The document summarizes Donald O. Hebb's theory of learning and memory from 1949. It discusses how Hebb proposed that learning occurs through synaptic plasticity and the formation of cell assemblies. Specifically, it outlines Hebb's theory that neurons that fire together wire together, and that repeated activation of synapses strengthens the connection between neurons. The document also addresses later critiques of Hebb's synaptic theory of memory and argues that the theory remains relevant when considering both synaptic plasticity and intracellular changes, as conceived in Hebb's original work.

1. 1
N E U R O S C I E N C E
O F L E A R N I N G
H E B B ’ S T H E O R Y
R A C H E L H O N G
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P S Y C H O L O G Y O F L E A R N I N G

2. 2
P R E S E N T A T I O N
C O N T E N T S
Donald O. Hebb’s Theory of Learning and Memory
Trettenbrein’s Critiques of the Neurophysiologic Explanation of Learning and Memory
Resolution of Recent Critiques Using Modern Neurophysiological Research
Reconciliation of the Differences Between Physiologists and Psychologists About the Role of Synaptic Plasticity in
Learning and Memory
Synaptic Change and the Formation of Cell Assemblies are Fundamental for Theories of Memory
Cell Assemblies Have Been Verified by Neuroimaging
Hebb’s Synaptic Learning Rule and Cell Assembly Theory are Used in Computational Neuroscience and
Robotics
Abnormalities in Synaptic Plasticity Underlie Cognitive and Motor Dysfunctions Pain Mechanisms and Drug
Addiction
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3. 3
H I G H L I G H T S
L I T E R A T U R E O N T H E N E U R O B I O L O G Y O F L E A R N I N G A N D M E M O R Y
There is a considerable literature
on the neurobiology of learning and memory
that shows the importance of synaptic plasticity
as the first step in the chain of cellular and
biochemical events involved in memory formation
Once memories are formed, synaptic modification is essential for their expression
(Langille & Brown, 2018).
The discussion will be in terms of Hebb’s (1949) neuropsychological theory.
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4. 4
D O N A L D O . H E B B ’ S ( 1 9 0 4 - 1 9 8 5 )
T H E O R Y O F L E A R N I N G A N D M E M O R Y
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N E U R O N S T H A T F I R E T O G E T H E R W I R E T O G E T H E R .

5. 5
H E B B ' S T H E O R Y
Hebb’s (1949) theory assumed that the neurophysiological changes underlying learning and memory occur
in three stages:
(1) synaptic changes
(2) formation of a “cell assembly”
(3) formation of a “phase sequence”
which link the neurophysiological changes underlying learning and memory as studied by
physiologists to the study of thought, and “mind” as conceived by cognitive psychologists.
Hebb’s neurophysiological assumption (Hebb, 1949) states that:
“When an axon of cell A is near enough to excite a cell B and repeatedly or persistently takes part in firing it,
some growth process or metabolic change takes place in one or both cells such that A’s efficiency, as one of the
cells firing B, is increased.”
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6. 6
H E B B ' S T H E O R Y
The cell assembly is a set of neurons and the
pathways connecting them, which act together
(Hebb, 1949),
such that a stimulus activating pathway 1 will
activate a reverberating circuit of N pathways
(in Hebb’s example, n = 15).
It is a hypothetical reverberating system, proposed
as a mediating process, an element of thought,
capable of holding an excitation and bridging a
gap in time between stimulus and response (Hebb,
1972).
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1, 4 5, 9
7, 15
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C E L L A S S E M B L Y

7. 7
H E B B ' S T H E O R Y
A series of cell assemblies, connected by neural activity over time is a “Phase Sequence,” which provides
the neural basis for a “train of thought” from one cell assembly to another (Hebb, 1949).
The cell assembly “relates the individual nerve cell to psychological phenomenon”
such that “a bridge has been thrown across the great gap between the details of neurophysiology and
the molar conceptions of psychology” (Hebb, 1949).
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P H A S E S E Q U E N C E

