: In this work we discuss Hebb's old ideas about cell assemblies in the light of recent results concerning temporal structure and correlations in neural signals. We want to give a conceptual, necessarily only rough picture, how ideas about `binding by synchronisation', `synfire chains', `local and global assemblies', `short and long term memory' and `behaviour' might be integrated into a coherent model of brain functioning based on neuronal assemblies. Keywords: cell assemblies, synchronization, gamma-oscillations, synfire chains, memory, behaviour 1 ASSEMBLIES AND ASSOCIATIVE MEMORIES 1.1 Cell Assemblies Cell assemblies have been introduced by Donald Hebb with the intention of providing a functional and at the same time structural model for cortical processes and neuronal representations of external events (Hebb, 1949). According to Hebb's ideas, stimuli, objects, things, but also more abstract entities like concepts, contextual relations, ideas, and so on are thought of being repre...

Illusory conjunctions are the incorrect perceptual combination of briefly presented colors and shapes. In the neon colors illusion, achromatic figures take on the color of an overlaid grid of colored lines. Both illusions are explained by a theory that assumes (a) poor location information or poor spatial resolution for some aspects of visual information and (b) that the spatial location of features is constrained by perceptual organization. Computer simulations demonstrate that the mechanisms suggested by the theory are useful in veridical perception and they are sufficient to produce illusory conjunctions. The theory suggests mechanisms that economically encode visual information in a way that filters noise and fills in missing data. Issues related to neural implementation are discussed. Four experiments illustrate the theory. Illusory conjunctions are shown to be affected by objective stimulus organization, by subjective organization, and by the linguistic structure of ambiguous Hebrew words. Neon colors are constrained by linguistic structure in the same way as illusory conjunctions. Marr (1982) suggested that the study of vision can be conceptualized in three related levels of analysis. At the computational level, a specific functional problem in

In order to gather evidence on functional subdivisions of the temporal lobe neocortex of the primate, the activity of more than 2600 single neurons was recorded in 10 myelo- and cytoarchitecturally defined subdivisions of the cortex in the superior temporal sulcus (STS) and inferior temporal gyrus of the anterior part of the temporal lobe of 5 hemispheres of 3 macaque monkeys. First, convergence of different mo-dalities into each area was investigated. Areas TS and TAa, in the upper part of this region, were found to receive visual as well as auditory inputs. Areas TPO, PGa, and IPa, in the depths of the STS, received visual, auditory, and somato-sensory inputs. Areas TEa, TEm, TE3, TE2, and TE 1, which extend from the ventral bank of the STS through the inferior temporal gyrus, were primarily unimodal visual areas. Sec-ond, of the cells with visual responses, it was found that

The existence of recurrent collateral connections between pyramidal cells within a cortical area and, in addition, reciprocal connections between connected cortical areas, is well established. In this work we analyse the properties of a tri-modular architecture of this type in which two input modules have convergent connections to a third module (which in the brain might be the next module in cortical processing or a bi-modal area receiving connections from two different processing pathways). Memory retrieval is analysed in this system which has Hebb-like synaptic modifiability in the connections and attractor states. Local activity features are stored in the intra-modular connections while the associations between corresponding features in different modules present during training are stored in the inter-modular connections. The response of the network when tested with corresponding and contradictory stimuli to the two input pathways is studied in detail. The model is solved quantitatively using techniques of statistical physics. In one type of test, a sequence of stimuli is applied, with a delay between them. It is found that if the coupling between the modules is low a regime exists in which they retain the capability to retrieve any of their stored features independently of the features being retrieved by the other modules. Although independent in this sense, the modules still influence each other in this regime through persistent modulatory currents which are strong enough to initiate recall in the whole network when only a single module is stimulated, and to raise the mean firing rates of the neurons in the attractors if the features in the different modules are corresponding. Some of these mechanisms might be useful for the description of many phenomena observe...

