for scientific and medical applications in humans, it is not easy to generalize to other species. Brain evidence may lend itself more easily to comparative testing. Human consciousness involves widespread, relatively fast low-amplitude interactions in the thalamocortical core of the brain, driven by current tasks and conditions. These features have also been found in other mammals, which suggests that consciousness (...) is a major biological adaptation in mammals. We suggest more than a dozen additional properties of human consciousness that may be used to test comparative predictions. Such homologies are necessarily more remote in non-mammals, which do not share the thalamocortical complex. However, as we learn more we may be able to make “deeper” predictions that apply to some birds, reptiles, large-brained invertebrates, and perhaps other species. (shrink)

Several theories claim that dreaming is a random by-product of REM sleep physiology and that it does not serve any natural function. Phenomenal dream content, however, is not as disorganized as such views imply. The form and content of dreams is not random but organized and selective: during dreaming, the brain constructs a complex model of the world in which certain types of elements, when compared to waking life, are underrepresented whereas others are over represented. Furthermore, dream content is consistently (...) and powerfully modulated by certain types of waking experiences. On the basis of this evidence, I put forward the hypothesis that the biological function of dreaming is to simulate threatening events, and to rehearse threat perception and threat avoidance. To evaluate this hypothesis, we need to consider the original evolutionary context of dreaming and the possible traces it has left in the dream content of the present human population. In the ancestral environment human life was short and full of threats. Any behavioral advantage in dealing with highly dangerous events would have increased the probability of reproductive success. A dream-production mechanism that tends to select threatening waking events and simulate them over and over again in various combinations would have been valuable for the development and maintenance of threat-avoidance skills. Empirical evidence from normative dream content, children's dreams, recurrent dreams, nightmares, post traumatic dreams, and the dreams of hunter-gatherers indicates that our dream-production mechanisms are in fact specialized in the simulation of threatening events, and thus provides support to the threat simulation hypothesis of the function of dreaming. Key Words: dream content; dream function; evolution of consciousness; evolutionary psychology; fear; implicit learning; nightmares; rehearsal; REM; sleep; threat perception. (shrink)

Dynamic systems theory offers conceptual andmathematical tools for describing the performance ofneural systems at very different levels oforganization. Three aspects of the dynamic paradigmare discussed, namely neural rhythms, neural andmental development, and macroscopic brain theories andmodels.

The explanatory gap and theknowledge argument are rooted in the conflationof propositional and phenomenal knowledge. Thebasic knowledge argument is based on theconsideration that ``physical information'' aboutthe nervous system is unable to provide theknowledge of a ``color experience'' . The implication is that physicalism isincomplete or false because it leaves somethingunexplained. The problem with Jackson'sargument is that physical information has theform of highly symbolic propositional knowledgewhereas phenomenal knowledge consists in innateneurophysiological processes. In addition totheir fundamental epistemological differences,clinical, anatomical, pathological and brainimaging (...) studies demonstrate that phenomenal andpropositional knowledge are fundamentallydifferent neurobiological processes. Propositional knowledge is phylogeneticallynew, highly symbolic, culturally acquired,exclusively human and expressible in differentnatural and artificial languages. By contrast,phenomenal knowledge consists inqualitative experiences and phenomenal conceptsthat provide an internal, language-independentreference to the properties of objects and theneeds of the organism. Language andpropositional knowledge are exclusively humanattributes implemented in specific regions ofthe dominant hemisphere. This contrastssharply with the phylogenicallysensory areas that are common to animals andhumans, which implement qualitativeexperiences. Experiences are hard-wiredneurobiological processes that can neither betransmitted nor re-created through thesymbolism of propositions. Thus, I concludethat the fallacy in the explanatory gap and inthe knowledge argument is a fallacy ofequivocation that results from ignoringfundamental neurobiological differences betweenphenomenal and propositional knowledge. (shrink)

We argue that computation via quantum mechanical processes is irrelevant to explaining how brains produce thought, contrary to the ongoing speculations of many theorists. First, quantum effects do not have the temporal properties required for neural information processing. Second, there are substantial physical obstacles to any organic instantiation of quantum computation. Third, there is no psychological evidence that such mental phenomena as consciousness and mathematical thinking require explanation via quantum theory. We conclude that understanding brain function is unlikely to require (...) quantum computation or similar mechanisms. (shrink)

