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Memory is a complex skill involving perception, encoding, storage, recall. If the underpinning it relies on fails, it doesn't work as it should. Worse, even at its best, it does not work like a video stored on a shelf somewhere, or a jpeg, stored on a hard drive, but it is fallible, as Elizabeth Loftus and her colleagues have shown, especially when we are young. But don't look for her name in this book, she is not there. And the title is misleadingly simple: the book is complex indeed.
Hasselmo is looking at the behavioral and neural dynamics of episodic memory, the memory of what has happened or what has yet to happen, rather than the overlearned memory of how to perform a skill or specific facts about things in the world. Space and time are coded in episodic memory pathways, linking events and episodes, with memory and drug-guided behavior all interlinked. Now you have an idea of the 7 chapter headings.
An episode is thus an experience we recall during a specific time interval and in a specific location. If you don't get excited about grid cells and their relationship to oscillatory dynamics in the entorhinal cortex, then this book is not for you but rather for those who are serious about the cornus ammoni, viz. Ammon's Horn, and all that this means for neuropsychology and neuroscience in general.
Essential to understanding this book is to understand the existence of multiple forms of memory, namely for the name of a state in the USA (semantic memory), or tracking a phone number in your head (working memory), or how to drive a car (procedural memory), or what you had for breakfast on your birthday (episodic memory) for which this book was written, in terms of matching the behavior of memory with the physiology of the substrates of the brain that produces it that constitutes then episodic memory.
In examining the neuronal pathways that attend memory in the hippocampus and the areas tending to it, especially the entorhinal pathways, Hasselmo draws frequently on an analogy of an entire orchestra, seating different instruments in different but connected and adjacent areas, and responding on different levels to different aspects of the conductors and transcription demands, including both pitch and timing in their instructions. He makes reference as well to the rhythms of the orchestra as emanating from Theta activity, and takes us through the complex mechanisms and structures we would encounter in negotiating our way through the CA fields mostly, and the surrounding areas. This model thus demonstrates the oscillatory dynamics of memory formation in the mesial temporal lobes where the hippocampus lives. Reciprocal richness of memory suggests connectedness to the sensory areas which provided the input in the first place. The 'what' of a memory and the 'where' of a memory are still however encoded separately. Here, he describes what grid cell spacing is, for those rendered curious by what I wrote early on. You will gather then that this book is not for beginners, but for serious students of the minutiae of memory formation down to cells and firing, and not to how one remembers that a rabbit is called a rabbit, and what one does down a maze chasing rats.
Central to episodic memory formation is the coding of space and time, and as I noted before, separate entities have to converge into a single recall. In this chapter, the idea of spiking and phase coding is reduced to the analogy of two toy monkeys, both clanging symbols at 1Hz. When the one monkey on a unicycle,(yes you heard me) speeds up its clanging in response to the phase angle a spot on its wheel moves through, and when it stops, although it was once in synch with the other monkey, it is now out of synch, the extent to which it is so can be predicted by the exact distance travelled by the red spot on the wheel. This is phase coding of spatial location by analogy. When the spiking of the first neuron is in synch with the second, their spikes will be in phase, but this will occur and fail recursively as the second moves and hence oscillates faster. The same will happen with Theta phasing, and I caution that the wavelength of Theta here is a bit wider than human, as much of this work is done on rodents.
The author then discusses how biophysical models of the entorhinal cortex, the most important neighbor of the hippocampus proper, can account for a range of intrinsic properties of the area. Hippocampal cells can read and store the code, since the phase of grid cell oscillations depends on location, and can be accessed by other neuronal structures for the encoding and retrieval of special location in episodic memory.
Encoding and retrieval of episodic trajectories is pursued in a similar vein. If you think of the analogy he uses around parking his car in different places in the same building on different days, you can see that simply place cells alone cannot encode the different locations without a trajectory model of how the car got there. With place cells alone, only the most recent trajectory can be followed. The more recent strength replaces the earlier strength in the memory trace, although you could think it through to find for instance your peppermints you dropped near yesterday's parking spot. Grid cells and place cells thus show stable firing over multiple different exposures to an environment even with long delays between exposures. This requires however association with the sensory input, as sensory cues are necessary to set the phase of firing to the same starting point to ensure stability.
