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(Updated 4/26/04)

Bernard J. Baars, James Newman, J.G. Taylor
Neuronal mechanisms of consciousness:
A Relational Global Workspace framework.

(Pages 269-278 in S. Hameroff, A. Kaszniak, J. Laukes, Toward a Science of Consciousness II: The second Tucson discussions and debates. Cambridge, MA: MIT Press, 1998.)
"This paper explores a remarkable convergence of ideas and evidence, previously presented in separate places by its authors. That convergence has now become so persuasive that we believe we are working within substantially the same broad framework. Taylor’s mathematical papers on neuronal systems involved in consciousness dovetail well with work by Newman and Baars on the thalamocortical system, suggesting a brain mechanism much like the global workspace architecture developed by Baars (see references below). This architecture is relational, in the sense that it continuously mediates the interaction of input with memory. While our approaches overlap in a number of ways, each of us tends to focus on different areas of detail. What is most striking, and we believe significant, is the extent of consensus, which we believe to be consistent with other contemporary approaches by Weiskrantz, Gray, Crick and Koch, Edelman, Gazzaniga, Newell and colleagues, Posner, Baddeley, and a number of others. We suggest that cognitive neuroscience is moving toward a shared understanding of consciousness in the brain."
...
"The brain stem-thalamocortical axis supports the state, but not the detailed contents of consciousness, which are produced by cortex." [Full Text]

John G. Taylor
Constructing the Relational Mind
PSYCHE, 4(10), June 1998
"The "relational mind" approach to the inner content of consciousness is developed in terms of various control structures and processing strategies and their possible neurobiological identifications in brain sites. This leads naturally to a division of consciousness into a passive and an active part. A global control structure for the "single strand" aspect of consciousness is proposed as the thalamo-nucleus reticularis thalami-cortex coupled system, which is related to experimental data on the electrical stimulation of awareness. Local control, in terms of excitatory transfer from pre-processing sites to posterior working memory regions, is supported by data on subliminal perception timing and disambiguation of poorly defined percepts. The inner content of consciousness is understood as arising from the resulting relational features between inputs and stored pre-processing and episodic memories. Strong analogies are drawn between emergent properties of the model and suggested properties of "raw feels", supporting the thesis that working memories are the initial sites for the emergence of phenomenal awareness, and the frontal lobes for its further adumbration in terms of higher cognitive processing, including the creation of self." [Full Text]

Baars BJ, Franklin S.
How conscious experience and working memory interact.
Trends Cogn Sci 2003 Apr;7(4):166-172
"Active components of classical working memory are conscious, but traditional theory does not account for this fact. Global Workspace theory suggests that consciousness is needed to recruit unconscious specialized networks that carry out detailed working memory functions. The IDA model provides a fine-grained analysis of this process, specifically of two classical working-memory tasks, verbal rehearsal and the utilization of a visual image. In the process, new light is shed on the interactions between conscious and unconscious aspects of working memory." [Abstract]

Baars BJ.
Tutorial commentary: surprisingly small subcortical structures are needed for the state of waking consciousness, while cortical projection areas seem to provide perceptual contents of consciousness.
Conscious Cogn 1995 Jun;4(2):159-62
"The evidence can therefore be summarized as follows: (1) RF, nRt, and ILN activity seem to be necessary but not sufficient for conscious experience. (2) Stimulus representation in primary sensory projection areas also seems to be necessary but not sufficient for conscious perceptual experience (Weiskrantz, 1980). The simplest hypothesis is that both components are necessary and sufficient to support conscious perceptual experience." [Abstract] [RF = reticular formation; nRt = nucleus reticularis thalami; ILN = thalamic intralaminar nuclei]

Cho SB, Baars BJ, Newman J.
A Neural Global Workspace Model for Conscious Attention.
Neural Netw 1997 Oct 1;10(7):1195-1206
"Considerable progress is being made in interdisciplinary efforts to develop a general theory of the neural correlates of consciousness. Developments of Baars' Global Workspace theory over the past decade are examples of this progress. Integrating experimental data and models from cognitive psychology, AI and neuroscience, we present a neurocognitive model in which consciousness is defined as a global integration and dissemination system - nested in a large-scale, distributed array of specialized bioprocessors - which controls the allocation of the processing resources of the central nervous system. It is posited that this global control is effected via cortical 'gating' of a strategic thalamic nucleus. The basic circuitry of this neural system is reasonably well understood, and can be modeled, to a first approximation, employing neural network principles." [Abstract]

Bernard J. Baars
IN THE THEATRE OF CONSCIOUSNESS
Global Workspace Theory,
A Rigorous Scientific Theory of Consciousness.

Journal of Consciousness Studies, 4, No. 4, 1997, pp. 292-309
"An array of evidence is beginning to reveal the role of consciousness in the nervous system, at least in outline. Conscious experience seems to create access to many independent knowledge sources in the brain, most of them quite unconscious. Humans seem to have a larger repertoire of uses for consciousness — including language and long-term planning, self-monitoring and self-reflection, inner speech, metaphor, symbolic representation of experience and deliberate use of imagery. When it comes to sensory consciousness, however, the brain shows little difference between humans and many other mammals." [PDF]

Baars BJ.
How does a serial, integrated and very limited stream of consciousness emerge from a nervous system that is mostly unconscious, distributed, parallel and of enormous capacity?
Ciba Found Symp 1993;174:282-90; discussion 291-303
"Much of the nervous system can be viewed as a massively parallel, distributed system of highly specialized but unconscious processors. Conscious experience on the other hand is traditionally viewed as a serial stream that integrates different sources of information but is limited to only one internally consistent content at any given moment. Global Workspace theory suggests that conscious experience emerges from a nervous system in which multiple input processors compete for access to a broadcasting capability; the winning processor can disseminate its information globally throughout the brain. Global workspace architectures have been widely employed in computer systems to integrate separate modules when they must work together to solve a novel problem or to control a coherent new response. The theory articulates a series of increasingly complex models, able to account for more and more evidence about conscious functioning, from perceptual consciousness to conscious problem-solving, voluntary control of action, and directed attention. Global Workspace theory is consistent with, but not reducible to, other theories of limited-capacity mechanisms. Global workspace architectures must show competition for input to a neural global workspace and global distribution of its output. Brain structures that are demonstrably required for normal conscious experience can carry out these two functions. The theory makes testable predictions, especially for newly emerging, high-speed brain imaging technology." [Abstract]