8. 8
H E B B ' S T H E O R Y
Hebb elaborated on
How this theory could account for learning and memory
How new learning could be associated with previous learning, and
How “quick learning” (similar to the single trial learning of Gallistel and Balsam (2014)) might occur
(Hebb, 1949).
Hebb’s cell assembly theory showed
how differences between psychologists and physiologists,
who use different definitions for the same phenomena,
could be reconciled into a theory of the neurophysiological basis of learning and memory.
Hebb’s assumption contains two concepts: synaptic plasticity and “some growth process or metabolic
change” in the neuron, which is “intrinsic plasticity” (Titley, Brunel, & Hansel, 2017).
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9. 9
1 0 Y E A R S
L A T E R
The only theory to realistically deal with problems of
behavior, thought process and learning
Theory has defects, but no real competitors
It is criticized, because it is difficult to experimentally prove
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10. 1 0
T R E T T E N B R E I N ’ S C R I T I Q U E S O F T H E
N E U R O P H Y S I O L O G I C E X P L A N A T I O N
O F L E A R N I N G A N D M E M O R Y
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The synapse-centered view of learning and memory is not
focused and that the neurobiological basis of learning and
memory is still unclear (Langille & Brown, 2018).
Trettenbrein (2016) argues that the concept of the synapse as
the locus of memory is not sensible and that a paradigm shift
is necessary.
However, no new paradigm is provided, but he suggests that
“the memory mechanism is (sub-) molecular in nature”.

11. 1 1
T R E T T E N B R E I N ’ S C R I T I Q U E S
There are six critiques of the synaptic plasticity theory of memory in Trettenbrein (2016)’s article:
(1) The synapse may not be the sole locus of learning and memory
(2) A synaptic locus of memory does not fit well with philosophical and cognitive theories of
learning and memory
(3) Memories survive despite synapse destruction and synaptic and (or) protein turn-over
(4) Evidence from spatial training suggests that there is a need to separate learning from memory
(5) Existing learning mechanisms cannot explain information that is encoded in a single trial
(Gallistel and Balsam, 2014)
(6) Memory may be sub-cellular in nature
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12. 1 2
R E S O L U T I O N
O F R E C E N T C R I T I Q U E S
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U S I N G M O D E R N
N E U R O P H Y S I O L O G I C A L
R E S E A R C H

13. 1 3
T R E T T E N B R E I N ’ S C R I T I Q U E S
The critique of synaptic plasticity theory proposed by
Trettenbrein (2016)
can be resolved using Hebb’s synaptic theory,
research based on cell assemblies as components of
neural networks,
and current research on the cellular and molecular
basis of memory formation
to show the nature of synaptic plasticity in understanding
the neurobiology of learning and memory.
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14. 1 4
R E C O N C I L I A T I O N O F T H E D I F F E R E N C E S
B E T W E E N P H Y S I O L O G I S T S A N D P S Y C H O L O G I S T S
A B O U T T H E R O L E O F S Y N A P T I C P L A S T I C I T Y I N
L E A R N I N G A N D M E M O R Y
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Critique of Trettenbrein (2016) focuses on “demise” (death, downfall, disappearance or
final fate) of the synaptic theory of memory.
The synaptic theory of memory has not disappeared, but that there are two components
of this theory: synaptic plasticity and intra-cellular biochemical changes.
The concern is whether “memory” consists of the synaptic changes activated by
intracellular biochemical changes OR the intracellular biochemical changes expressed via
synaptic plasticity (Langille & Brown, 2018).
Memory, as conceived by Hebb, consists inseparably of both synaptic plasticity and
“intrinsic plasticity” of the neurons (Lisman, Cooper, Sehgal, & Silva, 2018).