It is proposed that attention to an object requires the simultaneous activity of three brain regions that are interconnected by a triangular circuit. The regions are the cortical site of attentional expression, the thalamic enhancement structure, and the prefrontal area of control. It is also proposed that awareness of an object requires the additional component of attention directed to a representation of the self. The expression of attention to a self-representation may involve activations of cortical sites corresponding to the body (...) landscape and/or verbal-based memories of autobiographical episodes. As in the case of attention to objects, attention to the self is presumed to involve a triangular circuit. The proposed triangular circuit of attention is shown to be generally consistent with current knowledge of brain structures and with data from a broad range of experiments concerned with the functions of neurons in these structures. (shrink)

I examine critically the application of information‐theoretic ideas to biological communication during embryonic development and in the functioning central nervous system (CNS). I show that intercellular communication relies mostly on simple signals whose role is to effect a selection among predetermined cellular states. Hence, a crucial role is played by cellular memory, which stabilizes such states. Memory in cells is partly located in the nuclear DNA; no less important however is (phenotypic) memory lying in the cell's organelles and compartments. Because (...) of combinatorial effects in gene expression patterns, cell memory is an enormously powerful mechanism, which also underlies plasticity, and thus constitutes the factor unifying genetic determination, plasticity and learning. Communication in the CNS is analyzed in some detail: here, cellular memory is embodied in anatomy (i.e., cell shape) and neurochemistry. These are the major, relatively “static” factors affecting the routing of neural impulses in the adult CNS. In addition, however, faster channeling is also required: action potentials must be directed to their targets along a few of exponentially many paths, and this dynamic routing is crucial for proper operation. I suggest that collective oscillatory modes may play a role in solving this addressing problem, in the same way that the clock signal gates the operation of man‐made computers. BioEssays 25:699–708, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

The neuromodulatory effects of brain stimulation therapies notably involving repetitive transcranial magnetic stimulation on nocturnal sleep, which is critically disturbed in major depression and other neuropsychiatric disorders, remain largely undetermined. We have previously reported in major depression patients that prefrontal rTMS sessions enhanced their slow wave activity power, but not their sigma power which is related to sleep spindle activity, for electrodes located nearby the stimulation site. In the present study, we focused on measuring the spindle density to investigate cumulative (...) effects of prefrontal rTMS sessions on the sleep spindle activity. Fourteen male inpatients diagnosed with medication-resistant unipolar or bipolar depression were recruited and subjected to 10 daily rTMS sessions targeting the left dorsolateral prefrontal cortex. All-night polysomnography data was acquired at four time points: Adaptation, Baseline, Post-1, and Post-2. Clinical and cognitive evaluations were longitudinally performed at Baseline, Post-1, and Post-2 time points to explore associations with the spindle density changes. The PSG data from 12 of 14 patients was analyzed to identify sleep spindles across the sleep stages II–IV at four electrode sites: F3, F4, P3, and P4. Statistical analysis by two-way ANOVA revealed that spindle density at F3 increased at Post-1 but decreased at Post-2 time points. Moreover, the local and transient increase of spindle density at F3 was associated with the previously reported SWA power increase at F3, possibly reflecting a shared mechanism of thalamocortical synchronization locally enhanced by diurnal prefrontal rTMS sessions. Clinical and cognitive correlations were not observed in this dataset. These findings suggest that diurnal rTMS sessions transiently modulate nocturnal sleep spindle activity at the stimulation site, although clinical and cognitive effects of the local changes warrant further investigation. (shrink)

Various neurophysiological experiments have revealed remarkable correlations between cortical neuronal activity and subjective experiences. However, the mere presence of neuronal electrical activity does not appear to be sufficient to produce these experiences. It has been suggested that the explanation for the neural basis of consciousness might lie in understanding the reason that some types of neuronal activity possess subjective correlates and others do not. Here I propose and develop the idea that this difference may be caused by the existence of (...) an elementary nonarbitrary linkage between temporal or spatiotemporal patterns of neuronal activity and their subjective attributes. I also show how cortical neural circuits capable of generating experience-coding patterns could emerge during evolution and brain development, due to the presence of spontaneous stochastic neuronal activity and activity-dependent synaptic plasticity. This hypothesis leads to several testable predictions, principal among which is the idea that the neural correlates of consciousness are essentially innate and universal. (shrink)

Walker's target article proposes a refinement of the well known two-stage model of memory formation to explain the positive effects of sleep on consolidation. After a first stage in which a labile memory representation is formed, a further stabilisation of the memory trace takes place in the second stage, which is dependent on (REM) sleep. Walker has refined the latter stage into a stage in which a consolidation-based enhancement occurs. It is not completely clear what consolidation-based enhancement implies and how (...) it can be dissociated from a stage for memory-stabilisation. A more serious consideration, however, is whether a second stage in memory consolidation that is solely dependent on sleep, is really necessary. The classical, passive, interference theory is able to explain adequately the findings related to the effects of sleep and memory, and can lead perhaps better to an understanding of the highly variable data in this field. (shrink)