Baars BJ.
The conscious access hypothesis: origins and recent evidence.
Trends Cogn Sci 2002 Jan 1;6(1):47-52
"Consciousness might help to mobilize and integrate brain functions that are otherwise separate and independent. Evidence for this 'conscious access hypothesis' was described almost two decades ago, in a framework called global workspace theory. The theory had little impact at first, for three reasons: because consciousness was controversial; the evidence, though extensive, was indirect; and integrative theory was unfashionable. Recent neuroimaging evidence appears broadly to support the hypothesis, which has implications for perception, learning, working memory, voluntary control, attention and self systems in the brain." [Abstract] [PDF]

Dehaene S, Naccache L.
Towards a cognitive neuroscience of consciousness: basic evidence and a workspace framework.
Cognition 2001 Apr;79(1-2):1-37
"This introductory chapter attempts to clarify the philosophical, empirical, and theoretical bases on which a cognitive neuroscience approach to consciousness can be founded. We isolate three major empirical observations that any theory of consciousness should incorporate, namely (1) a considerable amount of processing is possible without consciousness, (2) attention is a prerequisite of consciousness, and (3) consciousness is required for some specific cognitive tasks, including those that require durable information maintenance, novel combinations of operations, or the spontaneous generation of intentional behavior. We then propose a theoretical framework that synthesizes those facts: the hypothesis of a global neuronal workspace. This framework postulates that, at any given time, many modular cerebral networks are active in parallel and process information in an unconscious manner. An information becomes conscious, however, if the neural population that represents it is mobilized by top-down attentional amplification into a brain-scale state of coherent activity that involves many neurons distributed throughout the brain. The long-distance connectivity of these 'workspace neurons' can, when they are active for a minimal duration, make the information available to a variety of processes including perceptual categorization, long-term memorization, evaluation, and intentional action. We postulate that this global availability of information through the workspace is what we subjectively experience as a conscious state. A complete theory of consciousness should explain why some cognitive and cerebral representations can be permanently or temporarily inaccessible to consciousness, what is the range of possible conscious contents, how they map onto specific cerebral circuits, and whether a generic neuronal mechanism underlies all of them. We confront the workspace model with those issues and identify novel experimental predictions. Neurophysiological, anatomical, and brain-imaging data strongly argue for a major role of prefrontal cortex, anterior cingulate, and the areas that connect to them, in creating the postulated brain-scale workspace." [Abstract] [PDF]

Stanislas Dehaene, Michel Kerszberg, and Jean-Pierre Changeux
A neuronal model of a global workspace in effortful cognitive tasks
PNAS 95: 14529-14534, 1998.
"A minimal hypothesis is proposed concerning the brain processes underlying effortful tasks. It distinguishes two main computational spaces: a unique global workspace composed of distributed and heavily interconnected neurons with long-range axons, and a set of specialized and modular perceptual, motor, memory, evaluative, and attentional processors. Workspace neurons are mobilized in effortful tasks for which the specialized processors do not suffice. They selectively mobilize or suppress, through descending connections, the contribution of specific processor neurons. In the course of task performance, workspace neurons become spontaneously coactivated, forming discrete though variable spatio-temporal patterns subject to modulation by vigilance signals and to selection by reward signals. A computer simulation of the Stroop task shows workspace activation to increase during acquisition of a novel task, effortful execution, and after errors. We outline predictions for spatio-temporal activation patterns during brain imaging, particularly about the contribution of dorsolateral prefrontal cortex and anterior cingulate to the workspace." [Full Text]

Damasio AR.
Investigating the biology of consciousness.
Philos Trans R Soc Lond B Biol Sci 1998 Nov 29;353(1377):1879-82
"The fact that consciousness is a private, first-person phenomenon makes it more difficult to study than other cognitive phenomena that, although being equally private, also have characteristic behavioural signatures. Nonetheless, by combining cognitive and neurobiological methods, it is possible to approach consciousness, to describe its cognitive nature, its behavioural correlates, its possible evolutionary origin and functional role; last but not least, it is possible to investigate its neuroanatomical and neurophysiological underpinnings. In this brief essay I distinguish between two kinds of consciousness: core consciousness and extended consciousness. Core consciousness corresponds to the transient process that is incessantly generated relative to any object with which an organism interacts, and during which a transient core self and transient sense of knowing are automatically generated. Core consciousness requires neither language nor working memory, and needs only a brief short-term memory. Extended consciousness is a more complex process. It depends on the gradual build-up of an autobiographical self, a set of conceptual memories pertaining to both past and anticipated experiences of an individual, and it requires conventional memory. Extended consciousness is enhanced by language." [Abstract] [Full Text]

Hans C. Lou, Bruce Luber, Michael Crupain, Julian P. Keenan, Markus Nowak, Troels W. Kjaer, Harold A. Sackeim, and Sarah H. Lisanby
Parietal cortex and representation of the mental Self
PNAS published April 19, 2004, 10.1073/pnas.0400049101
"For a coherent and meaningful life, conscious self-representation is mandatory. Such explicit "autonoetic consciousness" is thought to emerge by retrieval of memory of personally experienced events ("episodic memory"). During episodic retrieval, functional imaging studies consistently show differential activity in medial prefrontal and medial parietal cortices. With positron-emission tomography, we here show that these medial regions are functionally connected and interact with lateral regions that are activated according to the degree of self-reference. During retrieval of previous judgments of Oneself, Best Friend, and the Danish Queen, activation increased in the left lateral temporal cortex and decreased in the right inferior parietal region with decreasing self-reference. Functionally, the former region was preferentially connected to medial prefrontal cortex, the latter to medial parietal. The medial parietal region may, then, be conceived of as a nodal structure in self-representation, functionally connected to both the right parietal and the medial prefrontal cortices. To determine whether medial parietal cortex in this network is essential for episodic memory retrieval with self-representation, we used transcranial magnetic stimulation over the region to transiently disturb neuronal circuitry. There was a decrease in the efficiency of retrieval of previous judgment of mental Self compared with retrieval of judgment of Other with transcranial magnetic stimulation at a latency of 160 ms, confirming the hypothesis. This network is strikingly similar to the network of the resting conscious state, suggesting that self-monitoring is a core function in resting consciousness." [Abstract]