15. 1 5
S Y N A P T I C C H A N G E A N D T H E F O R M A T I O N O F C E L L A S S E M B L I E S A R E
F U N D A M E N T A L F O R T H E O R I E S O F M E M O R Y
Hebb (1949, 1959) realized that his theory would need revision for new discoveries.
His ideas on synaptic plasticity (Favero, Cangiano, & Busetto, 2014), cell assemblies (Wallace & Kerr, 2010) and
phase sequences (Almeida-Filho et al., 2014) continue to stimulate new research and discussion is a tribute to
his prescience.
Physiological mechanisms of learning and memory (Johansen et al., 2014)
Learning and development (Munakata and Pfaffly, 2004)
Memory span (Oberauer, Jones, & Lewandowsky, 2015)
Decision making (Wang, 2012)
Language learning (Wennekers, Garagnani, & Pulvermüller, 2006).
Posner and Rothbart (2004, 2007) suggested that using his ideas to integrate the disparate branches of
Psychology and Neuroscience.
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16. 1 6
C E L L A S S E M B L I E S H A V E B E E N V E R I F I E D B Y N E U R O I M A G I N G
Hebb’s theories on the neurophysiological basis of learning and memory integrate synaptic neurophysiology
with psychological concepts like attention, perception, thought and mind—the concepts which Pavlov avoided
in his objective approach to memory.
Hebb’s theory effectively integrated Pavlov’s concepts of the physiology of learning with Lashley’s (1932)
criticism that Pavlov ignored psychological concepts.
Neuroimaging studies have shown the usefulness of Hebb’s ideas for understanding both the psychological and
physiological mechanisms of memory.
Memory processes have been shown by fMRI and other neuroimaging methods to be distributed across
many cortical areas (Miyamoto, Osada, & Adachi, 2014).
Christophel, Klink, Spitzer, Roelfsema, & Haynes (2017) showed that different cortical neural networks
are activated in different types of working memory.
O’Neil et al. (2012) found that different cortical regions were activated in recognition memory.
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17. 1 7
T H E I M P O R T A N C E O F H E B B ’ S I D E A S
There is still existence on the Hebb synapse (Brown, 2020).
To name a few:
Graham Collingridge’s paper on Hebb synapses and beyond
Ole Paulson’s paper on Neuromodulation of Hebbian synapses
Zahid Padamsey’s presentation on a new framework for Hebbian
plasticity in the hippocampus.
For understanding cognitive (Takamiya, Yuki, Hirokawa, Manabe, &
Sakurai, 2019)
The focus on synaptic mechanisms is in learning and memory.
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18. 1 8
U S E S O F H E B B ’ S W O R K T O D A Y
Learning and memory
Long-term effects of the environment on
development
Aging
Neurocomputing
Artificial intelligence
Robotics
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19. 1 9
N E U R O P H Y S I O L O G Y O F L E A R N I N G A N D M E M O R Y
The proposal that long-term potentiation was a synaptic model of memory (Bliss, & Collingridge, 1993)
led to a number of examinations of Hebb’s concept of synaptic plasticity (Sweatt, 2016).
The concept of spike timing dependent plasticity (STDP) is built on the concept of the Hebb synapse,
producing the term “Hebbian STDP” (Brzosko, Mierau, & Paulsen, 2019).
The Dynamic Hebbian Learning Model (dynHebb) is developed to support for the complexities of
STDP (Olde Scheper, Meredith, Mansvelder, van Pelt, & van Ooyen, 2018).
McNaughton (2003) wrote about how Hebb’s theory stimulated his research on long-term potentiation
and memory.
Andersen, Krauth and Nabavi (2017) stated that “Hebbian plasticity, as represented by long-term
potentiation and long-term depression of synapses, is the most influential hypothesis to support for
encoding of memories.”
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20. 2 0
C E L L A S S E M B L Y
Cell assembly has led to new research on neural networks (Li, Liu, & Tsien, 2016) and the
molecular mechanisms underlying the cell assembly (Pulvermüller, Garagnani, & Wennekers,
2014).
Harris (2012) stated that “One of the most influential theories for cortical function is the ‘cell
assembly hypothesis’ first proposed over half a century ago (Hebb, 1949)”.
Harris (2005) proposed four experimental tests for the temporal organization of cell assemblies.
Eichenbaum (2018) proposed that cell assemblies
be studied as “units of information processing” to guide research on
“the structure and organization of neural representations in perception and cognition”.
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21. 2 1
C E L L A S S E M B L Y
Buzsaki (2010) defined the cell assembly as the neural syntax of the brain and
suggested ways in which the neural organization of cell assemblies could be understood in the context of both
brain function and brain-machine interfaces.
He proposed that cell assemblies were linked by “dynamically changing constellations of synaptic weights” which
he called “synapsembles” and
suggested that the objective identification of the cell assembly
requires a temporal framework and a reader mechanism
which can integrate the activity of cell assemblies over time.
The result has led to the consideration of Hebbian cell assemblies as the basis for “semantic circuits”
which define “the cortical locus of semantic knowledge” and
to the development of neurocomputational models of brain function (Tomasello, Garagnani, Wennekers, &
Pulvermüller, 2018).
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22. 2 2
P H A S E S E Q U E N C E S
Hebb’s concept of phase sequences as synchronized sets of cell assemblies
has been examined by recording action potentials
from the hippocampus and cortex of actively behaving rats (Almeida-Filho, et al., 2014).
The results suggest that the cell assemblies are the building blocks of neural representations,
