A minimal synaptic architecture is proposed for how the brain might perform path integration by computing the next internal representation of self-location from the current representation and from the perceived velocity of motion. In the model, a place-cell assembly called a “chart ” contains a twodimensional attractor set called an “attractor map ” that can be used to represent coordinates in any arbitrary environment, once associative binding has occurred between chart locations and sensory inputs. In hippocampus, there are different spatial relations among place fields in different environments and behavioral contexts. Thus, the same units may participate in many charts, and it is shown that the number of uncorrelated charts that can be encoded in the same recurrent network is potentially quite large. According to this theory, the firing of a given place cell is primarily a cooperative effect of the activity of its

Information in neuronal networks may be represented by the spatiotemporal patterns of spikes. Here we examined the temporal coordination of pyramidal cell spikes in the rat hippocampus during slow-wave sleep. In addition, rats were trained to run in a defined position in space (running wheel) to activate a selected group of pyramidal cells. A template-matching method and a joint probability map method were used for sequence search. Repeating spike sequences in excess of chance occurrence were examined by comparing the number of repeating sequences in the original spike trains and in surrogate trains after Monte Carlo shuffling of the spikes. Four different shuffling procedures were used to control for the population dynamics of Although it is a widely accepted notion that information is distributed in cell assemblies rather than encoded by single cells, the nature of coding in cell assembly has remained a major challenge for neuroscience research. Several explanations have been proposed on theoretical grounds, including frequency coding

The firing characteristics of 1437 CA1 pyramidal neurons were studied in relation to both spatial location and the phase of the theta rhythm in healthy young and old rats performing a simple spatial task on a rectangular track. The old rats had previously been found to be deficient on the Morris spatial learning task. Age effects on the theta rhythm per se were minimal. Theta amplitude and frequency during rapid eye movement sleep were virtually identical. During behavior, theta frequency was slightly reduced with age. In both groups, cell firing occurred at progressively earlier phases of the theta rhythm as the rat traversed the place field of the cell (i.e., there was “phase precession, ” as reported by others). The net phase shift did not differ between age groups. The main finding of the study was a Because effective spatial learning requires an intact hippocampus (e.g., O’Keefe and Nadel, 1978; Morris et al., 1982; Sutherland et al., 1982; Barnes, 1988; Jarrard, 1993), and because this crucial cognitive function is significantly impaired during normal aging, the study of how neuronal information processing in the hippocampus is altered with age is of considerable interest. In rodents, age deficits are observed in a variety of spatial learning

Although it is well known that hippocampal neurons code spatial information, it is less clear how these spatial representations are influenced by memory demands, especially in hippocampus-dependent tasks. Recently, our laboratory has demonstrated that hippocampal spatial representations are influenced by mnemonic factors in a T-maze continuous alternation task. Another unique experimental approach that might reveal the ways in which task-related factors impact hippocampal spatial representations is to compare firing patterns between events that require distinct episodic memory processes. Therefore, we recorded from CA1 single neurons during a discrete trial delayed-nonmatch-to-place task that allowed within-trial comparison between an encoding (sample) phase and a retrieval (choice) phase. A large subset of neurons that fired on the central stem of the maze showed dramatic selectivity for either the sample or choice phase of the trial. However, surprisingly, there were fewer neurons that showed differential firing rates between left- and rightbound trajectories. Our results suggest that trial-phase-selective coding is common in tasks that require rapid alternation between encoding and retrieval processes. Key words: hippocampus; context; delayed-nonmatch-to-place; episodic memory; encoding; retrieval

ABSTRACT: Although most observers agree that the hippocampus has a critical role in learning and memory, there remains considerable debate about the precise functional contribution of the hippocampus to these processes. Two of the most influential accounts hold that the primary function of the hippocampus is to generate cognitive maps and to mediate episodic memory processes. The well-documented spatial firing patterns (place fields) of hippocampal neurons in rodents, along with the spatial learning impairments observed with hippocampal damage support the cognitive mapping hypothesis. The amnesia for personally experienced events seen in humans with hippocampal damage and the data of animal models, which show severe memory deficits associated with hippocampal lesions, support the episodic memory account. Although an extensive literature supports each of these hypotheses, a specific contribution of place cells to episodic memory has not been clearly demonstrated. Recent data from our laboratory, together with previous findings, indicate that hippocampal place fields and neuronal responses to task-relevant stimuli are highly sensitive to the context, even when the contexts are defined by abstract task demands rather than the spatial geometry of the environment. On the basis of these findings, it is proposed that place fields reflect a more general context processing function of the hippocampus. Hippocampal context representations could serve to differentiate contexts and prime the relevant memories and behaviors. Since episodic memories, by definition, include information about the time and place where the episode occurred, contextual information is a necessary prerequisite for any episodic memory. Thus, place fields contribute importantly to episodic memory as part of the needed context representations. Additionally, recent findings indicate that hippocampal neurons differentiate contexts at progressively finer levels of detail, suggesting a hierarchical coding scheme which, if combined with temporal information, could provide a means of differentiating memory episodes. VC 2006 Wiley-Liss, Inc. KEY WORDS: hippocampus; context; discrimination; learning; spatial

