The brains of higher mammals are extraordinary integrative devices. Signals from large numbers of functionally specialized groups of neurons distributed over many brain regions are integrated to generate a coherent, multimodal scene. Signals from the environment are integrated with ongoing, patterned neural activity that provides them with a meaningful context. We review recent advances in neurophysiology and neuroimaging that are beginning to reveal the neural mechanisms of integration. In addition, we discuss concepts and measures derived from information theory that lend (...) a theoretical basis to the notion of complexity as integration of information and suggest new experimental tests of these concepts. (shrink)

Phillips & Singer's compelling presentation is weakest in its demonstration of commonalities between sensory plasticity and higher forms of learning and behavior. We propose that available data on schizophrenia can provide such evidence, because of the presence of impairments in a number of functions central to their model, and strong relationships between these dysfunctions and behavior.

Using robots that operate in the real world as opposed to computer simulations of animal behavior is a form of modeling that may provide some biological insights. However, since engineering principles and materials differ significantly from those used in biology, one should be extremely cautious in interpreting robot biomimicry as providing an explanation of biological mechanisms.

Webb's scheme for classifying behavioral models is applicable to a wide range of theories and simulations, nonrobotic as well as robotic. It is suggested that a meta-analysis of existing models, characterized according to the proposed scheme, could identify regions of the seven-dimensional modelling space that are particularly likely to lead to new insights in understanding behavior.

If the cortex is an associative memory, strongly connected cell assemblies will form when neurons in different cortical areas are frequently active at the same time. The cortical distributions of these assemblies must be a consequence of where in the cortex correlated neuronal activity occurred during learning. An assembly can be considered a functional unit exhibiting activity states such as full activation (“ignition”) after appropriate sensory stimulation (possibly related to perception) and continuous reverberation of excitation within the assembly (a putative (...) memory process). This has implications for cortical topographies and activity dynamics of cell assemblies forming during language acquisition, in particular for those representing words. Cortical topographies of assemblies should be related to aspects of the meaning of the words they represent, and physiological signs of cell assembly ignition should be followed by possible indicators of reverberation. The following postulates are discussed in detail: (1) assemblies representing phonological word forms are strongly lateralized and distributed over perisylvian cortices; (2) assemblies representing highly abstract words such as grammatical function words are also strongly lateralized and restricted to these perisylvian regions; (3) assemblies representing concrete content words include additional neurons in both hemispheres; (4) assemblies representing words referring to visual stimuli include neurons in visual cortices; and (5) assemblies representing words referring to actions include neurons in motor cortices. Two main sources of evidence are used to evaluate these proposals: (a) imaging studies focusing on localizing word processing in the brain, based on stimulus-triggered event-related potentials (ERPs), positron emission tomography (PET), and functional magnetic resonance imaging (fMRI), and (b) studies of the temporal dynamics of fast activity changes in the brain, as revealed by high-frequency responses recorded in the electroencephalogram (EEG) and magnetoencephalogram (MEG). These data provide evidence for processing differences between words and matched meaningless pseudowords, and between word classes, such as concrete content and abstract function words, and words evoking visual or motor associations. There is evidence for early word class-specific spreading of neuronal activity and for equally specific high-frequency responses occurring later. These results support a neurobiological model of language in the Hebbian tradition. Competing large-scale neuronal theories of language are discussed in light of the data summarized. Neurobiological perspectives on the problem of serial order of words in syntactic strings are considered in closing. Key Words: associative learning; cell assembly; cognition; cortex; ERP; EEG; fMRI; language; lexicon; MEG; PET; word category. (shrink)

