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lectures from Dr. Kathleen Munley
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What types of experiences can change the brain? What is the common feature of these experiences?
Development
Culture
Preferences
Coping
Learning is common feature of these experiences
What is neuroplasticity?
The brain’s lifelong ability to reorganize its structure, functions, and connections in response to experiences, learning, or injury
What is learning?
The active process of acquiring new understanding, behaviors, skills, or attitudes through study, experience, or practice
What is memory?
The mental faculty of encoding, storing, and retrieving information
Acts as record of experiences that guides future actions
What is an engram?
A mental representation of a previous experience
Memory trace
Corresponds to physical change in the brain, most likely involving synapses
What is Pavlovian conditioning?
A learning procedure in which neutral stimulus (CS) elicits a response (CR) because of its repeated pairing with an event (UCS)
Also called classical conditioning or respondent conditioning
What is the process of Pavlovian conditioning in terms stimulus and response?
CS + UCS → UCR
After several pairings: CS → CR
What is a conditioned stimulus (CS)?
An originally neutral stimulus that triggers conditioned response (CR) after association with unconditioned stimulus (UCS)
What is an unconditioned stimulus (UCS)?
A stimulus that naturally and automatically triggers an unconditioned response (UCR)
What is an unconditioned response (UCR)?
An unlearned, naturally occurring response to a UCS, such as salivation when food is in the mouth
What is a conditioned response (CR)?
A learned response to a formerly neutral CS
What is eyeblink conditioning?
Pavlovian conditioning of the blink response:
Tone (CS) is associated with a painless puff of air (UCS) to participant’s eye
Blinking is normal reaction (UCR) to a puff of air
Learning has occurred when blinking occurs due to CS alone (CR)

What is fear conditioning?
A behavioral learning process in which neutral stimulus becomes associated with an aversive event, causing the neutral stimulus to elicit fear responses (Eg. freezing, elevated heart rate)
What is operant conditioning?
A learning procedure that modifies voluntary behavior through consequences-reinforcement or punishment
Forms associations between actions and results to change behavior frequency
Is operant learning localized a specific brain circuit?
No, necessary circuits vary with task requirements
What is implicit memory?
Long-term, unconscious memory that allows individuals to perform tasks, skills, and emotional responses automatically without conscious effort
What is explicit memory?
Long-term, conscious, internal recollection of factual information, previous experiences, and concepts
What is priming?
Using a stimulus to sensitize the nervous system to later presentation of the same or a similar stimulus
Enables unconscious learning
What is the Gollin figure test?
A psychological assessment that measures visual perceptual skills and implicit memory
During retention test, participants identify image sooner, indicating some form of memory for the image
Subjects with amnesia also show improvement on test, even though they do not recall having taken it before
What is amnesia?
Partial or total loss of memory
What is the pursuit-rotor task?
A psychology assessment measuring hand-eye coordination and motor skill learning
How do amnesics perform on implicit memory tasks?
The same as non-amnesics
When presented with same task one week later, both controls and amnesics take less time to perform it
Amnesics fail to recall having performed the task before
What is declarative memory?
Long-term memory of facts, events, and concepts that can be consciously recalled and verbalized
What is procedural memory?
Long-term memory responsible for performing tasks automatically, without conscious thought
What is a learning set?
An implicit understanding of how problem can be solved using rule that can be applied in many different situations
What are difference in brain processing implicit vs explicit memory?
Implicit information is processed in a bottom-up or data-driven manner (via sensory input)
Explicit information is processed in a top-down or conceptually-driven manner (via prior knowledge)
What are the differences in task roles a person has with implicit vs explicit tasks?
In implicit tasks, person has passive role
In explicit tasks, person has active role
What is short-term memory? What parts of the brain are involved?
The brain’s temporary, limited capacity storage system in which information is held only for a few minutes, then discarded
Involves the frontal lobes
What is long-term memory? What part of the brain are involved?
The brain’s nearly permanent, high-capacity storage system, in which information is held indefinitely
Involves the temporal lobe
What parts of the brain are involved in short-term vs long-term memory?
Frontal lobes are involved in short-term memory
Temporal lobes are involved in long-term memory
What are the multiple memory systems summarized?