During slow wave sleep (SWS), traces of neuronal activity patterns from preceding behavior can be observed in rat hippocampus and neocortex. The spontaneous reactivation of these patterns is manifested as the reinstatement of the distribution of pairwise firing-rate correlations within a population of simultaneously recorded neurons. The effects of behavioral state [quiet wakefulness, SWS, and rapid eye movement (REM)], interactions between two successive spatial experiences, and global modulation during 200 Hz electroencephalographic (EEG) “ripples ” on pattern reinstatement were studied in CA1 pyramidal cell population recordings. Pairwise firing-rate correlations during often repeated experiences accounted for a significant proportion of the variance in these interactions in subsequent SWS or quiet wakefulness and, to a lesser degree, during SWS before the experience on a given day. The latter effect was absent for novel experiences, suggesting that a persistent memory trace develops with experience. Pattern The hippocampus is thought to play an important role in the acquisition and consolidation of certain forms of memory. Lesions of the hippocampus lead to a temporally graded retrograde amnesia, suggesting that the hippocampus plays a role in the initial encoding of a memory but that, with time, the memory becomes independent of the hippocampus (Scoville and Milner,

Previous studies in slices have shown that low-frequency stimulation at 5 Hz, i.e., theta pulse stimulation (TPS), completely reverses long-term potentiation (LTP) in area CA1 when delivered within 1–2 min after induction but produces progressively less depotentiation at longer delays, until it has no longer any impact at 30 min after induction. The present study examined whether LTP in the freely moving rat exhibits a similar timedependent susceptibility to reversal. Adult male Long–Evans rats with bilateral stimulating electrodes activating collateral/ commissural projections to area CA1 were used. A 1 min episode of TPS, ineffective when applied to naive pathways, was found to permanently erase LTP when delivered to the test pathway either 30 sec or 15 min after induction. Administered at a delay of 30 min, however, the same treatment no longer had Behavioral studies have shown that various manipulations can disrupt the stable encoding of memory if applied shortly after initial learning and thereby produce retrograde amnesia (Mc-Gaugh, 1966; McGaugh et al., 1993). The question naturally arises whether long-term potentiation (LTP) passes through a similar consolidation period during which it is susceptible to disruption. The first evidence suggestive of a vulnerable phase for LTP came from a study showing that transient hypoxia occurring within a few minutes after high-frequency stimulation prevented the stable formation of LTP in hippocampal area CA1 but was ineffective at longer delays (Arai et al., 1990b). An alternative approach to address the reversibility of LTP is to examine activity-dependent depotentiation, defined as a lasting decrement in synaptic efficacy induced by activation of the same set of pathways that were tetanized previously. Most initial observations on depotentiation were done in vitro and used stimulation protocols that differed in frequency, duration, and time of reversal attempt across laboratories, making it difficult for a coherent picture to emerge (Fujii et al., 1991; Wexler and Stanton, 1993;

Vestibular information influences spatial orientation and navigation in laboratory animals and humans. Neurons within the rat anterior thalamus encode the directional heading of the animal in absolute space. These neurons, referred to as head direction (HD) cells, fire selectively when the rat points its head in a specific direction in the horizontal plane with respect to the external laboratory reference frame. HD cells are thought to represent an essential component of a neural network that processes allocentric spatial information. The functional properties of HD cells may be dependent on vestibular input. Here, anterior thalamic HD cells were recorded before and after sodium arsanilate-induced vestibular system lesion. Vestibular lesions abolished the directional firing properties of HD cells. The time course of disruption in the directional firing properties paralleled the loss of vestibular function. Arsanilate-treated rats

ABSTRACT: The very different anatomical designs of the adjacent circuitries of the cortico-hippocampal pathway, along with their somewhat different synaptic plasticity mechanisms, suggest a nearly serial pathway of distinct memory circuits each contributing its own specialized processing operation to overall hippocampal function. Modeling and formal theoretical analysis of the prominent anatomical design features of particular circuits (piriform/entorhinal cortex; hippocampal field CA3; hippocampal field CA1) are found to identify potential emergent function not readily arrived at in the absence of these formal models, and yet which once derived can be seen potentially to confer unique capabilities to an integrated hippocampal mechanism for processing memories during behavior. r 1997 Wiley-Liss, Inc. KEY WORDS: LTP; Memory; corticohippocampal pathway; CA3; CA1; dentate gyrus; entorhinal cortex; perirhinal cortex; parahippocampal cortex

Advanced age in rats is associated with a decline in spatial memory capacities dependent on hippocampal processing. As yet, however, little is known about the nature of age-related alterations in the information encoded by the hippocampus. Young rats and aged rats identified as intact or impaired in spatial learning capacity were trained on a radial arm maze task, and then multiple parameters of the environmental cues were manipulated to characterize the changes in firing patterns of hippocampal neurons corresponding to the presence of particular cues or the spatial relationships among them. The scope of information encoded by the hippocampus was reduced in memory-impaired aged subjects, even though the Diminished memory capacity is a well known concomitant of