The role of the hippocampus in retaining relational information across short delays: the importance of memory load.

Patients with hippocampal damage are sometimes impaired at remembering information across delays as short as a few seconds. How are these impairments to be understood? One possibility is that retention of some kinds of information is critically dependent on the hippocampus, regardless of the retention interval and regardless of whether the task depends on working memory or long-term memory. Alternatively, retention may be dependent on the hippocampus only when the task involves a memory load large enough to exceed working memory capacity. To explore these possibilities, we assessed the performance of patients with hippocampal lesions on two tasks requiring retention of the same object-in-scene information across a brief delay. The tasks placed different demands on memory. In one task, which used a continuous recognition format, participants needed to try to hold up to nine scenes in mind, even when no scene intervened between the study scene and the corresponding test scene. Patients were impaired in this condition. In a second task, using a conventional study-test format, participants needed to hold in mind only one scene at a time for either 3 or 14 sec. With this procedure, patients performed as well as controls after a 3-sec delay but were impaired after a 14-sec delay. We suggest that retention of object-in-scene information is dependent on the hippocampus only when working memory is insufficient to support performance (because memory load is high or the retention interval is long). In these circumstances performance depends, at least in part, on long-term memory.

Intact working memory for relational information after medial temporal lobe damage.

Working memory has traditionally been viewed as independent of the hippocampus and related medial temporal lobe structures. Yet memory-impaired patients with medial temporal lobe damage are sometimes impaired at remembering relational information (e.g., an object and its location) across delays as short as a few seconds. This observation has raised the possibility that medial temporal lobe structures are sometimes critical for maintaining relational information, regardless of whether the task depends on working or long-term memory. An alternative possibility is that these structures are critical for maintaining relational information only when the task exceeds working memory capacity and depends instead on long-term memory. To test these ideas, we drew on a method used previously in a classic study of digit span in patient HM that distinguished immediate memory from long-term memory. In two experiments, we assessed the ability of four patients with medial temporal lobe lesions to maintain varying numbers of object-location associations across a 1 s retention interval. In both experiments, the patients exhibited a similar pattern of performance. They performed similarly to controls when only a small number of object-location associations needed to be maintained, and they exhibited an abrupt discontinuity in performance with larger set sizes. This pattern of results supports the idea that maintenance of relational information in working memory is intact after damage to the hippocampus and related medial temporal lobe structures and that damage to these structures impairs performance only when the task depends on long-term memory.

Role of the hippocampus in remembering the past and imagining the future.

It has been proposed that a core network of brain regions, including the hippocampus, supports both past remembering and future imagining. We investigated the importance of the hippocampus for these functions. Five patients with bilateral hippocampal damage and one patient with large medial temporal lobe lesions were tested for their ability to recount autobiographical episodes from the remote past, the recent past, and to imagine plausible episodes in the near future. The patients with hippocampal damage had intact remote autobiographical memory, modestly impaired recent memory, and an intact ability to imagine the future. The patient with large medial temporal lobe lesions had intact remote memory, markedly impaired recent memory, and also had an intact ability to imagine the future. The findings suggest that the capacity for imagining the future, like the capacity for remembering the remote past, is independent of the hippocampus.

An investigation of auditory dimensional interaction in a bivariate bilateral conditioning paradigm in the rabbit.

The current study adapted the Garner paradigm for diagnosing separable versus integral perceptual dimensions to the eye-blink classical conditioning paradigm using rabbits. Specifically, this study examined the ability of rabbits to categorize stimuli based on one auditory dimension while ignoring a second, irrelevant dimension by displaying an appropriate eye-blink for bilaterally conditioned discriminative responses. Tones used in training varied along two dimensions, starting frequency and magnitude of frequency sweep upwards from the start. Rabbits first learned to categorize along a single dimension (blinking one eye for one category response and the other eye for the other response) and then continued to categorize tones in a second phase in which the irrelevant dimension was varied. The variation of the irrelevant dimension did not disrupt performance, indicating that rabbits perceive these dimensions as separable.

Hippocampal response patterns during discriminative eyeblink/jaw movement conditioning in the rabbit.

Rabbits were given concurrent training in eyeblink (EB) and jaw movement (JM) conditioning in which 1 tone predicted an airpuff and another tone predicted water. After 10 days of discrimination training, the animals were given 10 days of reversal training. In the discrimination phase, acquisition of the 2 conditioned responses was not significantly different; however JM discrimination errors were much more frequent than were EB errors. In the reversal phase, correct performance on EB trials increased gradually, as was expected, whereas there was immediate behavioral reversal on JM trials. Differences in size and topography of dorsal CA1 multiple-unit responses reflected the ability of the hippocampus to discriminate between stimuli in trained animals, corresponding to the performance of the behavioral discrimination. During JM trials, the rhythmicity of the neural response was further modulated by the type of the prior trial, suggesting the coding of sequential events by the hippocampus. Thus, hippocampal conditioned activity can rapidly change its magnitude and pattern depending on the specific trial type during a concurrent EB/JM discrimination task and its reversal. (PsycINFO Database Record (c) 2008 APA, all rights reserved).

Nonpharmacological amelioration of age-related learning deficits: the impact of hippocampal theta-triggered training.

Age-related learning deficits are often attributed to deterioration of hippocampal function. Conversely, a well studied index of hippocampal activity, the rhythm, is known to enhance hippocampal plasticity and accelerate learning rate in young subjects, suggesting that manipulations of activity might be used as a means to counteract impairments related to the aging process. Here, young and older rabbits were given eyeblink conditioning trials either when exhibiting hippocampal (+) or regardless of hippocampal activity (yoked control). Although, as expected, older-yoked control animals showed a learning deficit, the older + group learned as fast as young controls, demonstrating that aging deficits, at least in eyeblink classical conditioning, can be overcome by giving trials during episodes of hippocampal activity. The use of several learning criteria showed that the benefits of hippocampal occur in multiple phases of learning that may depend on different cognitive or motor processes. Whereas there was a benefit of -triggered training in both age groups during the early phase of acquisition, the enhancement persisted in older animals, peaking during later performance. These findings have implications for theories of age-related memory deficits and may contribute to the development of beneficial treatments.