What is the semantic memory system?
The brain’s repository for general knowledge, facts, concepts, and word meanings
What brain regions are part of/involved with the semantic memory system?
Largely supported by network in left-lateralized temporal and inferior parietal cortex
Subregions relatively specialized for specific object characteristics or types of knowledge

When is the semantic memory system engaged?
It is similar to a default network that is active when participants are resting, rather than engaged in specific cognitive tasks
Semantic processing constitutes large component of cognitive activity during passive states
What is autobiographical memory?
Episodic memories (specific events) and semantic memories (facts) about one’s life
What parts of the brain are involved in autobiographical memory?
Involves ventromedial prefrontal cortex (vmPFC), hippocampus, and pathways between them
What is highly superior autobiographical memory (HSAM)?
People who display virtually complete recall for events in their lives, beginning around age 10
Many can describe any episode, including date and day of the week
Who was Henry Molaison (H.M.)? What happened to his memory?
He received bilateral medial temporal lobe resection due to recurrent seizures
After surgery, H. M. had severe amnesia and lacked explicit memory
Despite these deficits, H. M. had above- average IQ, performed well on perceptual tests, and could recall childhood events
H. M.’s performance on implicit memory tests was left intact
What are the primary structures for explicit memory?
Medial temporal region (Image of a monkey brain)
Hippocampus
Amygdala
Entorhinal cortex
Parahippocampal cortex
Perirhinal cortex
Prefrontal cortex
Other closely related structures

What is the parahippocampal cortex?
Medial temporal region involved in memory and visuospatial processing
Receives connections from parietal cortex
What is the perirhinal cortex?
Medial temporal region that specializes in visual object recognition, memory, and perception
Receives connections from visual regions of the ventral stream
What is the entorhinal cortex?
Medial temporal region that functions in episodic memory, navigation, and spatial awareness
Receives projections from parahippocampal and perirhinal cortices
What is visuospatial memory?
The cognitive system for storing and processing visual and spatial information, which is used to identify an object’s location in space
What part of the brain is involved in visuospatial memory?
The hippocampus is important for visuospatial memory
Laboratory animals and human patients with selective hippocampal injury have severe deficits in spatial memory
Monkeys with hippocampal lesions have difficulty learning location of objects
Animals with good spatial memory tend to have larger hippocampi than species with poorer spatial memories
What are the four classes of spatial cells?
Place cells
Place-by-direction cells (less focus here)
Head direction cells
Grid cells
What are place cells?
Spatial cells in the hippocampal formation that fire only when animal is in specific, localized area of its environment (place field), regardless of orientation
What are head direction cells?
Spatial cells in the hippocampal formation that function as “neural compass” and fire selectively when animal’s head faces specific direction
What are grid cells?
Spatial cells in the hippocampal formation that function as “neural GPS” and fire at many locations, forming virtual grid (grid field) invariant to changes in animal’s direction, movement, or speed
Where specifically are the spatial cells found in the hippocampal formation?
Place cells and head direction cells are found in the hippocampus and are closely related structures
Grid cells are found in the entorhinal cortex, a major afferent route into the hippocampus

How is the frontal lobe involved in short-term memory?
The frontal lobe participates in many forms of short-term memory
All sensory systems project to frontal lobes:
During tasks in which monkeys must keep information in short-term memory over a delay, some neurons in medial prefrontal cortex exhibit sustained firing
Animals that have not learned task show no such activity
What directions do the neural circuits travel for explicit vs implicit memory?
Neural circuit for explicit memory is reciprocal
Neural circuit for implicit memory is unidirectional
What is the reciprocal neural circuit for explicit memory?
Note that information flow begins with inputs from the sensory and motor systems, which are not considered part of the explicit memory circuit

What is the unidirectional neural circuit for implicit memory? What key structures are involved?
Basal ganglia
Ventral thalamus
Substantia nigra
Premotor cortex

What is the role of the basal ganglia in implicit memory?
Receive sensory and motor input from neocortex and send projections to ventral thalamus
What is the role of the ventral thalamus in implicit memory?
Sends projections to premotor cortex
What is the role of the substantia nigra in implicit memory?
Dopaminergic neurons project to basal ganglia
What is the role of the premotor cortex in implicit memory?
Stores short-term information that biases motor behavior and stimulus-induced learning
What is the neural circuit for emotional memory?