Episodes and events thus have to be linked in some way, since events are a central point in episodic memory, and we have to associate locations and times along a spatiotemporal trajectory with specific items and events. Here, he will describe the hippocampal circuitry for associations as well as the function of hippocampal subregions, including region CA3 and its role in associative memory. The interesting fact here is that after initial and complex excitation, the same memory can be re-associated by firing just a subset of the original circuits, and the recall is whole, even when the cues are incomplete, something called pattern completion. Elements of a memory may thus be used in a kind of priming to reproduce multiple elements of a more complex memory. This opens the door for imperfect memory, and the brain will accept false memories when they are full of complex detail.
This issue of how the CA3 works in associative memory is then discussed in detail, including the role of the dentate gyrus in sorting out the overlapping memory that will result. Region CA1 is also a target for extensive discussion, as it is intimately connected by the Schaffer collaterals, as strongly as are the excitatory connections from the CA3 to the Mossy fibers, but the pattern of connectivity is vastly different to the pattern in the CA1. The Mossy fiber input from the dentate to the CA3 stops before it reaches CA1, CA1 fields have only very weak recurrent connections, so the vast majority of synapses in the stratum radiatum of CA1 arise only from the Schaffer collateral pathway from CA3, and finally, CA1 receives input from layer III of the entorhinal cortex rather than II, and this input is segregated with input from medial entorhinal cortex entering areas of CA1 proximal to CA3, and input from the lateral entorhinal cortex entering CA1 more distal near the subiculum. Now this may seem meaningless or complex, but in reality to neuroscientists this describes markedly divergent functions in these two areas, which leads to implications for functional studies of the behavior of memory. Input from the CA3 thus can head off down the Mossy fibers, and through the fornix to the mammillary bodies, but not from the CA1 directly, suggesting different pathways for heteromodal information. There is in effect no direct connection from the CA3 to the entorhinal cortex other than via the CA1 to the subiculum. This will be of importance as the CA1 will act as a gating mechanism via Hebbian modification, mapping CA3 activity to the associated activity in entorhinal cortex and gating how strongly the CA3 output spreads back to influence the entorhinal cortex. In this way, information comes in from the perirhinal areas, richly encoded heteromodal information which is sent through the entorhinal area and via the CA3 and mossy fibers through the fornix arches and mammillary bodies, to be further encoded by long term potentiation. Another option opened up is that the CA1 and Schaffer collaterals provide access to the subiculum for further encoding, using the rich perceptual information provided by the sensorium, namely the tertiary association areas, and so the encoding and heteromodality of this process enriches learning and memory. I hope that summary is accurate, and helps demonstrate the value of what Hasselmo is describing.
Hasselmo examines the value of using drugs that can modify the dynamics of encoding and retrieval, looking at acetylcholine blockers and inhibition enhancers respectively, and how these work in relationship to other studies and to his modeling of the physiology of memory. The value of the models here is in solving the problems of the impairment or enhancement of memory using drugs that influence the acetylcholinergic and GABA receptors, and alter the Theta oscillations. The next two chapters examine then how the mechanisms of encoding and retrieval of episodic memory can be used for memory -- guided behavior in behavioral tasks, and also the mathematical models of memory that kind of prove his points, as an appendix.
By now, in my superficial glide over the book chapters you will have sensed that there is probably no more serious work on memory than this book, and not for the layman or undergraduate, not any more than say Gazzaniga's New Cognitive Neurosciences, or Kandel, Shwartz and Jessell's Principles of Neural Sciences are. This is dense, complex physiology that requires expert analysis, clearly written but requiring intense and detailed appreciation of the area in science, and an awareness of the literature and science evolution across the 2000's until now.
A very comprehensive volume that takes a lot of time and effort to absorb, but worth it for the serious researcher, and the knowledgeable neuroscience, but a tough read for those with little knowledge of the area. The ramifications of this book are unclear, apart from its role in educating those who want a detailed knowledge, and intend to study it further, but throughout are interesting, state of the art tidbits for the clinician, or those with a fine eye for diagnostic detail who are trying to make sense of complex presentations.
© 2012 Roy Sugarman
Roy Sugarman PhD, Director of Applied Neuroscience, Athletes Performance, USA.