Debra A. Gusnard, Erbil Akbudak, Gordon L. Shulman, and Marcus E. Raichle
Medial prefrontal cortex and self-referential mental activity: Relation to a default mode of brain function
PNAS 98: 4259-4264; published online before print as 10.1073/pnas.071043098
"Medial prefrontal cortex (MPFC) is among those brain regions having the highest baseline metabolic activity at rest and one that exhibits decreases from this baseline across a wide variety of goal-directed behaviors in functional imaging studies. This high metabolic rate and this behavior suggest the existence of an organized mode of default brain function, elements of which may be either attenuated or enhanced. Extant data suggest that these MPFC regions may contribute to the neural instantiation of aspects of the multifaceted "self." We explore this important concept by targeting and manipulating elements of MPFC default state activity. In this functional magnetic resonance imaging (fMRI) study, subjects made two judgments, one self-referential, the other not, in response to affectively normed pictures: pleasant vs. unpleasant (an internally cued condition, ICC) and indoors vs. outdoors (an externally cued condition, ECC). The ICC was preferentially associated with activity increases along the dorsal MPFC. These increases were accompanied by decreases in both active task conditions in ventral MPFC. These results support the view that dorsal and ventral MPFC are differentially influenced by attentiondemanding tasks and explicitly self-referential tasks. The presence of self-referential mental activity appears to be associated with increases from the baseline in dorsal MPFC. Reductions in ventral MPFC occurred consistent with the fact that attention-demanding tasks attenuate emotional processing. We posit that both self-referential mental activity and emotional processing represent elements of the default state as represented by activity in MPFC. We suggest that a useful way to explore the neurobiology of the self is to explore the nature of default state activity." [Full Text]

Zysset S, Huber O, Samson A, Ferstl EC, von Cramon DY.
Functional specialization within the anterior medial prefrontal cortex: a functional magnetic resonance imaging study with human subjects.
Neurosci Lett. 2003 Jan 2;335(3):183-6.
"This study investigated the functional neuroanatomy of the anterior medial prefrontal cortex (aMPFC). Previous studies have shown that the aMPFC is involved in evaluative judgment and self-referential processes. Specifically, different sections of the aMPFC are differentially influenced by attention demanding processes. Whereas the dorsal section is supposed to be involved in self-referential processes, the ventral section is assumed to be attenuated during attention demanding processes. The present study investigates the involvement of the dorsal and ventral aMPFC in evaluative judgment by using functional magnetic resonance imaging with spin-echo echo-planar-imaging. Processes involved in evaluative judgment are attention-demanding, self-referential and activate regions in the dorsal and ventral section of the aMPFC. Attention demanding tasks do not necessarily lead to an attenuation of the ventral section of the aMPFC, a region mainly involved in emotional and affective processing." [Abstract]

Reinders AA, Nijenhuis ER, Paans AM, Korf J, Willemsen AT, den Boer JA.
One brain, two selves.
Neuroimage. 2003 Dec;20(4):2119-25.
"Having a sense of self is an explicit and high-level functional specialization of the human brain. The anatomical localization of self-awareness and the brain mechanisms involved in consciousness were investigated by functional neuroimaging different emotional mental states of core consciousness in patients with Multiple Personality Disorder (i.e., Dissociative Identity Disorder (DID)). We demonstrate specific changes in localized brain activity consistent with their ability to generate at least two distinct mental states of self-awareness, each with its own access to autobiographical trauma-related memory. Our findings reveal the existence of different regional cerebral blood flow patterns for different senses of self. We present evidence for the medial prefrontal cortex (MPFC) and the posterior associative cortices to have an integral role in conscious experience." [Abstract]

Sterling C. Johnson, Leslie C. Baxter, Lana S. Wilder, James G. Pipe, Joseph E. Heiserman, and George P. Prigatano
Neural correlates of self-reflection

Brain 125: 1808-1814, 2002.
"The capacity to reflect on one’s sense of self is an important component of self-awareness. In this paper, we investigate some of the neurocognitive processes underlying reflection on the self using functional MRI. Eleven healthy volunteers were scanned with echoplanar imaging using the blood oxygen level-dependent contrast method. The task consisted of aurally delivered statements requiring a yes–no decision. In the experimental condition, participants responded to a variety of statements requiring knowledge of and reflection on their own abilities, traits and attitudes (e.g. ‘I forget important things’, ‘I’m a good friend’, ‘I have a quick temper’). In the control condition, participants responded to statements requiring a basic level of semantic knowledge (e.g. ‘Ten seconds is more than a minute’, ‘You need water to live’). The latter condition was intended to control for auditory comprehension, attentional demands, decision-making, the motoric response, and any common retrieval processes. Individual analyses revealed consistent anterior medial prefrontal and posterior cingulate activation for all participants. The overall activity for the group, using a random-effects model, occurred in anterior medial prefrontal cortex (t = 13.0, corrected P = 0.05; x, y, z, 0, 54, 8, respectively) and the posterior cingulate (t = 14.7, P = 0.02; x, y, z, –2, –62, 32, respectively; 967 voxel extent). These data are consistent with lesion studies of impaired awareness, and suggest that the medial prefrontal and posterior cingulate cortex are part of a neural system subserving self-reflective thought." [Abstract]

Wicker B, Ruby P, Royet JP, Fonlupt P.
A relation between rest and the self in the brain?
Brain Res Brain Res Rev. 2003 Oct;43(2):224-30.
"Neuroimaging techniques such as positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) are widely used to identify the cerebral correlates of cognitive tasks. The resting state presents the advantage to serve as a reference in all experiments but is also an ill-defined mental state because it may vary both from one subject to another and within the same subject. The most challenging question concerns the areas whose activity (revealed by PET or fMRI imaging) is greater in rest state than in an active condition. The present work reports the result of a meta-analysis including five previously published studies. The five different tasks involved are the following: attribution of intention, judgement of stimulus pleasantness, discrimination of spatial attributes, judgement of other peoples' belief and perception of gaze. For each study, the general linear model was used to assess statistical difference and a contrast resting state minus other conditions was calculated. The intersection of the five contrasts was used to search for the variation jointly observed across the different experiments. This lead to a reduced number of clusters: one cluster in the lower/anterior part of the cingulate gyrus and four clusters located in the medial/superior frontal gyrus, along the superior frontal sulcus. We discuss the location of these areas with respect to the location of activations induced by different tasks: externally focused attention, memory, general reasoning, theory of mind and self-referential tasks. We observed that medial prefrontal cortex exhibits a lower activity when the subject's attention is focused towards the external world than when the subject has to additionally refer to some internal states. By contrast, this activity is greater during resting state than during both externally directed and internally directed attention. Thus, we hypothesize that during rest, the subject is in a state where he refers only to his own self." [Abstract]

