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20 Minute Quiz. For each of the two questions, you can use text, diagrams, bullet points, etc. What are the main events in neural firing and transmission? Describe the main events in neural development. . How does activity lead to structural change?.

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20 minute quiz
20 Minute Quiz

For each of the two questions, you can use text, diagrams, bullet points, etc.

  • What are the main events in neural firing and transmission?
  • Describe the main events in neural development.
how does activity lead to structural change
How does activity lead to structural change?
  • The brain (pre-natal, post-natal, and adult) exhibits a surprising degree of activity dependent tuning and plasticity.
  • To understand the nature and limits of the tuning and plasticity mechanisms we study
    • How activity is converted to structural changes (say the ocular dominance column formation)
  • It is centrally important for us to understand these mechanisms to arrive at biological accounts of perceptual, motor, cognitive and language learning
    • Biological Learning is concerned with this topic.
learning and memory introduction







Learning and Memory: Introduction

facts about a situation

general facts


skill and fact learning may involve different mechanisms
Skill and Fact Learning may involve different mechanisms
  • Certain brain injuries involving the hippocampal region of the brain render their victims incapable of learning any new facts or new situations or faces.
    • But these people can still learn new skills, including relatively abstract skills like solving puzzles.
  • Fact learning can be single-instance based. Skill learning requires repeated exposure to stimuli.
  • Implications for Language Learning?
models of learning
Models of Learning
  • Hebbian ~ coincidence
  • Recruitment ~ one trial
  • Supervised ~ correction (backprop)
  • Reinforcement ~ delayed reward
  • Unsupervised ~ similarity
hebb s rule
Hebb’s Rule
  • The key idea underlying theories of neural learning go back to the Canadian psychologist Donald Hebb and is called Hebb’s rule.
  • From an information processing perspective, the goal of the system is to increase the strength of the neural connections that are effective.
hebb 1949
Hebb (1949)

“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”

From: The organization of behavior.

hebb s rule8
Hebb’s rule
  • Each time that a particular synaptic connection is active, see if the receiving cell also becomes active. If so, the connection contributed to the success (firing) of the receiving cell and should be strengthened. If the receiving cell was not active in this time period, our synapse did not contribute to the success the trend and should be weakened.
ltp and hebb s rule



LTP and Hebb’s Rule
  • Hebb’s Rule: neurons that fire together wire together
  • Long Term Potentiation (LTP) is the biological basis of Hebb’s Rule
  • Calcium channels are the key mechanism
chemical realization of hebb s rule
Chemical realization of Hebb’s rule
  • It turns out that there are elegant chemical processes that realize Hebbian learning at two distinct time scales
    • Early Long Term Potentiation (LTP)
    • Late LTP
  • These provide the temporal and structural bridge from short term electrical activity, through intermediate memory, to long term structural changes.
calcium channels facilitate learning
Calcium Channels Facilitate Learning
  • In addition to the synaptic channels responsible for neural signaling, there are also Calcium-based channels that facilitate learning.
    • As Hebb suggested, when a receiving neuron fires, chemical changes take place at each synapse that was active shortly before the event.
long term potentiation ltp
Long Term Potentiation (LTP)
  • These changes make each of the winning synapses more potent for an intermediate period, lasting from hours to days (LTP).
  • In addition, repetition of a pattern of successful firing triggers additional chemical changes that lead, in time, to an increase in the number of receptor channels associated with successful synapses - the requisite structural change for long term memory.
    • There are also related processes for weakening synapses and also for strengthening pairs of synapses that are active at about the same time.
the hebb rule is found with long term potentiation ltp in the hippocampus
The Hebb rule is found with long term potentiation (LTP) in the hippocampus

Schafer collateral pathway

Pyramidal cells

1 sec. stimuli

At 100 hz


During normal low-frequency trans-mission, glutamate interacts with NMDA and non-NMDA (AMPA) and metabotropic receptors.

With high-frequency stimulation

early and late ltp
Early and late LTP
  • (Kandel, ER, JH Schwartz and TM Jessell (2000) Principles of Neural Science. New York: McGraw-Hill.)
  • Experimental setup for demonstrating LTP in the hippocampus. The Schaffer collateral pathway is stimulated to cause a response in pyramidal cells of CA1.
  • Comparison of EPSP size in early and late LTP with the early phase evoked by a single train and the late phase by 4 trains of pulses.
computational models based on hebb s rule
Computational Models based onHebb’s rule

The activity-dependent tuning of the developing nervous system, as well as post-natal learning and development, do well by following Hebb’s rule.