An individual has a theory of mind if he imputes mental states to himself and others. A system of inferences of this kind is properly viewed as a theory because such states are not directly observable, and the system can be used to make predictions about the behavior of others. As to the mental states the chimpanzee may infer, consider those inferred by our own species, for example, purpose or intention, as well as knowledge, belief, thinking, doubt, guessing, pretending, liking, (...) and so forth. To determine whether or not the chimpanzee infers states of this kind, we showed an adult chimpanzee a series of videotaped scenes of a human actor struggling with a variety of problems. Some problems were simple, involving inaccessible food as in the original Kohler problems; others were more complex, involving an actor unable to extricate himself from a locked cage, shivering because of a malfunctioning heater, or unable to play a phonograph because it was unplugged. With each videotape the chimpanzee was given several photographs, one a solution to the problem, such as a stick for the inaccessible bananas, a key for the locked up actor, a lit wick for the malfunctioning heater. The chimpanzee's consistent choice of the correct photographs can be understood by assuming that the animal recognized the videotape as representing a problem, understood the actor's purpose, and chose alternatives compatible with that purpose. (shrink)

The additional data, perspectives, questions, and criticisms contributed by the commentaries strengthen our view that local cortical processors coordinate their activity with the context in which it occurs using contextual fields and synchronized population codes. We therefore predict that whereas the specialization of function has been the keynote of this century the coordination of function will be the keynote of the next.

It is worthwhile to search for forms of coding, processing, and learning common to various cortical regions and cognitive functions. Local cortical processors may coordinate their activity by maximizing the transmission of information coherently related to the context in which it occurs, thus forming synchronized population codes. This coordination involves contextual field (CF) connections that link processors within and between cortical regions. The effects of CF connections are distinguished from those mediating receptive field (RF) input; it is shown how CFs (...) can guide both learning and processing without becoming confused with the transmission of RF information. Simulations explore the capabilities of networks built from local processors with both RF and CF connections. Physiological evidence for synchronization, CFs, and plasticity of the RF and CF connections is described. Coordination via CFs is related to perceptual grouping, the effects of context on contrast sensitivity, amblyopia, implicit influences of color in achromotopsia, object and word perception, and the discovery of distal environmental variables and their interactions through self-organization. Cortical computation could thus involve the flexible evaluation of relations between input signals by locally specialized but adaptive processors whose activity is dynamically associated and coordinated within and between regions through specialized contextual connections. Key Words: cell assemblies; cerebral cortex; context; coordination; dynamic binding; epistemology; functional specialization; learning; neural coding; neural computation; neuropsychology; object recognition; perception; reading; self-organization; synaptic plasticity; synchronization. (shrink)

A general conceptual framework for large-scale neocortical dynamics based on data from many laboratories is applied to a variety of experimental designs, spatial scales, and brain states. Partly distinct, but interacting local processes (e.g., neural networks) arise from functional segregation. Global processes arise from functional integration and can facilitate (top down) synchronous activity in remote cell groups that function simultaneously at several different spatial scales. Simultaneous local processes may help drive (bottom up) macroscopic global dynamics observed with electroencephalography (EEG) or (...) magnetoencephalography (MEG). A local/global dynamic theory that is consistent with EEG data and the proposed conceptual framework is outlined. This theory is neutral about properties of neural networks embedded in macroscopic fields, but its global component makes several qualitative and semiquantitative predictions about EEG measures of traveling and standing wave phenomena. A more general “metatheory” suggests what large-scale quantitative theories of neocortical dynamics may be like when more accurate treatment of local and nonlinear effects is achieved. The theory describes the dynamics of excitatory and inhibitory synaptic action fields. EEG and MEG provide large-scale estimates of modulation of these synaptic fields around background levels. Brain states are determined by neuromodulatory control parameters. Purely local states are dominated by local feedback gains and rise and decay times of postsynaptic potentials. Dominant local frequencies vary with brain region. Other states are purely global, with moderate to high coherence over large distances. Multiple global mode frequencies arise from a combination of delays in corticocortical axons and neocortical boundary conditions. Global frequencies are identical in all cortical regions, but most states involve dynamic interactions between local networks and the global system. EEG frequencies may involve a “matching” of local resonant frequencies with one or more of the many, closely spaced global frequencies. Key Words: binding problem; cell assemblies; coherence; EEG; limit cycles; neocortical dynamics; pacemakers; phase locking; spatial scale; standing waves; synchronization. Footnotes1 The relationship between the synaptic action fields proposed in the target article and cell assemblies is clarified with Figure R1 (p. 416) of the Response. (This figure was not available to Commentators. (shrink)

While robotics has benefited from inspiration gained from biology, the opposite is not the case: there are few if any cases in which robotic models have lead to genuine insight into biology. We analyze the reasons why biorobotics has been essentially a one-way street. We argue that the development of better tools is essential for progress in this field.