Vogeley K, Fink GR.
Neural correlates of the first-person-perspective.
Trends Cogn Sci 2003 Jan;7(1):38-42
"Human self-consciousness depends on the metarepresentation of mental and bodily states as one's own mental and bodily states. First-person-perspective taking is not sufficient, but necessary for human self-consciousness. To assign a first-person-perspective is to center one's own multimodal experiential space upon one's own body, thus operating in an egocentric reference frame. The brain regions involved in assigning first-person-perspective comprise medial prefrontal, medial parietal and lateral temporoparietal cortex. These empirical findings complement recent neurobiologically oriented theories of self-consciousness which focus on the relation between the subject and his/her environment by supplying a neural basis for its key components." [Abstract]

Frith C.
Attention to action and awareness of other minds.
Conscious Cogn. 2002 Dec;11(4):481-7.
"We have only limited awareness of the system by which we control our actions and this limited awareness does not seem to be concerned with the control of action. Awareness of choosing one action rather than another comes after the choice has been made, while awareness of initiating an action occurs before the movement has begun. These temporal differences bind together in consciousness the intention to act and the consequences of the action. This creates our sense of agency. Activity in the anterior cingulate cortex and medial prefrontal cortex is associated with awareness of our own actions and also occurs when we think about the actions of others. I propose that the mechanism underlying awareness of how our own intentions lead to actions can also be used to represent the intentions that underlie the actions of others. This common system enables us to communicate mental states and thereby share our experiences." [Abstract]

Frith CD, Frith U.
Interacting minds--a biological basis.
Science. 1999 Nov 26;286(5445):1692-5.
"The ability to "mentalize," that is to understand and manipulate other people's behavior in terms of their mental states, is a major ingredient in successful social interactions. A rudimentary form of this ability may be seen in great apes, but in humans it is developed to a high level. Specific impairments of mentalizing in both developmental and acquired disorders suggest that this ability depends on a dedicated and circumscribed brain system. Functional imaging studies implicate medial prefrontal cortex and posterior superior temporal sulcus (STS) as components of this system. Clues to the specific function of these components in mentalizing come from single cell recording studies: STS is concerned with representing the actions of others through the detection of biological motion; medial prefrontal regions are concerned with explicit representation of states of the self. These observations suggest that the ability to mentalize has evolved from a system for representing actions." [Abstract]

Kelley WM, Macrae CN, Wyland CL, Caglar S, Inati S, Heatherton TF.
Finding the self? An event-related fMRI study.
J Cogn Neurosci. 2002 Jul 1;14(5):785-94.
"Researchers have long debated whether knowledge about the self is unique in terms of its functional anatomic representation within the human brain. In the context of memory function, knowledge about the self is typically remembered better than other types of semantic information. But why does this memorial effect emerge? Extending previous research on this topic (see Craik et al., 1999), the present study used event-related functional magnetic resonance imaging to investigate potential neural substrates of self-referential processing. Participants were imaged while making judgments about trait adjectives under three experimental conditions (self-relevance, other-relevance, or case judgment). Relevance judgments, when compared to case judgments, were accompanied by activation of the left inferior frontal cortex and the anterior cingulate. A separate region of the medial prefrontal cortex was selectively engaged during self-referential processing. Collectively, these findings suggest that self-referential processing is functionally dissociable from other forms of semantic processing within the human brain." [Abstract]

Johnson SC, Baxter LC, Wilder LS, Pipe JG, Heiserman JE, Prigatano GP.
Neural correlates of self-reflection.
Brain. 2002 Aug;125(Pt 8):1808-14.
"The capacity to reflect on one's sense of self is an important component of self-awareness. In this paper, we investigate some of the neurocognitive processes underlying reflection on the self using functional MRI. Eleven healthy volunteers were scanned with echoplanar imaging using the blood oxygen level-dependent contrast method. The task consisted of aurally delivered statements requiring a yes-no decision. In the experimental condition, participants responded to a variety of statements requiring knowledge of and reflection on their own abilities, traits and attitudes (e.g. 'I forget important things', 'I'm a good friend', 'I have a quick temper'). In the control condition, participants responded to statements requiring a basic level of semantic knowledge (e.g. 'Ten seconds is more than a minute', 'You need water to live'). The latter condition was intended to control for auditory comprehension, attentional demands, decision-making, the motoric response, and any common retrieval processes. Individual analyses revealed consistent anterior medial prefrontal and posterior cingulate activation for all participants. The overall activity for the group, using a random-effects model, occurred in anterior medial prefrontal cortex (t = 13.0, corrected P = 0.05; x, y, z, 0, 54, 8, respectively) and the posterior cingulate (t = 14.7, P = 0.02; x, y, z, -2, -62, 32, respectively; 967 voxel extent). These data are consistent with lesion studies of impaired awareness, and suggest that the medial prefrontal and posterior cingulate cortex are part of a neural system subserving self-reflective thought." [Abstract]

Maguire EA, Mummery CJ.
Differential modulation of a common memory retrieval network revealed by positron emission tomography.
Hippocampus. 1999;9(1):54-61.
"Functional neuroimaging is uniquely placed to examine the dynamic nature of normal human memory, the distributed brain networks that support it, and how they are modulated. Memory has traditionally been classified into context-specific memories personally experienced ("episodic memory") and impersonal non-context-specific memories ("semantic memory"). However, we suggest that another useful distinction is whether events are personally relevant or not. Typically the factors of personal relevance and temporal context are confounded, and it is as yet not clear the precise influence of either on how memories are stored or retrieved. Here we focus on the retrieval of real-world memories unconfounding personal relevance and temporal context during positron emission tomography (PET) scanning. Memories differed along two dimensions: They were personally relevant (or not) and had temporal specificity (or not). Recollection of each of the resultant four memory subtypes-autobiographical events, public events, autobiographical facts, and general knowledge-was associated with activation of a common network of brain regions. Within this system, however, enhanced activity was observed for retrieval of personally relevant, time-specific memories in left hippocampus, medial prefrontal cortex, and left temporal pole. Bilateral temporoparietal junctions were activated preferentially for personal memories, regardless of time specificity. Finally, left parahippocampal gyrus, left anterolateral temporal cortex, and posterior cingulate cortex were involved in memory retrieval irrespective of person or time. Our findings suggest that specializations in memory retrieval result from associations between subsets of regions within a common network. We believe that these findings throw new light on an old debate surrounding episodic and declarative theories of memory and the precise involvement of the hippocampus." [Abstract]