Working memory has been linked to elevated single neuron discharge in monkeys and to oscillatory changes in the human EEG, but the relation between these effects has remained largely unexplored. We addressed this question by measuring local field potentials and single unit activity simultaneously from multiple electrodes placed in extrastriate visual cortex while monkeys were performing a working memory task. We describe a significant enhancement in theta band energy during the delay period. Theta oscillations had a systematic effect on single neuron activity, with neurons emitting more action potentials near their preferred angle of each theta cycle. Sample-selective de-lay activity was enhanced if only action potentials emitted near the preferred theta angle were consid-ered. Our results suggest that extrastriate visual cor-tex is involved in short-term maintenance of informa-tion and that theta oscillations provide a mechanism for structuring the recurrent interaction between neurons in different brain regions that underlie working memory.

A population of neurons in anterior inferotemporal cortex has sustained activity following the presentation of specific visual stimuli when monkeys perform a delayed match-to-sample (DMS) task. Typically, only stimuli that are repeatedly shown elicit robust delay activity. When the sample stimuli were shown in a fixed temporal order, the few images that evoked delay activity in the same neuron were often neighboring stimuli in the sequence. Therefore, this delay activity was suggested as the neural correlate of associative long-term memory of visual images. We report here that stimulus selective sustained activity is evident also following the presentation of the test stimulus in the same DMS task. We demonstrate, using a neural network model, that the persistence of stimulus selective activity across the inter-trial-interval can lead to similar menmonic representations (distribution of delay activity across the neural population) for neighboring visual stimuli. Thus, the neural machin...

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Previous research has identified the prefrontal cortex (PFC) as a brain region that is critical for cognitive control. Currently, theorists remain divided about whether to view the PFC as primarily a coordinative, mnemonic, or inhibitory structure. A theory is presented that attempts to resolve some of the apparent conflicts between the predominant views on PFC control functions. In this theory, PFC is proposed to actively maintain representations of context information. These maintained representations provide a mechanism of control by serving as a top-down bias on the local competitive interactions that occur during processing. As such, it is suggested that PFC performs both mnemonic and inhibitory functions in the service of control, and that each is preferentially observable under different task situations. A series of behavioral, computational, and neuroimaging studies are presented that demonstrates how this theory can account for a wide range of data associated with performance ...

Recordings from cells in the associative cortex of monkeys performing visual working memory tasks link persistent neuronal activity, long-term memory and associative memory. In particular, delayed pair-associate tasks have revealed neuronal correlates of long-term memory of associations between stimuli. Here, a recurrent cortical network model with Hebbian plastic synapses is subjected to the pair-associate protocol. In a ®rst stage, learning leads to the appearance of delay activity, representing individual images ('retrospective' activity). As learning proceeds, the same learning mechanism uses retrospective delay activity together with choice stimulus activity to potentiate synapses connecting neural populations representing associated images. As a result, the neural population corresponding to the pair-associate of the image presented is activated prior to its visual stimulation ('prospective ' activity). The probability of appearance of prospective activity is governed by the strength of the inter-population connections, which in turn depends on the frequency of pairings during training. The time course of the transitions from retrospective to prospective activity during the delay period is found to depend on the fraction of slow, N-methyl-D-aspartate-like receptors at excitatory synapses. For fast recurrent excitation, transitions are abrupt; slow recurrent excitation renders transitions gradual. Both scenarios lead to a gradual rise of delay activity when averaged over many trials, because of the stochastic nature of the transitions. The model reproduces most of the neuro-physiological data obtained during such tasks, makes experimentally testable predictions and demonstrates how persistent activity (working

Iconic memory, the short-lasting visual memory of a briefly flashed stimulus, is an important component of most models of visual perception. Here we investigate what physiological mechanisms underlie this capacity by showing rapid serial visual presentation (RSVP) sequences with and without interstimulus gaps to human observers and macaque monkeys. For gaps of up to 93 ms between consecutive images, human observers and neurones in the temporal cortex of macaque monkeys were found to continue processing a stimulus as if it was still present on the screen. The continued firing of neurones in temporal cortex may therefore underlie iconic memory. Based on these findings, a neurophysiological vision of iconic memory is presented. Iconic memory and information persistence In the world around us, many important events can be fleetingly brief. A gazelle might get a brief