The prefrontal cortex has long been suspected to play an important role in cognitive control, in the ability to orchestrate thought and action in accordance with internal goals. Its neural basis, however, has remained a mystery. Here, we propose that cognitive control stems from the active maintenance of patterns of activity in the prefrontal cortex that represent goals and the means to achieve them. They provide bias signals to other brain structures whose net effect is to guide the flow of (...) activity along neural pathways that establish the proper mappings between inputs, internal states, and outputs needed to perform a given task. We review neurophysiological, neurobiological, neuroimaging, and computational studies that support this theory and discuss its implications as well as further issues to be addressed. (shrink)

This paper analyzes an explicit instantiation of the program of neurophenomenology in a neuroscientific protocol. Neurophenomenology takes seriously the importance of linking the scientific study of consciousness to the careful examination of experience with a specific first-person methodology. My first claim is that such strategy is a fruitful heuristic because it produces new data and illuminates their relation to subjective experience. My second claim is that the approach could open the door to a natural account of the structure of human (...) experience as it is mobilized in itself in such methodology. In this view, generative passages define the type of circulation which explicitly roots the active and disciplined insight the subject has about his/her experience in a biological emergent process. (shrink)

This paper analyzes an explicit instantiation of the program of “neurophenomenology” in a neuroscientific protocol. Neurophenomenology takes seriously the importance of linking the scientific study of consciousness to the careful examination of experience with a specific first-person methodology. My first claim is that such strategy is a fruitful heuristic because it produces new data and illuminates their relation to subjective experience. My second claim is that the approach could open the door to a natural account of the structure of human (...) experience as it is mobilized in itself in such methodology. In this view, generative passages define the type of circulation which explicitly roots the active and disciplined insight the subject has about his/her experience in a biological emergent process. (shrink)

This article summarizes a variety of current as well as previous research in support of a new theory of consciousness. Evidence has been steadily accumulating that information about a stimulus complex is distributed to many neuronal populations dispersed throughout the brain and is represented by the departure from randomness of the temporal pattern of neural discharges within these large ensembles. Zero phase lag synchronization occurs between discharges of neurons in different brain regions and is enhanced by presentation of stimuli. This (...) evidence further suggests that spatiotemporal patterns of coherence, which have been identified by spatial principal component analysis, may encode a multidimensional representation of a present or past event. How such distributed information is integrated into a holistic percept constitutes the binding problem. How a percept defined by a spatial distribution of nonrandomness can be subjectively experienced constitutes the problem of consciousness. Explanations based on a discrete connectionistic network cannot be reconciled with the relevant facts. Evidence is presented herein of invariant features of brain electrical activity found to change reversibly with loss and return of consciousness in a study of 176 patients anesthetized during surgical procedures. A review of relevant research areas, as well as the anesthesia data, leads to a postulation that consciousness is a property of quantumlike processes, within a brain field resonating within a core of structures, which may be the neural substrate of consciousness. This core includes regions of the prefrontal cortex, the frontal cortex, the pre- and paracentral cortex, thalamus, limbic system, and basal ganglia. (shrink)

The core issue of our target article concerns how relational complexity should be assessed. We propose that assessments must be based on actual cognitive processes used in performing each step of a task. Complexity comparisons are important for the orderly interpretation of research findings. The links between relational complexity theory and several other formulations, as well as its implications for neural functioning, connectionist models, the roles of knowledge, and individual and developmental differences, are considered.

Current EEG research emphasizes gamma band coherence as a signature of functional integration, that is, the solution to the binding problem. We note that spatial patterns of coherent neural activity are also observed at other EEG frequencies. If these oscillations reflect Nunez's resonant modes, they offer a solution to the binding problem that emerges naturally from the architecture of cortical connections.