Bernard J. Baars
Understanding Subjectivity: Global Workspace Theory and the Resurrection of the Observing Self
Journal of Consciousness Studies, 3, No. 3, 1996, pp. 211-16
"Why is the problem of subjectivity so hard, as David Chalmers claims? This essay suggests that it becomes hard when we adopt an implausible, perfectionistic standard. In the last two decades the standard has come to be 'observer empathy' -- the ability to know what it's like to be a bat or another human. That makes understanding consciousness difficult indeed. Far more practical criteria are used every day in medicine and scientific studies of consciousness, and indeed traditional philosophy from Kant to James took a much more relaxed view of subjectivity. Once we adopt these more workable standards, subjectivity is suddenly revealed to involve a familiar concept, namely 'the self as observer' of conscious experiences. Contrary to some, this sense of self is conceptually coherent and well-supported by hard evidence. For example, the 'left-hemisphere interpreter' in split-brain patients behaves as one such self. Given a modest and practical approach, we can expect to make progress toward understanding subjectivity." [Full Text]

Adam Zeman
Consciousness
Brain 124: 1263-1289, 2001.
"Consciousness is topical, for reasons including its renewed respectability among psychologists, rapid progress in the neuroscience of perception, memory and action, advances in artificial intelligence and dissatisfaction with the dualistic separation of mind and body. Consciousness is an ambiguous term. It can refer to (i) the waking state; (ii) experience; and (iii) the possession of any mental state. Self-consciousness is equally ambiguous, with senses including (i) proneness to embarrassment in social settings; (ii) the ability to detect our own sensations and recall our recent actions; (iii) self-recognition; (iv) the awareness of awareness; and (v) self-knowledge in the broadest sense. The understanding of states of consciousness has been transformed by the delineation of their electrical correlates, of structures in brainstem and diencephalon which regulate the sleep–wake cycle, and of these structures' cellular physiology and regional pharmacology. Clinical studies have defined pathologies of wakefulness: coma, the persistent vegetative state, the `locked-in' syndrome, akinetic mutism and brain death. Interest in the neural basis of perceptual awareness has focused on vision. Increasingly detailed neuronal correlates of real and illusory visual experience are being defined. Experiments exploiting circumstances in which visual experience changes while external stimulation is held constant are tightening the experimental link between consciousness and its neural correlates. Work on unconscious neural processes provides a complementary approach. `Unperceived' stimuli have detectable effects on neural events and subsequent action in a range of circumstances: blindsight provides the classical example. Other areas of cognitive neuroscience also promise experimental insights into consciousness, in particular the distinctions between implicit and explicit memory and deliberate and automatic action. Overarching scientific theories of consciousness include neurobiological accounts which specify anatomical or physiological mechanisms for awareness, theories focusing on the role played by conscious processes in information processing and theories envisaging the functions of consciousness in a social context. Whether scientific observation and theory will yield a complete account of consciousness remains a live issue. Physicalism, functionalism, property dualism and dual aspect theories attempt to do justice to three central, but controversial, intuitions about experience: that it is a robust phenomenon which calls for explanation, that it is intimately related to the activity of the brain and that it has an important influence on behaviour." [Full Text]

Dennett D.
Are we explaining consciousness yet?
Cognition 2001 Apr;79(1-2):221-37
"Theorists are converging from quite different quarters on a version of the global neuronal workspace model of consciousness, but there are residual confusions to be dissolved. In particular, theorists must resist the temptation to see global accessibility as the cause of consciousness (as if consciousness were some other, further condition); rather, it is consciousness. A useful metaphor for keeping this elusive idea in focus is that consciousness is rather like fame in the brain. It is not a privileged medium of representation, or an added property some states have; it is the very mutual accessibility that gives some informational states the powers that come with a subject's consciousness of that information. Like fame, consciousness is not a momentary condition, or a purely dispositional state, but rather a matter of actual influence over time. Theorists who take on the task of accounting for the aftermath that is critical for consciousness often appear to be leaving out the Subject of consciousness, when in fact they are providing an analysis of the Subject, a necessary component in any serious theory of consciousness." [Abstract] [Full Text]

Publications by Daniel Dennett

Searle JR.
How to study consciousness scientifically.
Philos Trans R Soc Lond B Biol Sci 1998 Nov 29;353(1377):1935-42
"The neurosciences have advanced to the point that we can now treat consciousness as a scientific problem like any other. The problem is to explain how brain processes cause consciousness and how consciousness is realized in the brain. Progress is impeded by a number of philosophical mistakes, and the aim of this paper is to remove nine of those mistakes: (i) consciousness cannot be defined; (ii) consciousness is subjective but science is objective; (iii) brain processes cannot explain consciousness; (iv) the problem of 'qualia' should be set aside; (v) consciousness is epiphenomenal; (vi) consciousness has no evolutionary function; (vii) a causal account of consciousness is necessarily dualistic; (viii) science is reductionistic, so a scientific account of consciousness would show it reducible to something else; and (ix) an account of consciousness must be an information processing account." [Full Text]

Searle, John R.
Consciousness
Annu. Rev. Neurosci. 2000 23: 557-578
"Until recently, most neuroscientists did not regard consciousness as a suitable topic for scientific investigation. This reluctance was based on certain philosophical mistakes, primarily the mistake of supposing that the subjectivity of consciousness made it beyond the reach of an objective science. Once we see that consciousness is a biological phenomenon like any other, then it can be investigated neurobiologically. Consciousness is entirely caused by neurobiological processes and is realized in brain structures. The essential trait of consciousness that we need to explain is unified qualitative subjectivity. Consciousness thus differs from other biological phenomena in that it has a subjective or first-person ontology, but this subjective ontology does not prevent us from having an epistemically objective science of consciousness. We need to overcome the philosophical tradition that treats the mental and the physical as two distinct metaphysical realms. Two common approaches to consciousness are those that adopt the building block model, according to which any conscious field is made of its various parts, and the unified field model, according to which we should try to explain the unified character of subjective states of consciousness. These two approaches are discussed and reasons are given for preferring the unified field theory to the building block model. Some relevant research on consciousness involves the subjects of blindsight, the split-brain experiments, binocular rivalry, and gestalt switching." [Abstract]

