Neuroscience of Learning: Hebb's Theory

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PSYCHOLOGY OF LEARNING
NEUROSCIENCE OF LEARNING HEBBS THEORY
HEBB'S THEORY
NEUROSCIENCE OF LEARNING
RACHEL HONG
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• Donald O. Hebbs Theory of Learning and Memory
• Trettenbreins 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
• Hebbs 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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HEBB'S THEORY
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• 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 Hebbs (1949)
  neuropsychological theory.

NEUROSCIENCE OF LEARNING
HEBB'S THEORY
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NEUROSCIENCE OF LEARNING
HEBB'S THEORY
DONALD O. HEBBS (1904 - 1985) THEORY OF LEARNING
AND MEMORY
NEURONS THAT FIRE TOGETHER WIRE TOGETHER.
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HEBB'S THEORY

• Hebbs (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.
• Hebbs 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 As efficiency, as one of the cells
  firing B, is increased.

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HEBB'S THEORY
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HEBB'S THEORY

• 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 Hebbs 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).

CELL ASSEMBLY
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5, 9
NEUROSCIENCE OF LEARNING
HEBB'S THEORY
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2, 14
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HEBB'S THEORY

• 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).

NEUROSCIENCE OF LEARNING
HEBB'S THEORY
PHASE SEQUENCE
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HEBB'S THEORY

• 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).
• Hebbs 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.
• Hebbs 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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HEBB'S THEORY
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• 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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HEBB'S THEORY
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TRETTENBREINS CRITIQUES OF THE NEUROPHYSIOLOGIC
EXPLANATION OF LEARNING AND MEMORY

• 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.

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HEBB'S THEORY
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TRETTENBREINS CRITIQUES

• 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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HEBB'S THEORY
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RESOLUTION OF RECENT CRITIQUES
USING MODERN NEUROPHYSIOLOGICAL RESEARCH
NEUROSCIENCE OF LEARNING
HEBB'S THEORY
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TRETTENBREINS CRITIQUES

• The critique of synaptic plasticity theory
  proposed by Trettenbrein (2016)
• can be resolved using Hebbs 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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HEBB'S THEORY
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RECONCILIATION OF THE DIFFERENCES BETWEEN
PHYSIOLOGISTS AND PSYCHOLOGISTS ABOUT THE ROLE OF
SYNAPTIC PLASTICITY IN LEARNING AND MEMORY

• 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).

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HEBB'S THEORY
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SYNAPTIC CHANGE AND THE FORMATION OF CELL
ASSEMBLIES ARE FUNDAMENTAL FOR THEORIES OF MEMORY

• 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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HEBB'S THEORY
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CELL ASSEMBLIES HAVE BEEN VERIFIED BY NEUROIMAGING

• Hebbs theories on the neurophysiological basis
  of learning and memory integrate synaptic
  neurophysiology with psychological concepts like
  attention, perception, thought and mindthe
  concepts which Pavlov avoided in his objective
  approach to memory.
• Hebbs theory effectively integrated Pavlovs
  concepts of the physiology of learning with
  Lashleys (1932) criticism that Pavlov ignored
  psychological concepts.
• Neuroimaging studies have shown the usefulness of
  Hebbs 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.
• ONeil et al. (2012) found that different
  cortical regions were activated in recognition
  memory.

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HEBB'S THEORY
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THE IMPORTANCE OF HEBBS IDEAS

• There is still existence on the Hebb synapse
  (Brown, 2020).
• To name a few
• Graham Collingridges paper on Hebb synapses and
  beyond
• Ole Paulsons paper on Neuromodulation of Hebbian
  synapses
• Zahid Padamseys 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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HEBB'S THEORY
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USES OF HEBBS WORK TODAY

• Learning and memory
• Long-term effects of the environment on
  development
• Aging
• Neurocomputing
• Artificial intelligence
• Robotics

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HEBB'S THEORY
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NEUROPHYSIOLOGY OF LEARNING AND MEMORY

• The proposal that long-term potentiation was a
  synaptic model of memory (Bliss, Collingridge,
  1993)
• led to a number of examinations of Hebbs 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 Hebbs 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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HEBB'S THEORY
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CELL ASSEMBLY

• 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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HEBB'S THEORY
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CELL ASSEMBLY

• 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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PHASE SEQUENCES

• Hebbs 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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HEBB'S THEORY
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NEUROCOMPUTING

• 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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HEBB'S THEORY
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HEBBS SYNAPTIC LEARNING RULE AND CELL ASSEMBLY
THEORY ARE USED IN COMPUTATIONAL NEUROSCIENCE AND
ROBOTICS

• 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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HEBB'S THEORY
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ABNORMALITIES IN SYNAPTIC PLASTICITY UNDERLIE
COGNITIVE AND MOTOR DYSFUNCTIONS PAIN MECHANISMS
AND DRUG ADDICTION

• 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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HEBB'S THEORY
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• 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 As 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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HEBB'S THEORY
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• 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, Parkinsons disease,
  Huntingtons disease and multiple sclerosis
  (Torres, Vallejo, Inestrosa, 2017)

• 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 Alzheimers
  disease (Beatty, Salmon, Butters, Heindel,
  Granholm, 1988).

NEUROSCIENCE OF LEARNING
HEBB'S THEORY

• A synaptic plasticity theory of memory can
  demonstrate the memory impairments in
  neuropathologic conditions like Alzheimers
  disease.

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HEBB'S THEORY
- REMEMBER -
Practice Makes Perfect!
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HEBB'S THEORY
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