On the basis of neuroiinaging studies, Posner & Raichle summarily report that the prefrontal cortex is involved in executive functioning and attention. In contrast to that superficial view, we briefly describe a testable model of the kinds of representations that are stored in prefrontal cortex, which, when activated, are expressed via plans, actions, thematic knowledge, and schemas.

There is no best scientific model of anything; there are only models more or less good for different purposes. Thus, there is no general answer to the question of whether one should model biological behavior using computer simulations or robots. It all depends on what one wants to learn. This is not a question about models, but about scientific goals.

This paper explores the difference between Connectionist proposals for cognitive a r c h i t e c t u r e a n d t h e s o r t s o f m o d e l s t hat have traditionally been assum e d i n c o g n i t i v e s c i e n c e . W e c l a i m t h a t t h (...) e m a j o r d i s t i n c t i o n i s t h a t , w h i l e b o t h Connectionist and Classical architectures postulate representational mental states, the latter but not the former are committed to a symbol-level of representation, or to a ‘language of thought’: i.e., to representational states that have combinatorial syntactic and semantic structure. Several arguments for combinatorial structure in mental representations are then reviewed. These include arguments based on the ‘systematicity’ of mental representation: i.e., on the fact that cognitive capacities always exhibit certain symmetries, so that the ability to entertain a given thought implies the ability to entertain thoughts with semantically related contents. We claim that such arguments make a powerful case that mind/brain architecture is not Connectionist at the cognitive level. We then consider the possibility that Connectionism may provide an account of the neural (or ‘abstract neurological’) structures in which Classical cognitive architecture is implemented. We survey a n u m b e r o f t h e s t a n d a r d a r g u m e n t s t h a t h a v e b e e n o f f e r e d i n f a v o r o f Connectionism, and conclude that they are coherent only on this interpretation. (shrink)

William James described the stream of thought as having two components: (1) a nucleus of highly conscious, often perceptual material; and (2) a fringe of dimly felt contextual information that controls the entry of information into the nucleus and guides the progression of internally directed thought. Here I examine the neural and cognitive correlates of this phenomenology. A survey of the cognitive neuroscience literature suggests that the nucleus corresponds to a dynamic global buffer formed by interactions between different regions of (...) the brain, while the fringe corresponds to a set of mechanisms in the frontal and medial temporal lobes that control the contents of this global buffer. A consequence of this account is that there might be conscious imagistic representations that are not part of the nucleus. I argue that phenomenology can be linked to psychology and neuroscience and a meaningful way that illuminates both. (shrink)

Theories of binding have recently come into the focus of the consciousness debate. In this review, we discuss the potential relevance of temporal binding mechanisms for sensory awareness. Specifically, we suggest that neural synchrony with a precision in the millisecond range may be crucial for conscious processing, and may be involved in arousal, perceptual integration, attentional selection and working memory. Recent evidence from both animal and human studies demonstrates that specific changes in neuronal synchrony occur during all of these processes (...) and that they are distinguished by the emergence of fast oscillations with frequencies in the gamma-range. (shrink)

This paper starts with one of Chalmers’ basic points: first-hand experience is an irreducible field of phenomena. I claim there is no ‘theoretical fix’ or ‘extra ingredient’ in nature that can possibly bridge this gap. Instead, the field of conscious phenomena requires a rigorous method and an explicit pragmatics for its exploration and analysis. My proposed approach, inspired by the style of inquiry of phenomenology, I have called neurophenomenology. It seeks articulations by mutual constraints between phenomena present in experience and (...) the correlative field of phenomena established by the cognitive sciences. It needs to expand into a widening research community in which the method is cultivated further. (shrink)

[opening paragraph] -- Clark: The `astonishing hypothesis' which you put forward in your book, and which you obviously feel is very controversial, is that `You, your joys and sorrows, your memories and ambitions, your sense of personal identity and free will are, in fact, no more than the behaviour of a vast assembly of nerve cells. As Lewis Carroll's Alice might have phrased it: `You're nothing but a pack of neurons'.' But it seems to me that this is not so (...) astonishing a statement for a scientist to make. Isn't this what reductionist science has always believed? (shrink)