The influence of prior experience on some forms of behavior and cognition is drastically affected by damage to the hippocampal system. However, if the hippocampal system is left intact both during the experience and for a period of time thereafter, subsequent damage can have much less or even no effect. Such findings suggest that memory traces change over time in a way that makes them less dependent on the hippocampal system. This process of change has often been called consolidation. Consolidation is a very gradual process; in humans, it appears to span up to 15 years. This article asks what consolidation is and why it occurs. We take as our point of departure the view that the initial memory trace that results from a relevant experience consists of changes to the strengths of the connections among neurons in the hippocampal system. Bidirectional connections between the neocortex and the hippocampus allow these initial traces to mediate the reinstatement of representations of events o...

The authors draw together the results of a series of detailed computational studies and show how they are contributing to the development of a theory of hippocampal function. A new part of the theory introduced here is a quantitative analysis of how backprojections from the hippocampus to the neocortex could lead to the recall of recent memories. The theory is then compared with other theories of hippocampal function. First, what is computed by the hippocampus is considered. The hypothesis the authors advocate, on the basis of the effects of damage to the hippocampus and neuronal activity recorded in it, is that it is involved in the formation of new memories by acting as an intermediate-term buffer store for information about episodes, particularly for spatial, but probably also for some nonspatial, information. The authors analyze how the hippocampus could perform this function, by producing a computational theory of how it operates, based on neuroanatomical and neurophysiological information about the different neuronal systems con-tained within the hippocampus. Key hypotheses are that the CA3 pyramidal cells operate as a single autoassociation network to store new episodic information as it arrives via a number of specialized preprocessing stages from many association areas of the cerebral cortex, and that the dentate

The hippocampus and related structures are thought to be capable of 1) representing cortical activity in a way that minimizes overlap of the representations assigned t ~ different cortical patterns (pattern separation); and 2) modifying synaptic connections so that these representations can later be reinstated from partial or noisy versions of the cortical activity pattern that was present at the time of storage (pattern completion). We point out that there is a trade-off between pattern separation and completion and propose that the unique anatomical and physiological properties of the hippocampus might serve to minimize this trade-off. We use analytical methods to determine quantitative estimates of both separation and completion for specified parameterized models of the hippocampus. These estimates are then used to evaluate the role of various properties and of the hippocampus, such as the activity levels seen in different hippocampal regions, synaptic potentiation and depression, the multi-layer connectivity of the system, and the relatively focused and strong mossy fiber projections. This analysis is focused on the feedforward pathways from the entorhinal cortex (EC) to the dentate gyrus (DG) and region CA3. Among our results are the following: 1) Hebbian synaptic modification (LTP) facilitates completion but reduces separation, unless the

We present a theoretical framework for understanding the roles of the hippocampus and neocortex in learning and memory. This framework incorporates a theme found in many theories of hippocampal function, that the hippocampus is responsible for developing conjunctive representations binding together stimulus elements into a unitary rep- resentation that can later be recalled from partial input cues. This idea appears problematic, however, because it is contradicted by the fact that hippocampally lesioned rats can learn nonlinear discrimination problems that require conjunctive representations. Our framework accommodates this finding by establishing a principled division of labor between the cortex and hippocampus, where the cortex is responsible for slow learning that integrates over multiple experiences to extract generalities, while the hippocampus performs rapid learning of the arbitrary contents of individual experiences. This framework shows that nonlinear discrimination problems are not good tests of hippocampal function, and suggests that tasks involving rapid, incidental conjunctive learning are better. We implement this framework in a computational neural network model, and show that it can account for a wide range of data in animal learning, thus validating our theoretical ideas, and providing a number of insights and predictions about these learning phenomena.

