Role of the hippocampus in goal representation : Insights from behavioural and electrophysiological approaches

The hippocampus plays an important role in spatial cognition, as supported by the location-specific firing of hippocampal place cells. In random foraging tasks, each place cell fires at a specific position (‘place field’) while other hippocampal pyramidal neurons remain silent. A recent study evidenced a reliable extra-field activity in most CA1 place cells of rats waiting for reward delivery in an uncued goal zone. While the location-specific activity of place cells is thought to underlie a flexible representation of space, the nature of this goal-related signal remains unclear. To test whether hippocampal goal-related activity reflects a representation of goal location or a reward-related signal, we designed a two-goal navigation task in which rats were free to choose between two uncued spatial goals to receive a reward. The magnitude of reward associated to each goal zone was modulated, therefore changing the goal value. We recorded CA1 and CA3 unit activity from rats performing this task. Behaviourally, rats were able to remember each goal location and flexibly adapt their choices to goal values. Electrophysiological data showed that a large majority of CA1-CA3 place and silent cells expressed goal-related activity. This activity was independent from goal value and rats’ behavioural choices. Importantly, a large proportion of cells expressed a goal-related activity at one goal zone only. Altogether, our findings suggest that the hippocampus processes and stores relevant information about the spatial characteristics of the goal. This goal representation could be used in cooperation with structures involved in decision-making to optimise goal-directed navigation

Social Spaces: Place Cells Represent the Locations of Others

How does the brain represent the location of others? Recordings in rats and bats show that, along with representing self-location in an environment, some hippocampal neurons are modulated by the position of another individual

Volumetric spatial behaviour in rats reveals the anisotropic organisation of navigation

We investigated how access to the vertical dimension influences the natural exploratory and foraging behaviour of rats. Using high-accuracy three-dimensional tracking of position in two- and three-dimensional environments, we sought to determine (i) how rats navigated through the environments with respect to gravity, (ii) where rats chose to form their home bases in volumetric space, and (iii) how they navigated to and from these home bases. To evaluate how horizontal biases may affect these behaviours, we compared a 3D maze where animals preferred to move horizontally to a different 3D configuration where all axes were equally energetically costly to traverse. Additionally, we compared home base formation in two-dimensional arenas with and without walls to the three-dimensional climbing mazes. We report that many behaviours exhibited by rats in horizontal spaces naturally extend to fully volumetric ones, such as home base formation and foraging excursions. We also provide further evidence for the strong differentiation of the horizontal and vertical axes: rats showed a horizontal movement bias, they formed home bases mainly in the bottom layers of both mazes and they generally solved the vertical component of return trajectories before and faster than the horizontal component. We explain the bias towards horizontal movements in terms of energy conservation, while the locations of home bases are explained from an information gathering view as a method for correcting self-localisation

Insensitivity of place cells to the value of spatial goals in a two-choice flexible navigation task

Hippocampal place cells show position-specific activity, thought to reflect a self-localization signal. Several reports also point to some form of goal encoding by place cells. We investigated this by asking whether they also encode the value of spatial goals, which is a crucial information for optimizing goal-directed navigation. We used a continuous place navigation task in which male rats navigate to one of two (freely chosen) unmarked locations and wait, triggering the release of reward which is then located and consumed elsewhere. This allows sampling of place fields, and dissociates spatial goal from reward consumption. The two goals varied in the amount of reward provided, allowing assessment of whether the rats factored goal value into their navigational choice, and of possible neural correlates of this value. Rats successfully learned the task, indicating goal localization, and they preferred higher-value goals, indicating processing of goal value. Replicating previous findings, there was goal-related activity in the out-of-field firing of CA1 place cells, with a ramping-up of firing rate during the waiting period, but no general over-representation of goals by place fields, an observation that we extended to CA3 place cells. Importantly, place cells were not modulated by goal value. This suggests that dorsal hippocampal place cells encode space independently of its associated value, despite the effect of that value on spatial behavior. Our findings are consistent with a model of place cells in which they provide a spontaneously constructed value-free spatial representation, rather than encoding other navigationally relevant, but non-spatial, information

Contribution of Cerebellar Sensorimotor Adaptation to Hippocampal Spatial Memory

Complementing its primary role in motor control, cerebellar learning has also a bottom-up influence on cognitive functions, where high-level representations build up from elementary sensorimotor memories. In this paper we examine the cerebellar contribution to both procedural and declarative components of spatial cognition. To do so, we model a functional interplay between the cerebellum and the hippocampal formation during goal-oriented navigation. We reinterpret and complete existing genetic behavioural observations by means of quantitative accounts that cross-link synaptic plasticity mechanisms, single cell and population coding properties, and behavioural responses. In contrast to earlier hypotheses positing only a purely procedural impact of cerebellar adaptation deficits, our results suggest a cerebellar involvement in high-level aspects of behaviour. In particular, we propose that cerebellar learning mechanisms may influence hippocampal place fields, by contributing to the path integration process. Our simulations predict differences in place-cell discharge properties between normal mice and L7-PKCI mutant mice lacking long-term depression at cerebellar parallel fibre-Purkinje cell synapses. On the behavioural level, these results suggest that, by influencing the accuracy of hippocampal spatial codes, cerebellar deficits may impact the exploration-exploitation balance during spatial navigation