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...

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.

We present a computational neural network model of recognition memory based on the biological structures of the hippocampus and medial temporal lobe cortex (MTLC), which perform complementary learning functions. The hippocampal component of the model contributes to recognition by recalling specific studied details. MTLC can not support recall, but it is possible to extract a scalar familiarity signal from MTLC that tracks how well the test item matches studied items. We present simulations that establish key qualitative differences in the operating characteristics of the hippocampal recall and MTLC familiarity signals, and we identify several manipulations (e.g., target-lure similarity, interference) that differentially affect the two signals. We also use the model to address the stochastic relationship between recall and familiarity (i.e., are they independent), and the effects of partial vs. complete hippocampal

Structural priming in language production is a tendency to recreate a recently uttered syntactic structure in different words. This tendency can be seen independent of specific lexical items, thematic roles, or word sequences. Two alternative proposals about the mechanism behind structural priming include (a) short-term activation from a memory representation of a priming structure and (b) longer term adaptation within the cognitive mechanisms for creating sentences, as a form of procedural learning. Two experiments evaluated these hypotheses, focusing on the persistence of structural priming. Both experiments yielded priming that endured beyond adjacent sentences, persisting over 2 intervening sentences in Experiment 1 and over 10 in Experiment 2. Although memory may have short-term consequences for some components of this kind of priming, the persisting effects are more compatible with a learning account than a transient memory account. Speakers repeat themselves. Sometimes their repetitions are intentional, made for emphasis or other stylistic and social purposes (Giles & Powesland, 1975; Tannen, 1987), and sometimes they are accidental. They may involve almost

this article. Because there is no visible cue in the hidden-platform water maze task, it would not help the animal find the platform. 3. Route system. Routes stored in the hippocampus can be written out to the cortex, so that directions necessary to reach a goal are associated with local views. This is the system detailed in section 2.5 (see also section 4.3). This system requires training for each step the animal must take; it cannot learn to associate local views with directions to distant goals without hippocampal help (through route replay). The Role of the Hippocampus 97 If there were a way to show the animal the route to the goal, it might be possible to train the route system even without a hippocampus. Whishaw, Cassell, and Jarrard (1995) and Schallert, Day, Weisend, and Sutherland (1996) both showed ways to train the route system directly and found that animals could learn to solve the water maze even with hippocampal lesions. Whishaw et al. (1995) trained animals with fimbria/fornix lesions to find a visible platform and then removed the visible platform. These animals concentrated their search where the platform had been. Schallert et al. (1996) used animals with kainate-colchicine hippocampal lesions. The animals were first trained with a large platform that filled almost the entire maze. Once the animals could reach that platform reliably, it was shrunk trial by trial until it was the same size as a typical platform in a water maze task. Again, the animals could learn to solve the water maze without a hippocampus. 4.3 Where Is the Route System? Although the data are not yet conclusive, we suggest that the most likely candidate for anatomical instantiation of the route system is from posterior parietal to posterior cingulate cortex. There is a lot of evide...

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Psycholinguistic research has shown that the influence of abstract syntactic knowledge on performance is shaped by particular sentences that have been experienced. To explore this idea, the authors applied a connectionist model of sentence production to the development and use of abstract syntax. The model makes use of (a) error-based learning to acquire and adapt sequencing mechanisms and (b) meaning–form mappings to derive syntactic representations. The model is able to account for most of what is known about structural priming in adult speakers, as well as key findings in preferential looking and elicited production studies of language acquisition. The model suggests how abstract knowledge and concrete experience are balanced in the development and use of syntax.

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Early neuroimaging studies often failed to obtain evidence of medial temporal lobe (MTL) activation during episodic encoding or retrieval, but a growing number of studies using functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) have provided such evidence. We review data from fMRI studies that converge on the conclusion that posterior MTL is associated with episodic encoding; too few fMRI studies of retrieval have reported MTL activations to allow firm conclusions about their exact locations. We then turn to a recent meta-analysis of PET studies (Lepage et al., Hippocampus 1998;8:313-- 322) that appears to contradict the fMRI encoding data. Based on their analysis of the rostrocaudal distribution of activations reported during episodic encoding or retrieval, Lepage et al. (1998) concluded that anterior MTL is strongly associated with episodic encoding, whereas posterior MTL is strongly associated with episodic retrieval. After considering the evidence reviewed by Lepage et al. (1998) along with additional studies, we conclude that PET studies of encoding reveal both anterior and posterior MTL activations. These observations indicate that the contradiction between fMRI and PET studies of encoding was more apparent than real. However, PET studies have reported anterior MTL encoding activations more frequently than have fMRI studies. We consider possible sources of these differences. Hippocampus 1999;9:7--24.

The medial temporal lobe (MTL) has been studied extensively at all levels of analysis, yet its function remains unclear. Theory regarding the cognitive function of the MTL has centered along 3 themes. Different authors have emphasized the role of the MTL in episodic recall, spatial navigation, or relational memory. Starting with the temporal context model (M.W. Howard and M. J. Kahana, 2002), a distributed memory model that has been applied to benchmark data from episodic recall tasks, the authors propose that the entorhinal cortex supports a gradually changing representation of temporal context and the hippocampus proper enables retrieval of these contextual states. Simulation studies show this hypothesis explains the firing of place cells in the entorhinal cortex and the behavioral effects of hippocampal lesion in relational memory tasks. These results constitute a first step towards a unified computational theory of MTL function that integrates neurophysiological, neuropsychological and cognitive findings.