The neuroanatomical substrates controlling and regulating sleeping and waking, and thus consciousness, are located in the brain stem. Most crucial for bringing the brain into a state conducive for consciousness and information processing is the mesencephalic part of the brain stem. This part controls the state of waking, which is generally associated with a high degree of consciousness. Wakefulness is accompanied by a low-amplitude, high-frequency electroencephalogram, due to the fact that thalamocortical neurons fire in a state of tonic depolarization. Information (...) can easily pass the low-level threshold of these neurons, leading to a high transfer ratio. The complexity of the electroencephalogram during conscious waking is high, as expressed in a high correlation dimension. Accordingly, the level of information processing is high. Spindles, and alpha waves in humans, mark the transition from wakefulness to sleep. These phenomena are related to drowsiness, associated with a reduction in consciousness. Drowsiness occurs when cells undergo moderate hyperpolarizations. Increased inhibitions result in a reduction of afferent information, with a lowered transfer ratio. Information processing subsides, which is also expressed in a diminished correlation dimension. Consciousness is further decreased at the onset of slow wave sleep. This sleep is controlled by the medullar reticular formation and is characterized by a high-voltage, low-frequency electroencephalogram. Slow wave sleep becomes manifest when neurons undergo a further hyperpolarization. Inhibitory activities are so strong that the transfer ratio further drops, as does the correlation dimension. Thus, sensory information is largely blocked and information processing is on a low level. Finally, rapid eye movement sleep is regulated by the pontine reticular formation and is associated with a ''wake-like'' electroencephalographic pattern. Just as during wakefulness, this is the expression of a depolarization of thalamocortical neurons. The transfer ratio of rapid eye movement sleep has not yet been determined, but seems to vary. Evidence exists that this type of sleep, associated with dreaming, with some kind of perception and consciousness, is involved in processing of ''internal'' information. In line with this, rapid eye movement sleep has higher correlation dimensions than slow-wave sleep and sometimes even higher than wakefulness. It is assumed that the ''near-the-threshold'' depolarized state of neurons in the thalamus and cerebral cortex is a necessary condition for perceptual processes and consciousness, such as occurs during waking and in an altered form during rapid eye movement sleep. (shrink)

Philosophers have been talking about brain states for almost 50 years and as of yet no one has articulated a theoretical account of what one is. In fact this issue has received almost no attention and cognitive scientists still use meaningless phrases like 'C-fiber firing' and 'neuronal activity' when theorizing about the relation of the mind to the brain. To date when theorists do discuss brain states they usually do so in the context of making some other argument with the (...) result being that any discussion of what brain states are has a distinct en passant flavor. In light of this it is a goal of mine to make brain states the center of attention by providing some general discussion of them. I briefly look at the argument of Bechtel and Mundale, as I think that they expose a common misconception philosophers had about brain states early on. I then turn to briefly examining Polger's argument, as I think he offers an intuitive account of what we expect brain states to be as well as a convincing argument against a common candidate for knowledge about brain states that is currently "on the scene." I then introduce a distinction between brain states and states of the brain: Particular brain states occur against background states of the brain. I argue that brain states are patterns of synchronous neural firing, which reflects the electrical face of the brain; states of the brain are the gating and modulating of neural activity and reflect the chemical face of the brain. (shrink)

Surgical patients under anesthesia can wake up unpredictably and be exposed to intense, traumatic pain. Current medical techniques cannot maintain depth of anesthesia at a perfectly stable and safe level; the depth of unconsciousness may change from moment to moment. Without an effective consciousness monitor anesthesiologists may not be able to adjust dosages in time to protect patients from pain. An estimated 40,000 to 200,000 midoperative awakenings may occur in the United States annually. E. R. John and coauthors present the (...) scientific basis of a practical ''consciousness monitor'' in two articles. One article is empirical and shows widespread and consistent electrical field changes across subjects and anesthetic agents as soon as consciousness is lost; these changes reverse when consciousness is regained afterward. These findings form the basis of a surgical consciousness monitor that recently received approval from the U.S. Food and Drug Administration. This may be the first practical application of research on the brain basis of consciousness. The other John article suggests theoretical explanations at three levels, a neurophysiological account of anesthesia, a neural dynamic account of conscious and unconscious states, and an integrative field theory. Of these, the neurophysiology is the best understood. Neural dynamics is evolving rapidly, with several alternative points of view. The field theory sketched here is the most novel and controversial. (shrink)