Explicit Memory in mammals appears to involve LTP in the Hippocampus.

Many computational systems for modeling incorporate versions of Hebb’s rule.

  • Winner-Take-All:
      • Units compete to learn, or update their weights.
      • The processing element with the largest output is declared the winner
      • Lateral inhibition of its competitors.
  • Recruitment Learning
      • Learning Triangle Nodes
  • LTP in Episodic Memory Formation
hebbian learning takes place
Hebbian learning takes place






Category node 2 now represents ‘at’

category 1 is established through hebbian learning as well
Category 1 is established through Hebbian learning as well






Category node 1 now represents ‘to’

distributed vs localist rep n36
What happens if you want to represent a group?

How many persons can you represent with n bits? 2^n

What happens if one neuron dies?

How many persons can you represent with n bits? n

Distributed vs Localist Rep’n
recruiting connections
Recruiting connections
  • Given that LTP involves synaptic strength changes and Hebb’s rule involves coincident-activation based strengthening of connections
    • How can connections between two nodes be recruited using Hebbs’s rule?







Finding a Connection in Random Networks

For Networks with N nodes and branching factor,

there is a high probability of finding good links.

(Valiant 1995)


Recruiting a Connection in Random Networks

  • Informal Algorithm
  • Activate the two nodes to be linked
  • Have nodes with double activation strengthen their active synapses (Hebb)
  • There is evidence for a “now print” signal based on LTP (episodic memory)










hebb s rule is not sufficient
Hebb’s rule is not sufficient
  • What happens if the neural circuit fires perfectly, but the result is very bad for the animal, like eating something sickening?
    • A pure invocation of Hebb’s rule would strengthen all participating connections, which can’t be good.
    • On the other hand, it isn’t right to weaken all the active connections involved; much of the activity was just recognizing the situation – we would like to change only those connections that led to the wrong decision.
  • No one knows how to specify a learning rule that will change exactly the offending connections when an error occurs.
    • Computer systems, and presumably nature as well, rely upon statistical learning rules that tend to make the right changes over time. More in later lectures.
hebb s rule is insufficient


tastes rotten

eats food

gets sick

drinks water

Hebb’s rule is insufficient
  • should you “punish” all the connections?
models of learning48
Models of Learning
  • Hebbian ~ coincidence
  • Recruitment ~ one trial
  • Next Lecture: Supervised ~ correction (backprop)
  • Reinforcement ~ delayed reward
  • Unsupervised ~ similarity
constraints on connectionist models
Constraints on Connectionist Models

100 Step Rule

Human reaction times ~ 100 milliseconds

Neural signaling time ~ 1 millisecond

Simple messages between neurons

Long connections are rare

No new connections during learning

Developmentally plausible

5 levels of neural theory of language
5levels of Neural Theory of Language

Pyscholinguistic experiments

Spatial Relation

Motor Control



Cognition and Language


Structured Connectionism


Neural Net and learning


Triangle Nodes

Computational Neurobiology


Neural Development




short term memory
Short term memory
  • How do we remember someone’s telephone number just after they tell us or the words in this sentence?
  • Short term memory is known to have a different biological basis than long term memory of either facts or skills.
    • We now know that this kind of short term memory depends upon ongoing electrical activity in the brain.
    • You can keep something in mind by rehearsing it, but this will interfere with your thinking about anything else. (Phonological Loop)
long term memory
Long term memory
  • But we do recall memories from decades past.
    • These long term memories are known to be based on structural changes in the synaptic connections between neurons.
    • Such permanent changes require the construction of new protein molecules and their establishment in the membranes of the synapses connecting neurons, and this can take several hours.
  • Thus there is a huge time gap between short term memory that lasts only for a few seconds and the building of long-term memory that takes hours to accomplish.
  • In addition to bridging the time gap, the brain needs mechanisms for converting the content of a memory from electrical to structural form.
situational memory
Situational Memory
  • Think about an old situation that you still remember well. Your memory will include multiple modalities- vision, emotion, sound, smell, etc.
  • The standard theory is that memories in each particular modality activate much of the brain circuitry from the original experience.
  • There is general agreement that the Hippocampal area contains circuitry that can bind together the various aspects of an important experience into a coherent memory.
  • This process is believed to involve the Calcium based potentiation (LTP).
dreaming and memory
Dreaming and Memory
  • There is general agreement and considerable evidence that dreaming involves simulating experiences and is important in consolidating memory.