Engram (neuropsychology)

An engram is a unit of cognitive information imprinted in a physical substance, theorized to be the means
by which memories are stored[1] as biophysical or biochemical[2] changes in the brain or other biological
tissue, in response to external stimuli.

Demonstrating the existence of, and the exact mechanism and location of, neurologically defined engrams
has been a focus of persistent research for many decades.[3]

Contents
History
Overview
See also
References
Further reading

History
The term "engram" was coined by memory researcher Richard Semon in reference to the physical substrate
of memory in the organism. Semon warned, however: "In animals, during the evolutionary process, one
organic system—the nervous system—has become specialised for the reception and transmission of stimuli.
No monopoly of this function by the nervous system, however, can be deduced from this specialisation, not
even in its highest state of evolution, as in Man."[4] One of the first ventures on identifying the location of a
memory in the brain was undertaken by Karl S. Lashley who removed portions of the brain in rodents. In
Lashley's experiments, rats were trained to run through a maze and then tissue was removed from their
cerebral cortex. Increasing the amount of tissue removed increased the degradation of memory, but more
remarkably, where the tissue was removed from made no difference. His search thus proved unsuccessful,
and his conclusion – that memory is diffusely distributed in the brain – became widely influential.[5]
However, today we appreciate that memory is not completely but only largely distributed in the brain; this,
together with its dynamic nature makes engrams challenging to identify using traditional scientific
methods.[5][6]

Later, Richard F. Thompson sought the engram in the cerebellum, rather than the cerebral cortex. He used
classical conditioning of the eyelid response in rabbits in search of the engram. He puffed air upon the
cornea of the eye and paired it with a tone. After a number of experiences associating it with a tone, the
rabbits became conditioned to blink when they heard the tone even without a puff. One region that
Thompson's group studied was the lateral interpositus nucleus (LIP). When it was deactivated chemically,
the conditioned response disappeared; when re-activated, they responded again, demonstrating that the LIP
is a key element of the engram for this response.[7] This approach, targeting the cerebellum, though
successful, examines only basic, automatic responses, which virtually all animals possess. However,
engrams of specific types of memory are found in the subsystems mediating that learning process and as
such solely engrams of simple conditioning are associated with the LIP but not, for instance, engrams of
semantic memory.
Overview
Neuroscience acknowledges the existence of many types of memory and their physical location within the
brain is likely to be dependent on the respective system mediating the encoding of this memory.[8] Such
brain parts as the cerebellum, striatum, cerebral cortex, hippocampus, and amygdala are thought to play an
important role in memory. For example, the hippocampus is believed to be involved in spatial and
declarative memory, as well as consolidating short-term into long-term memory.

Studies have shown that declarative memories move between the limbic system, deep within the brain, and
the outer, cortical regions. These are distinct from the mechanisms of the more primitive cerebellum, which
dominates in the blinking response and receives the input of auditory information directly. It does not need
to "reach out" to other brain structures for assistance in forming some memories of simple association.

An MIT study found that behavior based on high-level cognition, such as the expression of a specific
memory, can be generated in a mammal by highly specific physical activation of a specific small
subpopulation of brain cells. By reactivating these cells by physical means in mice, such as shining light on
neurons affected by optogenetics, a long-term fear-related memory appears to be recalled.[9]

Another study used optogenetics and chemogenetics to control neuronal activity in animals encoding and
recalling the memory of a spatial context to investigate how the brain determines the lifetime of memories.
The results found by the researchers have defined a role for specific hippocampal inhibitory cells
(somatostatin expressing cells) in restricting the number of neurons involved in the storage of spatial
information and limiting the duration of the associated memory.[10]

In 2016, an MIT study found that memory loss in early stages of Alzheimer's disease could be reversed by
strengthening specific memory engram cell connections in the brains of Alzheimer mouse models.[11]