Hippocampal ‘place ’ cells and the head-direction cells of the dorsal presubiculum and related neocortical and thalamic areas appear to be part of a preconfigured network that generates an abstract internal representation of two-dimensional space whose metric is self-motion. It appears that viewpoint-specific visual information (e.g. landmarks) becomes secondarily bound to this structure by associative learning. These associations between landmarks and the preconfigured path integrator serve to set the origin for path integration and to correct for cumulative

The CA3 network in the hippocampus may operate as an autoassociator, in which declarative memories, known to be dependent on hippocampal processing, could be stored, and subsequently retrieved, using modifiable synaptic efficacies in the CA3 recurrent collateral system. On the basis of this hypothesis, the authors explore the computational relevance of the extrinsic afferents. to the CA3 network. A quantitative statistical analysis of the information that may be relayed by such afferent connections reveals the need for two distinct systems of input synapses. The synapses of the first system need to be strong (but not associatively modifiable) in order to force, during learning, the CA3 cells into a pattern of activity relatively independent of any inputs being received from the recurrent collaterals, and which thus reflects sizable amounts of new information. It is proposed that the mossy fiber system performs this function. A second system, with a large number of associatively modifiable synapses on each receiving cell, is needed in order to relay a signal specific enough to initiate the retrieval process. This may be identified, we propose, with the perforant path input to CA3. Key words: hippocampus, autoassociative memory, attractor neural networks, associative synapses, information storage

In the past decade, a large number of robots has been built that explicitly implement biological navigation behaviours. We review these biomimetic approaches using a framework that allows for a common description of biological and technical navigation behaviour. The review shows that biomimetic systems make significant contributions to two fields of research: First, they provide a real world test of models of biological navigation behaviour; second, they make new navigation mechanisms available for technical applications, most notably in the field of indoor robot navigation. While simpler insect navigation behaviours have been implemented quite successfully, the more complicated way-finding capabilities of vertebrates still pose a challenge to current systems. ©2000 Elsevier Science B.V. All rights reserved.

In this paper we describe a model that applies Marr's theory of hippocampal function to the problem of map based navigation. Like many others we attribute a spatial memory function to the hippocampus, but we suggest that the additional functional components required for map based navigation are located elsewhere in the brain. One of the key functional components in this model is an egocentric map of space, located in the neocortex, that is continuously updated using ideothetic (self motion) information. The hippocampus stores snapshots of this egocentric map. The modelled activity pattern of head direction cells is used to set the best egocentric map rotation to match the snapshots stored in the hippocampus, resulting in place cells with a non-directional firing pattern. We describe an evaluation of this model using a mobile robot, and demonstrate that with this model the robot can recognise an environment and find a hidden goal. This model is discussed in the context of prior experime...

Human cognition is unique in the way in which it relies on combinatorial (or compositional) structures. Language provides ample evidence for the existence of combinatorial structures, but they can also be found in visual cognition. To understand the neural basis of human cognition, it is therefore essential to understand how combinatorial structures can be instantiated in neural terms. In his recent book on the foundations of language, Jackendoff formulated four fundamental problems for a neural instantiation of combinatorial structures: the massiveness of the binding problem, the problem of 2, the problem of variables and the transformation of combinatorial structures from working memory to long-term memory. This paper aims to show that these problems can be solved by means of neural ‘blackboard ’ architectures. For this purpose, a neural blackboard architecture for sentence structure is presented. In this architecture, neural structures that encode for words are temporarily bound in a manner that preserves the structure of the sentence. It is shown that the architecture solves the four problems presented by Jackendoff. The ability of the architecture to instantiate sentence structures is illustrated with examples of sentence complexity observed in human language performance. Similarities exist between the architecture for sentence structure and blackboard architectures for combinatorial structures in visual cognition, derived from the structure of the visual cortex. These architectures are briefly discussed, together with an example of a combinatorial structure in which the blackboard architectures for language and vision are combined. In this way, the architecture for language is grounded in perception. 2 Content

O'Keefe (1991) has proposed that spatial information in rats might be represented as phasors: phase and amplitude of a sine wave encoding angle and distance to a landmark. We describe computer simulations showing that operations on phasors can be efficiently realized by arrays of spiking neurons that re-code the temporal dimension of the sine wave spatially. Some cells in motor and parietal cortex exhibit response properties compatible with this proposal. 1 Address all correspondence to the first author. Electronic mail address: dst@cs.cmu.edu. This work was supported by a contract from Fujitsu Corporation. Hank Wan and David Redish were supported by NSF Graduate Fellowships. The views and conclusions contained in this document are those of the authors and should not be interpreted as representing the official policies, either expressed or implied, of the Fujitsu Corporation, the National Science Foundation, or the U.S. government. Keywords: Neural Modelling, Spatial Reasoning, Pari...