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The present study describes the spatial firing properties of neurons in the lateral septum (LS). LS neuronal activity was recorded in rats as they performed a spatial navigation task in an open field. In this task, the rat acquired an intracranial self-stimulation reward when it entered a certain place, a location that varied randomly from trial to trial. Of 193 neurons recorded in the LS, 81 showed place-related activity. The majority of the tested neurons changed place-related activity when spatial relations between environmental cues were altered by rotating intrafield (proximal) cues. The comparison of place activities between LS place-related neurons recorded in the present study and hippocampal place cells recorded in our previous study, using identical behavioral and recording procedures, revealed that spatial parameters (spatial information content, coherence, and cluster size) were smaller in the LS than in the hippocampus. Of the 193 LS neurons, 86 were influenced by intracranial self-stimulation rewards; 31 of these 86 were also place-related. These results, together with previous anatomical and behavioral observations, suggest that the spatial information sent from the hippocampus to the LS is modulated by and interacts with signals related to reward in the LS.

The hippocampal formation (HF) is hypothesized as a neu- ronal substrate of a cognitive map, which represents environmental spa- tial information by an ensemble of neural activity. However, the relation- ships between the hippocampal place cells and the cognitive map have not been clarified in monkeys. The present study was designed to inves- tigate how activity patterns of place-selective neurons encode spatial relationships of various environmental stimuli; to do this, we used multi- dimensional scaling (MDS) for hippocampal neuronal activity in the mon- key during the performance of real and virtual translocation. Of 389 neurons recorded from the monkey HF and parahippocampal gyrus (PH), 166 had place fields that displayed increased activity in a specific area of an experimental field and/or on a monitor (place-selective neurons). The MDS transformed relationships among the 16 places in the experimental field and the monitor, expressed as correlation coefficients between all possible pairs of two places based on the 166 place-selective responses, into geometric relationships in a two-dimensional MDS space. In the real translocation tasks, the 16 places were distributed throughout the MDS space, and their relative positions were well correlated to real positions in the experimental laboratory. However, the correlation between the MDS space and real arrangements was significantly smaller in virtual than real translocation tasks. The present results strongly suggest that activity pat- terns of the HF and PH neurons represent spatial information and might provide a neurophysiological basis for a cognitive map. Hippocampus 2003;13:190 -196. © 2003 Wiley-Liss, Inc.

The septal complex, having close and reciprocal connections with the hippocampus, is known to play an important role in learning and memory. Anatomically, the septal complex is divided into the medial and lateral areas (MS and LS). In the present study, in order to elucidate functional differences between the MS and LS, we recorded single unit activity in the MS or LS and electroencephalogram (EEG) in the hippocampus simultaneously while the rats performed the following 2 spatial tasks in an open-field chamber. In task 1, the rat received rewarding intracranial electrical stimulation (ICES) when it entered in a reward place that was set randomly in the open field in each trial. In task 2, the rat received rewarding ICES when it alternately visited two fixed reward places in the open field. Unit activity was analyzed in relation to the pattern of hippocampal EEG, and rat's location, locomotion direction and locomotion speed in the spatial tasks. A total of 47 neurons were recorded in the septal complex (MS, 19; LS, 28). The majority of neurons with activity correlated with hippocampal EEG were found in the MS (14/19). All of the neurons with place-related activity (an increase in unit activity when the rat was in a specific location in the open field) were found in the LS (n = 15). The majority of neurons with direction-related activity were found in the LS (18/23). Twenty-one neurons displayed speed-related activity (MS, 9; LS, 12). The present results indicate that (1) the MS is directly involved in the formation and control of hippocampal EEG patterns, and (2) the LS is important for the processing and integration of spatial information in the environment. Hippocampus 1999; 9:220–234. © 1999 Wiley-Liss, Inc.

The purpose of this study was to investigate, during the performance of an object discrimination task, responses of neurons in the monkey hippocampal formation to the sight of several objects that have biological meaning, and compare these responses with those of amygdalar neurons studied previously using the same task. Neuronal activity in the hippocampal formation of conscious monkeys was recorded during performance of a task that led to presentation of familiar rewarding, familiar aversive, or unfamiliar objects. Of 864 neurons recorded in the hippocampal formation and adjacent cortices, 160 (18.5%) responded to the sight of a certain object(s). Responses to the sight of different kinds of objects were analyzed in detail. Nondifferential neurons (n = 73) responded to different objects with no significant difference in response magnitudes, and differential neurons (n = 87) responded to different objects with different response magnitudes. Of the differential neurons, 23 responded more strongly to rewarding objects than to other objects (rewarding-object-dominant neurons), but the magnitude of responses to objects did not necessarily correlates with the order of preferences to the objects as determined from observation of animal behavior. Aversive-object-dominant neurons (n = 13) responded more to aversive objects than to other objects. Unfamiliar-object-dominant neurons (n = 7) responded more to unfamiliar objects than to familiar objects. Selective neurons (n = 10) responded selectively to only one object or one category of objects. Fourteen of the rewarding- or averse-object-dominant neurons were tested in extinction or reversal trials. In 12 of 14 neurons, responses to a rewarding or aversive object did not change, or slightly weakened, in extinction or reversal trials. The results suggest the following. (1) Responses of rewarding- or aversive-object-dominant neurons may be involved in object-reward or object-aversion association. However, responses of many of these neurons might reflect past inputs to reinforcement rather than extant emotional processing. (2) Responses of unfamiliar-object-dominant neurons may be involved in recognition of objects based on their familiar or unfamiliar aspects. These results are further discussed and compared with responsiveness of amygdalar neurons.