See also
Multiple trace theory
Samskara

References
1. Liu, Xu; Ramirez, Steve; Pang, Petti T.; Puryear, Corey B.; Govindarajan, Arvind; Deisseroth,
Karl; Tonegawa, Susumu (22 March 2012). "Optogenetic stimulation of a hippocampal
engram activates fear memory recall" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC33319
14). Nature. 484 (7394): 381–385. Bibcode:2012Natur.484..381L (https://ui.adsabs.harvard.e
du/abs/2012Natur.484..381L). doi:10.1038/nature11028 (https://doi.org/10.1038%2Fnature1
1028). PMC 3331914 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3331914).
PMID 22441246 (https://pubmed.ncbi.nlm.nih.gov/22441246).
2. Ryan, T. J.; Roy, D. S.; Pignatelli, M.; Arons, A.; Tonegawa, S. (28 May 2015). "Engram cells
retain memory under retrograde amnesia" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC55
83719). Science. 348 (6238): 1007–1013. Bibcode:2015Sci...348.1007R (https://ui.adsabs.h
arvard.edu/abs/2015Sci...348.1007R). doi:10.1126/science.aaa5542 (https://doi.org/10.112
6%2Fscience.aaa5542). PMC 5583719 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC558
3719). PMID 26023136 (https://pubmed.ncbi.nlm.nih.gov/26023136).
3. Levy, Adam (14 January 2021). "Memory, the mystery" (https://knowablemagazine.org/articl
e/mind/2021/memory-mystery). Knowable Magazine. doi:10.1146/knowable-011421-3 (http
s://doi.org/10.1146%2Fknowable-011421-3). Retrieved 25 March 2022.
 4. Semon, Richard (1921). "Chapter II. Engraphic Action of Stimuli on the Individual" (https://ar
chive.org/stream/cu31924100387210#page/n31/mode/2up). The Mneme. London: George
Allen & Unwin. p. 24; trans by Louis Simon.
5. Sa, Josselyn; S, Köhler; Pw, Frankland (September 2015). "Finding the Engram" (https://pub
med.ncbi.nlm.nih.gov/26289572/). Nature reviews. Neuroscience. PMID 26289572 (https://p
ubmed.ncbi.nlm.nih.gov/26289572). Retrieved 2020-06-02.
6. Bruce, Darryl (1 December 2001). "Fifty Years Since Lashley's In Search of the Engram:
Refutations and Conjectures". Journal of the History of the Neurosciences. 10 (3): 308–318.
doi:10.1076/jhin.10.3.308.9086 (https://doi.org/10.1076%2Fjhin.10.3.308.9086).
PMID 11770197 (https://pubmed.ncbi.nlm.nih.gov/11770197).
7. James W. Kalat, Biological Psychology p. 392–393
8. Gerrig and Zimbardo (2005) Psychology and Life (17th edition: International edition)
9. Costandi, Mo (6 April 2012). "Light brings back bad memories" (https://www.theguardian.co
m/science/neurophilosophy/2012/apr/06/1). The Guardian.
10. Stefanelli, Thomas; Bertollini, Cristina; Lüscher, Christian; Muller, Dominique; Mendez,
Pablo (March 2016). "Hippocampal Somatostatin Interneurons Control the Size of Neuronal
Memory Ensembles" (https://doi.org/10.1016%2Fj.neuron.2016.01.024). Neuron. 89 (5):
1074–1085. doi:10.1016/j.neuron.2016.01.024 (https://doi.org/10.1016%2Fj.neuron.2016.01.
024). PMID 26875623 (https://pubmed.ncbi.nlm.nih.gov/26875623).
11. Roy, Dheeraj S.; Arons, Autumn; Mitchell, Teryn I.; Pignatelli, Michele; Ryan, Tomás J.;
Tonegawa, Susumu (March 2016). "Memory retrieval by activating engram cells in mouse
models of early Alzheimer's disease" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC48477
31). Nature. 531 (7595): 508–512. Bibcode:2016Natur.531..508R (https://ui.adsabs.harvard.
edu/abs/2016Natur.531..508R). doi:10.1038/nature17172 (https://doi.org/10.1038%2Fnature
17172). PMC 4847731 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4847731).
PMID 26982728 (https://pubmed.ncbi.nlm.nih.gov/26982728).

Further reading
Forgotten Ideas, Neglected Pioneers: Richard Semon and the Story of Memory, Daniel
Schacter, 2001 ISBN 1-84169-052-X
Dudai, Y (2004). "The neurobiology of consolidations, or, how stable is the engram?".
Annual Review of Psychology. 55: 51–86. doi:10.1146/annurev.psych.55.090902.142050 (ht
tps://doi.org/10.1146%2Fannurev.psych.55.090902.142050). PMID 14744210 (https://pubme
d.ncbi.nlm.nih.gov/14744210).
Gonzalez, Robbie (July 25, 2013). Memory implantation is now officially real (https://gizmod
o.com/memory-implantation-is-now-officially-real-909746570) Gizmodo. Retrieved
December 3, 2021.
Josselyn, S.A. (2010). "Continuing the search for the engram: examining the mechanism of
fear memories" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2895151). Journal of
Psychiatry and Neuroscience. 35 (4): 221–228. doi:10.1503/jpn.100015 (https://doi.org/10.1
503%2Fjpn.100015). PMC 2895151 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC289515
1). PMID 20569648 (https://pubmed.ncbi.nlm.nih.gov/20569648).

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