David J. Chalmers
On the Search for the Neural Correlate of Consciousness
Toward a Science of Consciousness II: The Second Tucson Discussions and Debates (S. Hameroff, A. Kaszniak, and A.Scott, eds), published with MIT Press in 1998
"Once one recognizes the central role that pre-experimental assumptions play in the search for the NCC, one realizes that there are some limitations on just what we can expect this search to tell us. Still, whether or not the NCC is the Holy Grail, I hope that I have said enough to make it clear that the quest for it is likely to enhance our understanding considerably. And I hope to have convinced you that there are important ways in which philosophy and neuroscience can come together to help clarify some of the deep problems involved in the study of consciousness." [Full Text]

David J. Chalmers
What is a Neural Correlate of Consciousness?
Neural Correlates of Consciousness: Empirical and Conceptual Questions (T. Metzinger, ed), published with MIT Press in 2000
"The search for neural correlates of consciousness (or NCCs) is arguably the cornerstone in the recent resurgence of the science of consciousness. The search poses many difficult empirical problems, but it seems to be tractable in principle, and some ingenious studies in recent years have led to considerable progress. A number of proposals have been put forward concerning the nature and location of neural correlates of consciousness." [Full Text]

Online papers on consciousness, part 3: Science of consciousness
Compiled by David Chalmers

Consciousness and the Brain
Annotated Biography

Compiled by Ralph D. Ellis and Natika Newton

“Consciousness”
Selected Bibliography
1970 - 2003
Compiled by Thomas Metzinger

Science And Consciousness Review

Crick F, Koch C.
Consciousness and Neuroscience
Cerebral Cortex, 8:97-107, 1998
"The explanation of consciousness is one of the major unsolved problems of modern science. After several thousand years of speculation, it would be very gratifying to find an answer to it." [Full Text]

Perner J, Dienes Z.
Developmental aspects of consciousness: How much theory of mind do you need to be consciously aware?
Conscious Cogn 2003 Mar;12(1):63-82
"When do children become consciously aware of events in the world? Five possible strategies are considered for their usefulness in determining the age in question. Three of these strategies ask when children show signs of engaging in activities for which conscious awareness seems necessary in adults (verbal communication, executive control, explicit memory), and two of the strategies consider when children have the ability to have the minimal form of higher-order thought necessary for access consciousness and phenomenal consciousness, respectively. The tentative answer to the guiding question is that children become consciously aware between 12 and 15 months (+/-3 months)." [Abstract]

Cooney JW, Gazzaniga MS.
Neurological disorders and the structure of human consciousness.
Trends Cogn Sci 2003 Apr;7(4):161-165
"Recent studies that identify distinct neural correlates of perceptual awareness offer a promising step towards improved understanding of the neurological underpinnings of conscious experience. Such studies indicate that perceptual awareness is modular in nature, with neural correlates of awareness consisting of the specialized structures involved in perceptual processing. However, the integrative, multimodal nature of conscious experience appears to require a functional architecture that overcomes this modular segregation of function. We propose a model in which experience emerges from the dynamic interactions of specialized component processes via a distributed neural network. Such a model offers a mechanism to explain several empirical observations of the neural correlates of perceptual awareness, cognitive function, and symptoms of neurological damage." [Abstract]

Taylor JG.
The central role of the parietal lobes in consciousness.
Conscious Cogn 2001 Sep;10(3):379-417
"There are now various approaches to understand where and how in the brain consciousness arises from neural activity, none of which is universally accepted. Difficulties among these approaches are reviewed, and a missing ingredient is proposed here to help adjudicate between them, that of "perspectivalness." In addition to a suitable temporal duration and information content of the relevant bound brain activity, this extra component is posited as being a further important ingredient for the creation of consciousness from neural activity. It guides the development of what is termed the "Central Representation," which is supposed to be present in all mammals and extended in humans to support self-consciousness as well as phenomenal consciousness. Experimental evidence and a theoretical framework for the existence of the central representation are presented, which relates the extra component to specific buffer working memory sites in the inferior parietal lobes, acting as attentional coordinators on the spatial maps making up the central representation. The article closes with a discussion of various open questions." [Abstract]

J.G. Taylor
Paying Attention to Consciousness
Trends in Cognitive Sciences Vol. 6 No.5 May 2002
"Despite being much studied by cognitive neuroscience, consciousness has resisted attempts to understand it. Recent neuroscientific papers on the problem have surprisingly neglected attention as a guide to consciousness. A new neural mechanism is proposed here, guided by a control approach to attention, which identifies the source of consciousness, especially that of the ownership of experience." [PDF]

J.G. Taylor
From Matter To Mind
Journal of Consciousness Studies 2002 Apr;9(4):3-22
"The relation between mind and matter is considered in terms of recent ideas from both phenomenology and brain science. Phenomenology is used to give clues to help bridge the brain–mind gap by providing constraints on any underlying neural architecture suggested from brain science. A tentative reduction of mind to matter is suggested and used to explain various features of phenomenological experience and of ownership of conscious experience. The crucial mechanism is the extended duration of the corollary discharge of attention movement, with its gating of activity for related content. Aspects of experience considered in terms of the model are the discontinuous nature of consciousness, immunity to error through misidentification, and the state of ‘pure’ consciousness as experienced through meditation. Corollary discharge of attention movement is proposed as the key idea bringing together basic features of meditation, consciousness and neuroscience, and helping to bridge the gap between mind and matter." [Abstract]

Journal of Consciousness Studies



Rosenthal DM.
How many kinds of consciousness?
Conscious Cogn 2002 Dec;11(4):653-65
"Ned Block's influential distinction between phenomenal and access consciousness has become a staple of current discussions of consciousness. It is not often noted, however, that his distinction tacitly embodies unargued theoretical assumptions that favor some theoretical treatments at the expense of others. This is equally so for his less widely discussed distinction between phenomenal consciousness and what he calls reflexive consciousness. I argue that the distinction between phenomenal and access consciousness, as Block draws it, is untenable. Though mental states that have qualitative character plainly differ from those with no mental qualities, a mental state's being conscious is the same property for both kinds of mental state. For one thing, as Block describes access consciousness, that notion does not pick out any property that we intuitively count as a mental state's being conscious. But the deeper problem is that Block's notion of phenomenal consciousness, or phenomenality, is ambiguous as between two very different mental properties. The failure to distinguish these results in the begging of important theoretical questions. Once the two kinds of phenomenality have been distinguished, the way is clear to explain qualitative consciousness by appeal to a model such as the higher-order-thought hypothesis." [Abstract]

