Choices, Choices

Hippocampal maps happen in many modes and dimensions. How do they relate to conscious navigation?

How do our brains work? A question that was once mystical is now tantalizingly concrete. Neurobiology is, thanks to the sacrifices of countless rats, mice, and undergraduate research subjects, slowly bringing to light mechanisms by which thoughts flit about the brain. The parallel processing of vision, progressively through the layers of the visual cortex, was one milestone. Another has been work in the hippocampus, which is essential for memory formation as well as mapping and navigation. Several kinds of cells have been found there (or in associated brain areas) which fire when the animal is in a certain place, or crosses a subjective navigational grid boundary, or points its head in a certain direction. 

A recent paper reviewed recent findings about how such navigation signals are bound together and interact with the prefrontal cortex during decision making. One is that locations are encoded in a peculiar way, within the brain wave known as the theta oscillation. These run at about 4 to 12 cycles per second, and as an animal moves or thinks, place cells corresponding to locations behind play at the trough of the cycle, while locations progressively closer, and then in front of the animal play correspondingly higher on the wave. So the conscious path that the animal is contemplating is replayed on a sort of continuous loop in highly time-compressed fashion. And this happens not only while the animal is on the path, but at other times as well, if it is dreaming about its day, or is resting and thinking about its future options.

"For hippocampal place cells to code for both past and future trajectories while the animal navigates through an environment, the hippocampus needs to integrate multiple sensory inputs and self-generated cues by the animal’s movement for both retrospective and prospective coding."

These researchers describe a new piece of the story, that alternate theta cycles can encode different paths. That is, as the wave repeats, the first cycle may encode one future path out of a T-maze, while the next may encode another path out of the same maze, and then repeating back to A, B, etc. It is evident that the animal is trying to decide what to do, and its hippocampus (with associated regions) is helpfully providing mappings of the options. Not only that, but the connecting brain areas heading towards the prefrontal cortex (the nucleus reuniens, entorhinal cortex, and parahippocampal gyrus) separate these path representations into different cell streams, (still on the theta oscillation), and progressively filter one out. Ultimately, the prefrontal cortex represents only one path ... the one that the rat actually chooses to go down. The regions are connected in both directions, so there is clearly top-down as well as bottom-up processing going on. The conclusion is that in general, the hippocampus and allied areas provide relatively unbiased mapping services, while the cortex does the decision making about where to go, and while it may receive.

    "This alternation between left and right begins as early as 25 cm prior to the choice point and will continue until the animal makes its turn"

A rat considers its options. Theta waves are portrayed, as they appear in different anatomical locations in the brain. Hippocampal place cells, on the bottom right, give a mapping of the relevant path repeatedly encoded across single theta wave cycles. One path is encoded in one cycle, the other in the next. Further anatomical locations (heading left) separate the maps into different channels / cells, from which the prefrontal cortex finally selects only the one it intends to actually use.

The hippocampus is not just for visual navigation, however. It is now known to map many other senses in spatial terms, like sounds, smells. It also maps the flow of time in cognitive space, such as in memories, quite apart from spatial mapping. It seems to be a general facility to create cognitive maps of the world, given whatever the animal has experienced and is interested in, at any scale, and in many modalities. The theta wave embedding gives a structure that is highly compressed, and repeated, so that it is available to higher processing levels for review, re-enactment, dreaming, and modification for future planning. 

Thus using the trusty maze test on rats and mice, neuroscientists are slowly, and very painfully, getting to the point of deciphering how certain kinds of thoughts happen in the brain- where they are assembled, how their components combine, and how they relate to behavior. How they divide between conscious and unconscious processes naturally awaits more insight into what this dividing line really consists of.

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