Study Notes on Synaptic Plasticity, Memory, and Learning
Synaptic Plasticity Overview
Synaptic plasticity refers to the changes in the strengths of synaptic connections in response to experience and neuronal activity.
Key forms of synaptic plasticity include:
Hebb’s rule (1949): Formulated the idea that an increase in the synaptic strength arises from the repeated and persistent stimulation of one neuron by another.
Long-Term Potentiation (LTP) (1970): A long-lasting enhancement in signal transmission between two neurons that results from stimulating them simultaneously.
Long-Term Depression (LTD) (1978): A long-lasting decrease in synaptic strength following a pattern of activity.
Spike-Timing-Dependent Plasticity (STDP) (1997): A biological learning rule that expresses how the relative timing of spikes from the pre- and postsynaptic neurons determines the sign and magnitude of the change in synaptic strength.
Homeostatic plasticity (1998): The ability of neurons to stabilize their activity by adjusting their synaptic strengths in response to changes in overall network activity.
Structural plasticity: Changes in the number and structure of synapses in the brain as a response to learning and memory.
Relationship Between Learning and Memory
Spatial Learning and Memory
The relationship between hippocampal LTP and spatial memory is crucial for understanding memory encoding.
Important concepts:
Memory engram: The physical embodiment of memory in the brain, often theorized to reside within specific neurons and synapses that undergo plastic changes during learning.
Hippocampal LTP is fundamental for spatial learning: For example, experiments show that manipulations that alter hippocampal LTP also impact spatial memory performance.
Research Insights and Methodologies
Experimental Methods to Assess Memory:
Morris Water Maze: An experimental tool used to measure spatial learning and memory, where rats learn to find a hidden platform within a pool of water, demonstrating learning via time to find the platform.
Example data from the Morris Water Maze experiment shows control rats are able to locate the hidden platform faster than hippocampal-lesioned rats, indicating the critical role of the hippocampus in spatial memory.
Control rat: Time to find platform is significantly lower (e.g., 60 seconds).
Hippocampal-lesioned rat: Longer time, reaching up to 110 seconds in some trials.
LTP Mechanisms: Phases and Elements
Early Phase of LTP: Triggered by repeated high-frequency stimulation (HFS), increasing Ca2+ influx through NMDA receptors, leading to activation of adenylyl cyclase, and subsequent cAMP and PKA activity.
Late Phase of LTP: Involves gene expression changes mediated by CREB and c-FOS, solidifying synaptic changes.
Roles of calcium signaling and AMPA receptor insertion in the postsynaptic membrane are emphasized in LTP consolidation.
Molecular Mechanisms of Memory Encoding
CaMKII: A critical molecule for memory; acts as a historical record of local calcium signaling within the neuron.
Autophosphorylated CaMKII can phosphorylate AMPA receptors, increasing their availability on the postsynaptic membrane, potentially enhancing synaptic strength and efficacy.
Hypotheses on Memory and Synaptic Connections
Hypothesis I: Memory is stored in the strength of synaptic connections within neural circuits.
The synaptic weight matrix model could theoretically store vast amounts of information by linking specific input patterns (events) to particular output patterns (memories).
Hypothesis II: Learning processes modify the strengths of these synaptic connections.
This suggests a causal relationship between learning experiences and alterations in synaptic weights.
Experimental Evidence of Learning-Induced LTP
Studies question whether learning induces hippocampal LTP.
Utilizing multielectrode recordings at CA1 to measure neural activity before and after training provides insights.
Findings indicate that a small fraction of electrodes from trained rats exhibited detectable potentiation post-training, supporting the hypothesis of induced synaptic changes.
Saturation of LTP:
Examined how residual LTP affects further learning; findings suggested that minimal residual LTP may indicate saturation and limit the capacity for further synaptic strengthening during learning.
Cellular Mechanisms of Memory Recall
Investigating whether specific neuron populations active during learning can be reactivated to trigger memory recall.
Tagging Neurons: Techniques such as using Fos-tTA mice, where immediate early gene c-Fos under controlled conditions marks active neurons during learning, allows tracking and retrieval experiments.
Expression under specific conditions (absence of doxycycline, dox) confirms the successful tagging and activation of targeted neurons.
Contextual Fear Conditioning:
Activation of neuron populations engaged during fear conditioning can evoke fear responses, demonstrating the functional role of these previously active circuits in memory recall.