Faw B.
Pre-frontal executive committee for perception, working memory, attention, long-term memory, motor control, and thinking: A tutorial review.
Conscious Cogn 2003 Mar;12(1):83-139
"As an explicit organizing metaphor, memory aid, and conceptual framework, the prefrontal cortex may be viewed as a five-member 'Executive Committee,' as the prefrontal-control extensions of five sub-and-posterior-cortical systems: (1) the 'Perceiver' (dominant-right-hemisphere ventral-lateral prefrontal cortex-VL/PERC-PFC) is the frontal extension of the ventral perceptual stream (the VL/PERC system) which represents the world and self in object coordinates; (2) the 'Verbalizer' (dominant-left-hemisphere ventral-lateral prefrontal cortex system-VL/VERB-PFC) is the frontal extension of the language stream (the VL/VERB system) which represents the world and self in language coordinates; (3) the 'Motivator' (ventral/medial-orbital pre-frontal cortex-VMO-PFC) is the frontal cortical extension of a subcortical extended-amygdala stream (the VMO system) which represents the world and self in motivational/emotional coordinates; (4) the 'Attender' (dorsal-medial/anterior cingulate-DM/AC-PFC) is the frontal cortical extension of a subcortical extended-hippocampal stream (the DM/AC system) which represents the world and self in spatiotemporal coordinates and directs attention to internal and external events; and (5) the 'Coordinator' (the dorsolateral prefrontal cortex-DL-PFC) is the frontal extension of the dorsal perceptual stream (the DL system) which represents the world and self in body- and eye-coordinates and controls willed action and working memory. This tutorial review examines the interacting roles of these five systems in perception, working memory, attention, long-term memory, motor control, and thinking." [Abstract]

M. Steriade
Impact of Network Activities on Neuronal Properties in Corticothalamic Systems
J Neurophysiol 86: 1-39, 2001.
"Data from in vivo and in vitro experiments are discussed to emphasize that synaptic activities in neocortex and thalamus have a decisive impact on intrinsic neuronal properties in intact-brain preparations under anesthesia and even more so during natural states of vigilance. Thus the firing patterns of cortical neuronal types are not inflexible but may change with the level of membrane potential and during periods rich in synaptic activity. The incidences of some cortical cell classes (defined by their responses to depolarizing current pulses) are different in isolated cortical slabs in vivo or in slices maintained in vitro compared with the intact cortex of naturally awake animals. Network activities, which include the actions of generalized modulatory systems, have a profound influence on the membrane potential, apparent input resistance, and backpropagation of action potentials. The analysis of various oscillatory types leads to the conclusion that in the intact brain, there are no "pure" rhythms, generated in simple circuits, but complex wave sequences (consisting of different, low- and fast-frequency oscillations) that result from synaptic interactions in corticocortical and corticothalamic neuronal loops under the control of activating systems arising in the brain stem core or forebrain structures. As an illustration, it is shown that the neocortex governs the synchronization of network or intrinsically generated oscillations in the thalamus. The rhythmic recurrence of spike bursts and spike trains fired by thalamic and cortical neurons during states of decreased vigilance may lead to plasticity processes in neocortical neurons. If these phenomena, which may contribute to the consolidation of memory traces, are not constrained by inhibitory processes, they induce seizures in which the neocortex initiates the paroxysms and controls their thalamic reflection. The results indicate that intact-brain preparations are necessary to investigate global brain functions such as behavioral states of vigilance and paroxysmal activities." [Full Text]

Steriade M.
Corticothalamic resonance, states of vigilance and mentation.
Neuroscience 2000;101(2):243-76
"During various states of vigilance, brain oscillations are grouped together through reciprocal connections between the neocortex and thalamus. The coherent activity in corticothalamic networks, under the control of brainstem and forebrain modulatory systems, requires investigations in intact-brain animals. During behavioral states associated with brain disconnection from the external world, the large-scale synchronization of low-frequency oscillations is accompanied by the inhibition of synaptic transmission through thalamocortical neurons. Despite the coherent oscillatory activity, on the functional side there is dissociation between the thalamus and neocortex during slow-wave sleep. While dorsal thalamic neurons undergo inhibitory processes due to the prolonged spike-bursts of thalamic reticular neurons, the cortex displays, periodically, a rich spontaneous activity and preserves the capacity to process internally generated signals that dominate the state of sleep. In vivo experiments using simultaneous intracellular recordings from thalamic and cortical neurons show that short-term plasticity processes occur after prolonged and rhythmic spike-bursts fired by thalamic and cortical neurons during slow-wave sleep oscillations. This may serve to support resonant phenomena and reorganize corticothalamic circuitry, determine which synaptic modifications, formed during the waking state, are to be consolidated and generate a peculiar kind of dreaming mentation. In contrast to the long-range coherent oscillations that occur at low frequencies during slow-wave sleep, the sustained fast oscillations that characterize alert states are synchronized over restricted territories and are associated with discrete and differentiated patterns of conscious events." [Abstract]

Niedermeyer E.
Electrophysiology of the frontal lobe.
Clin Electroencephalogr 2003 Jan;34(1):5-12
"The electrophysiology of the frontal lobe appears to be unimpressive when the view is limited to the routine EEG recording of a healthy waking adult. There is usually low voltage fast activity, which becomes more pronounced when recorded with depth leads. Three special EEG patterns of marginal to slightly abnormal character are discussed: a) rhythmical midfrontal 6-7/sec activity of juveniles, b) rhythmical midfrontal sharp 4-6/sec activity of infancy and early childhood with arousal from sleep, and c) frontal intermittent rhythmical delta activity (FIRDA) in waking adults with frontopolar maximum, possibly related to thought processes under abnormal conditions. With extension of the frequency range, ultraslow (DC-like) as well as fast beta (gamma, 40-80/sec) and ultrafast activity (80-1000/sec) are found particularly over the frontal lobes. Ultraslow baseline shifts are arousal-related and mixed with overlying ultrafast waves. Attention control and the "working memory" involve chiefly the dorsolateral prefrontal cortex, investigated with P300 responses and likely to show ultrafast spectra. Perception-related 40-80/sec gamma activity has been thought to be associated with the entrance into consciousness. Initiation and design of motor activity spreads from prefrontal to the frontomotor cortex, associated with powerful event-related potentials: contingent negative variation (CNV) and "Bereitschafts potential" ("readiness potential," RP). Neuroscientific research of the highest frontal lobe functions has become a very active domain of neuroimaging. With the use of the extended frequency range, EEG and also evoked potential studies could add further information with acquisition in real time. Ultrafast frequency ranges presented in computerized frequency analysis and mapping might show impressive correlates of highest frontal lobe functions." [Abstract]

