Prelude. Cycle 1. Introduction. Cycle 2. Structure defines function. Cycle 3. Diversity of cortical functions is provided by inhibition. Cycle 4. Windows on the brain. Cycle 5. A system of rhythms: from simple to complex dynamics. Cycle 6. Synchronization by oscillation. Cycle 7. The brain's default state: self-organized oscillations in rest and sleep. Cycle 8. Perturbation of the default patterns by experience. Cycle 9. The gamma buzz: gluing by oscillations in the waking brain. Cycle 10. Perceptions and actions are brain state-dependent. Cycle 11. Oscillations in the "other cortex:" navigation in real and memory space. Cycle 12. Coupling of systems by oscillations. Cycle 13. The tough problem. References.

Rhythms of the brain

The ability to find one's way depends on neural algorithms that integrate information about place, distance and direction, but the implementation of these operations in cortical microcircuits is poorly understood. Here we show that the dorsocaudal medial entorhinal cortex (dMEC) contains a directionally oriented, topographically organized neural map of the spatial environment. Its key unit is the 'grid cell', which is activated whenever the animal's position coincides with any vertex of a regular grid of equilateral triangles spanning the surface of the environment. Grids of neighbouring cells share a common orientation and spacing, but their vertex locations (their phases) differ. The spacing and size of individual fields increase from dorsal to ventral dMEC. The map is anchored to external landmarks, but persists in their absence, suggesting that grid cells may be part of a generalized, path-integration-based map of the spatial environment.

/pdf/microstructure-of-a-spatial-map-in-the-entorhinal-cortex-511e186u79.pdf

Microstructure of a spatial map in the entorhinal cortex

The hippocampal formation can encode relative spatial location, without reference to external cues, by the integration of linear and angular self-motion (path integration). Theoretical studies, in conjunction with recent empirical discoveries, suggest that the medial entorhinal cortex (MEC) might perform some of the essential underlying computations by means of a unique, periodic synaptic matrix that could be self-organized in early development through a simple, symmetry-breaking operation. The scale at which space is represented increases systematically along the dorsoventral axis in both the hippocampus and the MEC, apparently because of systematic variation in the gain of a movement-speed signal. Convergence of spatially periodic input at multiple scales, from so-called grid cells in the entorhinal cortex, might result in non-periodic spatial firing patterns (place fields) in the hippocampus.

Path integration and the neural basis of the 'cognitive map'

More than three decades of research have demonstrated a role for hippocampal place cells in representation of the spatial environment in the brain. New studies have shown that place cells are part of a broader circuit for dynamic representation of self-location. A key component of this network is the entorhinal grid cells, which, by virtue of their tessellating firing fields, may provide the elements of a path integration-based neural map. Here we review how place cells and grid cells may form the basis for quantitative spatiotemporal representation of places, routes, and associated experiences during behavior and in memory. Because these cell types have some of the most conspicuous behavioral correlates among neurons in nonsensory cortical systems, and because their spatial firing structure reflects computations internally in the system, studies of entorhinal-hippocampal representations may offer considerable insight into general principles of cortical network dynamics.

/pdf/place-cells-grid-cells-and-the-brain-s-spatial-4ey9f1b6vx.pdf

Place Cells, Grid Cells, and the Brain's Spatial Representation System

http://eebweb.arizona.edu/Faculty/Dornhaus/courses/materials/papers/Cheng%20spatial%20learning%20module%20in%20rats.pdf

A purely geometric module in the rat's spatial representation*

Homing by path integration in a mammal

The paper presents experimental evidence for homing by path integration in a bird. Using a mammalian model case, the essentials of a cybernetical theory of this type of navigation are developed. As a consequence of its application to the extant data on geese, the necessary information about the translatory component of the animal’s movement along its path appears to be provided by the visual system, viz. the translatory component of the visual flow, whereas the rotatory information must (also) have non-visual sources, e.g. the semicircular canals or the magnetic field.

Homing by Path Integration

When vision is excluded humans are still able to walk back to a starting point or to a previously seen target. This performance may be mediated by path integration, based on information about movement with respect to the ground or to inertial space, that is, on substratal or inertial idiothetic cues. We intend to unravel whether, and how accurately, these two inputs act and interact on the translatory component of this navigation performance. Subjects were asked 1) to reproduce a path they had walked, and 2) to indicate the location of a target they had seen before being blindfolded by (i) walking there, (ii) treading a motor-driven conveyor belt until they imagine they are there, and (iii) reporting, while being driven in a trolley, when they seem to pass the target. The estimation of path length turns out to vary as a function of walking velocity, step length, and step rate. The estimate becomes virtually veridical when subjects walk at their normal pace, but it overshoots at lower and undershoots at higher values of these variables. Veridicality at near normal speeds is also found with passive transport (iii), but with a reverse dependence on velocity. It is concluded that in these paradigms path control and perception are mediated by an open-loop performance of the underlying path integration system, calibrated in such a way as to yield veridical estimates during normal walking. Either inertial or substratal idiothetic information is sufficient for this performance. However, the quantitative relations found argue in favor of the hypothesis that substratal idiothetic information predominates when both are available. In spite of its limitations the capability shown here may serve as an essential constituent of navigation by path integration in humans.

Marie Luise Mittelstaedt

Papers

Homing by path integration in a mammal

Homing by Path Integration

Idiothetic navigation in humans: estimation of path length.

Idiothetic navigation in Gerbils and Humans

Mechanismen der Orientierung ohne richtende Außenreize