while the phase sequences that link cell assemblies are modifiable by new experiences,
modulating the neural connections of cognition and behaviour.
This approach has been used to apply Hebbian learning and cell assemblies
to the construction of neurocomputational models of language learning which simulate the
brain mechanisms of word meaning in “semantic hubs” (Tomasello, et al., 2018).
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23. 2 3
N E U R O C O M P U T I N G
The concept of Hebbian learning is used in
neurocomputing and the development of artificial neural networks (Kuriscak, Marsalek, Stroffek, & Toth,
2015).
The mathematical definition of the change in activity at a Hebb synapse through “synaptic scaling”
has allowed for the quantitative definition of a Hebbian Cell Assembly (Tetzlaff, Dasgupta, Kulvicius, &
Wörgötter, 2015) for use in robotics and artificial intelligence.
Virtual Cell Assembly Robots (CABots)
have been built using cell assemblies as the basis of short- and long-term artificial memories (Huyck, &
Mitchell, 2018) and
the cell assembly has been proposed as the basis for computer simulation of human brain function
(Huyck, 2019).
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24. 2 4
H E B B ’ S S Y N A P T I C L E A R N I N G R U L E A N D C E L L A S S E M B L Y T H E O R Y
A R E U S E D I N C O M P U T A T I O N A L N E U R O S C I E N C E A N D R O B O T I C S
Cell assemblies and phase sequences are used to develop
theories of the cortical control of behavior (Palm, Knoblauch, Hauser, & Schüz, 2014)
network theories of memory (Fuster, 1997) and
computer models of memory processes (Lansner, 2009).
Driven by neurophysiological and biophysical findings, they concern the basic neuronal mechanisms and the detailed temporal
processes of neuronal activation and interaction, and by computational arguments and requirements.
Cell assembly theory has helped in developing the anatomical features that underlie the location of memory storage in the
cortex (Palm et al., 2014).
Hebbian learning rules and cell assemblies
Are applied in computer models of the brain to build neural networks based on STDP (Markram, Gerstner, W., & Sjöström,
2011).
Are currently used in robotics (Calderon, Baidyk, & Kussul, 2013).
Hebbian learning rules are used to control brain-robot interfaces in neurorehabilitation (Takeuchi & Izumi, 2015).
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25. 2 5
A B N O R M A L I T I E S I N S Y N A P T I C P L A S T I C I T Y U N D E R L I E C O G N I T I V E A N D
M O T O R D Y S F U N C T I O N S P A I N M E C H A N I S M S A N D D R U G A D D I C T I O N
The activation of the network of synaptic connections in a cell assembly
requires changes in synaptic strength
to establish the connectivity of the neurons in the cell assembly.
Cell assemblies are a collection of activated synapses and the sufficiently strong activation of these
synapses
causes biochemical changes in the neurons of the cell assembly.
Biochemical changes and gene activation within the neurons of a cell assembly are required to maintain
memories (Li, Liu, & Tsien, 2016).
These involve complex interactions between excitatory and inhibitory synapses (Barron, Vogels,
Behrens, & Ramaswami, 2017).
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26. 2 6
The biochemical changes in the neurons of a cell assembly that are activated by transient changes in synaptic
activity involve epigenetic mechanisms including chromatin remodeling
which drives changes in the transcription and translation of information in the DNA, protein synthesis
and cellular changes underlying learning and memory formation (Vogel-Ciernia & Wood, 2014).
Hebb (1949) stated that the synaptic changes following repeated stimulation at a synapse lead to “some growth
process or metabolic change… in one or both cells such that A’s efficiency, as one of the cells firing B, is increased.”
Neuroscientific research on the cellular and molecular basis of memory in the last 70 years has been finding
these growth processes and metabolic changes that underlie memory (Poo et al., 2016).
Synaptic change is not limited to learning and memory, but forms the basis of neural changes in
perception (Yang, Weiner, Zhang, Cho, & Bao 2011), pain (Luo, Kuner, & Kuner, 2014) and drug addiction
(Lüscher, 2013).
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27. 2 7
Neurological disorders which involve cognitive or motor
dysfunction are the result of synaptic abnormalities (Kouroupi et
al., 2017).
Synaptic dysfunction underlies
neurodevelopmental disorders like autism, Rett syndrome,
Down syndrome and ADHD (Moretto, Murru, Martano,
Sassone, & Passafaro, 2018) and
neurological disorders of adulthood and aging, including
Alzheimer disease, Parkinson’s disease, Huntington’s disease
and multiple sclerosis (Torres, Vallejo, & Inestrosa, 2017)
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Impaired hippocampal long-term potentiation and
consolidation
may struggle in forming new, lasting memories
(Weintraub, Wicklund, & Salmon, 2012), termed
“anterograde amnesia”.
The decreases in synaptic strength (and removal of the
physical substrates of memories) and synapse loss
may erase the past memories in retrograde amnesia,
Both of which are characteristic of Alzheimer’s disease
(Beatty, Salmon, Butters, Heindel, & Granholm, 1988).
A synaptic plasticity theory of memory can demonstrate the memory impairments in neuropathologic conditions like Alzheimer’s disease.

28. 2 8
- R E M E M B E R -
Practice Makes Perfect!
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R E F E R E N C E S
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T H A N K Y O U !
Hope You Enjoy this Presentation!
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