A unifying theory of general anesthetic-induced unconsciousness must explain the common mechanism through which various anesthetic agents produce unconsciousness. Functional-brain-imaging data obtained from 11 volunteers during general anesthesia showed specific suppression of regional thalamic and midbrain reticular formation activity across two different commonly used volatile agents. These findings are discussed in relation to findings from sleep neurophysiology and the implications of this work for consciousness research. It is hypothesized that the essential common neurophysiologic mechanism underlying anesthetic-induced unconsciousness is, as with (...) sleep-induced unconsciousness, a hyperpolarization block of thalamocortical neurons. A model of anesthetic-induced unconsciousness is introduced to explain how the plethora of effects anesthetics have on cellular functioning ultimately all converge on a single neuroanatomic/neurophysiologic system, thus providing for a unitary physiologic theory of narcosis related to consciousness. (shrink)

This is a bibliography of books and articles on consciousness in philosophy, cognitive science, and neuroscience over the last 30 years. There are three main sections, devoted to monographs, edited collections of papers, and articles. The first two of these sections are each divided into three subsections containing books in each of the main areas of research. The third section is divided into 12 subsections, with 10 subject headings for philosophical articles along with two additional subsections for articles in cognitive (...) science and neuroscience. Of course the division is somewhat arbitrary, but I hope that it makes the bibliography easier to use. This bibliography has first been compiled by Thomas Metzinger and David Chalmers to appear in print in two philosophical anthologies on conscious experience . From 1995 onwards it has been continuously updated by Thomas Metzinger, and now is freely available as a PDF-, RTF-, or HTML-file. This bibliography mainly attempts to cover the Anglo-Saxon and German debates, in a non-annotated, fully formatted way that makes it easy to "cut and paste" from the original file. To a certain degree this bibliography also contains items in other languages than English and German - all submissions in other languages are welcome. Last update of current version: July 13th, 2001. (shrink)

1 INSERM-CEA Unit 562, Cognitive Neuroimaging, Service Hospitalier Fre´de´ric Joliot, Orsay, France, 2 CNRS URA2182 Re´cepteurs and Cognition, Institut Pasteur, Paris, France.

Different anesthetics are known to modulate different types of membrane-bound receptors. Their common mechanism of action is expected to alter the mechanism for consciousness. Consciousness is hypothesized as the integral of all the units of internal sensations induced by reactivation of inter-postsynaptic membrane functional LINKs during mechanisms that lead to oscillating potentials. The thermodynamics of the spontaneous lateral curvature of lipid membranes induced by lipophilic anesthetics can lead to the formation of non-specific inter-postsynaptic membrane functional LINKs by different mechanisms. These (...) include direct membrane contact by excluding the inter-membrane hydrophilic region and readily reversible partial membrane hemifusion. The constant reorganization of the lipid membranes at the lateral edges of the postsynaptic terminals (dendritic spines) resulting from AMPA receptor-subunit vesicle exocytosis and endocytosis can favor the effect of anesthetic molecules on lipid membranes at this location. Induction of a large number of non-specific LINKs can alter the conformation of the integral of the units of internal sensations that maintain consciousness. Anesthetic requirement is reduced in the presence of dopamine that causes enlargement of dendritic spines. Externally applied pressure can transduce from the middle ear through the perilymph, cerebrospinal fluid, and the recently discovered glymphatic pathway to the extracellular matrix space, and finally to the paravenular space. The pressure gradient reduce solubility and displace anesthetic molecules from the membranes into the paravenular space, explaining the pressure reversal of anesthesia. Changes in membrane composition and the conversion of membrane hemifusion to fusion due to defects in the checkpoint mechanisms can lead to cytoplasmic content mixing between neurons and cause neurodegenerative changes. The common mechanism of anesthetics presented here can operate along with the known specific actions of different anesthetics. (shrink)

During sleep, consciousness is reduced, associated with a diminished connection of the brain to the environment. This is due to the blocking of the thalamocortical pathway by inhibitory mechanisms. This “thalamic gating” during sleep, however, is not complete, but partial. The stream of information is considerably reduced, but the brain is not fully disconnected from the environment. An inspection of the environment takes place to optimize safety during sleep. Stimuli having a meaning for the individual, or signaling danger, are recognized, (...) and may enter into awareness, leading to a wake-up call, which allows the individual to react. This subconscious stimulus evaluation is regarded as having a guardian function for sleep. The recognition of stimuli during sleep points toward the presence of a sort of consciousness without awareness in the sleeping brain. (shrink)