McAlonan K, Brown VJ.
The thalamic reticular nucleus: more than a sensory nucleus?
Neuroscientist 2002 Aug;8(4):302-5
"Sensory information is routed to the cortex via the thalamus, but despite this sensory bombardment, animals must attend selectively to stimuli that signal danger or opportunity. Sensory input must be filtered, allowing only behaviorally relevant information to capture limited attentional resources. Located between the thalamus and cortex is a thin lamina of neurons called the thalamic reticular nucleus (Rt). The thalamic reticular nucleus projects exclusively to thalamus, thus forming an essential component of the circuitry mediating sensory transmission. This article presents evidence supporting a role for Rt beyond the mere relay of sensory information. Rather than operating as a component of the sensory relay, the authors suggest that Rt represents an inhibitory interface or "attentional gate," which regulates the flow of information between the thalamus and cortex. Recent findings have also implicated Rt in higher cognitive functions, including learning, memory, and spatial cognition. Drawing from recent insights into the dynamic nature of the thalamic relay in awake, behaving animals, the authors present a speculative account of how Rt might regulate thalamocortical transmission and ultimately the contents of consciousness." [Abstract]

Llinas R, Ribary U, Contreras D, Pedroarena C.
The neuronal basis for consciousness.
Philos Trans R Soc Lond B Biol Sci 1998 Nov 29;353(1377):1841-9
"Attempting to understand how the brain, as a whole, might be organized seems, for the first time, to be a serious topic of inquiry. One aspect of its neuronal organization that seems particularly central to global function is the rich thalamocortical interconnectivity, and most particularly the reciprocal nature of the thalamocortical neuronal loop function. Moreover, the interaction between the specific and non-specific thalamic loops suggests that rather than a gate into the brain, the thalamus represents a hub from which any site in the cortex can communicate with any other such site or sites. The goal of this paper is to explore the basic assumption that large-scale, temporal coincidence of specific and non-specific thalamic activity generates the functional states that characterize human cognition." [Abstract] [Full Text]

Jones EG.
Thalamic circuitry and thalamocortical synchrony.
Philos Trans R Soc Lond B Biol Sci 2002 Dec;357(1428):1659-73
"The corticothalamic system has an important role in synchronizing the activities of thalamic and cortical neurons. Numerically, its synapses dominate the inputs to relay cells and to the gamma-amino butyric acid (GABA)ergic cells of the reticular nucleus (RTN). The capacity of relay neurons to operate in different voltage-dependent functional modes determines that the inputs from the cortex have the capacity directly to excite the relay cells, or indirectly to inhibit them via the RTN, serving to synchronize high- or low-frequency oscillatory activity respectively in the thalamocorticothalamic network. Differences in the alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA) subunit composition of receptors at synapses formed by branches of the same corticothalamic axon in the RTN and dorsal thalamus are an important element in the capacity of the cortex to synchronize low-frequency oscillations in the network. Interactions of focused corticothalamic axons arising from layer VI cortical cells and diffuse corticothalamic axons arising from layer V cortical cells, with the specifically projecting core relay cells and diffusely projecting matrix cells of the dorsal thalamus, form a substrate for synchronization of widespread populations of cortical and thalamic cells during high-frequency oscillations that underlie discrete conscious events." [Abstract]

Balkin TJ, Braun AR, Wesensten NJ, Jeffries K, Varga M, Baldwin P, Belenky G, Herscovitch P.
The process of awakening: a PET study of regional brain activity patterns mediating the re-establishment of alertness and consciousness.
Brain 2002 Oct;125(Pt 10):2308-19
"Awakening from sleep entails rapid re-establishment of consciousness followed by the relatively slow (20-30 min later) re-establishment of alertness--a temporal dissociation that facilitates specification of the physiological underpinnings of each of these facets of the awakening process. H(2)(15)O PET was used to assess changes in regional cerebral blood flow (rCBF) upon awakening from stage 2 sleep. Cerebral blood flow (CBF) was most rapidly re-established in centrencephalic regions (e.g. brainstem and thalamus), suggesting that the reactivation of these regions underlies the re-establishment of conscious awareness. Across the ensuing 15 min of wakefulness, further increases in CBF were evident primarily in anterior cortical regions, suggesting that the dissipation of sleep inertia effects (post-awakening performance and alertness deficits) is effected by reactivation of these regions. Concomitant shifts in correlation patterns of regional brain activity across the post-awakening period [in particular, a waning negative correlation between prefrontal cortex and mesencephalic reticular formation (RF) activity, and a waxing positive correlation between prefrontal cortex and ventromedial caudate nucleus (CAUD) activity] suggest that the post-awakening reversal of sleep inertia effects may be mediated by more than mere reactivation--it may also involve the functional reorganization of brain activity. Conversely, stable post-awakening correlations--such as those found between the anterior cingulate cortex (ACC) and most other brain regions--may denote the pattern of functional connectivity that underlies consciousness itself." [Abstract]

Steriade M.
The corticothalamic system in sleep.
Front Biosci 2003 May 1;8:D878-99
"The transition from wakefulness to NREM sleep is associated with typical signs of brain electrical activity, characterized by prolonged periods of hyperpolarization and increased membrane conductance in thalamocortical (TC) neurons, with the consequence that incoming messages are inhibited and the cerebral cortex is deprived of signals from the outside world. There are three major oscillations during NREM sleep. Spindles are generated within the thalamus, due to thalamic reticular (RE) neurons that impose rhythmic inhibitory sequences onto TC neurons, but the widespread synchronization of this rhythm is governed by corticothalamic projections. There are two types of delta activity: clock-like waves generated in TC neurons by the interplay between two hyperpolarization-activated inward currents; and cortical waves that survive extensive thalamectomy. The hallmark of NREM sleep activity is the slow oscillation, generated intracortically, which has the virtue of grouping the other types of sleep activities, thus leading to a coalescence of different rhythms that can only be observed in intact-brain animals and humans. Far from being epiphenomena, with no functional role, NREM sleep oscillations, particularly spindles and their experimental model augmenting responses, produce synaptic plasticity in target cortical neurons and resonant activity in corticothalamic loops, as in "memory" processes. Upon brain arousal, spindles are blocked by inhibition of RE neurons, the spindles' pacemakers; clock-like delta rh