What is non-associative learning?
A form of learning that involves change in behavioral response to a single, repeated stimulus
What is habituation?
A form of non-associative learning in which neural responses to repeated, irrelevant stimuli are suppressed over time
Allows animals to filter noise and focus on salient signals
What is the neural basis of habituation?
Lies in change in presynaptic calcium (Ca2+) channels
Involves reduced sensitivity of calcium channels and decrease in neurotransmitter release
What is sensitization?
A form of non-associative learning characterized by enhanced response to a noxious stimulus
Serves as adaptive mechanism to detect and protect against damage
What is the neural basis for sensitization?
Neural circuits participating in sensitization differ from those for habituation
Involves delayed opening of potassium (K+) channels, resulting in longer-lasting action potential
What is associative learning?
The process in which animal learns to connect stimuli or events, resulting in lasting changes in behavior
Produces enduring neural change in postsynaptic cell after EPSP from presynaptic cell crosses synaptic cleft
What is long-term potentiation (LTP)?
The persistent strengthening of synapse based on recent, high-frequency activity pattern
What is long-term depression (LTD)?
The selective weakening of synapse based on low-frequency stimulation
What is the basis for understanding synaptic changes that underly learning and memory?
LTP is associated with memory formation, whereas LTD is associated with forgetting or clearing memory traces
Enhanced LTP in recruited pathways is exhibited when animals learn to solve problems
LTP produces enduring changes in synaptic morphology, resembling those seen in memory
What are the two receptor types that glutamate acts on?
AMPA receptors
NDMA receptors
What are AMPA receptors?
Receptors on the postsynaptic membrane that respond normally to glutamate
Mediate fast synaptic transmission
What are NMDA receptors?
Doubly gated channels that are typically blocked by magnesium (Mg2+) ions
Serve as coincidence detectors and play role in learning and memory
What two events must occur for NMDA receptors to open?
Depolarization of postsynaptic membrane, which displaces Mg2+ from pore (strong electrical stimulus)
Activation by glutamate from presynaptic neuron (weak electrical stimulus)
How are the lasting effects of glutamate seen at different levels of electrical stimulation?

What receptors and ions are required for long-term potentiation (LTP)?
NMDA receptors and calcium influx are required for LTP

What actions do GABA nterneurons show?
Similar phenomena to LTP and LTD
Plasticity of GABAergic (inhibitory) synapses modulate networks of excitatory neurons
What is inhibitory long-term potentiation (iLTP)?
Activity-dependent, long-lasting increase in strength of inhibitory synapses
What is inhibitory long-term depression (iLTD)?
Persistent, activity-dependent reduction in strength of inhibitory synapses
How are existing neural circuits modified?
Neurons change their structure in response to changing experiences
Changes in number of dendrites can be used to infer synaptic changes (Eg. more dendrites provide more connections)
New synapses can form between already connected neurons or between neurons that were not previously connected

What neuroanatomical/structural changes are seen in learning and memory of sea slugs (Aplysia)?
In Aplysia, neuroanatomical changes can occur in sensory and motor neurons involved in gill and siphon withdrawal reflex
During habituation, number of synapses between sensory and motor neuron decrease
During sensitization, number of synapses between sensory and motor neuron increase
Structural changes may underlie enduring memories

How are new neural circuits created in adulthood?
Adult neurogenesis occurs in mammals
Occurs in olfactory bulb, hippocampalformation, and possibly neocortex
What is BrdU (5-boromo-2’-deoxyuridine) staining?
A histological technique for detecting proliferating cells
how is adult neurogenesis beneficial?
It may enhance brain plasticity, especially processes underlying learning and memory
Why are their structural differences in cortical neurons of male and female rats?
Differences depend on gonadal hormones (e.g., androgens and estrogens)
How is cortical cell structure influenced in human females?
Gonadal hormones influence cell structure and behavior in human females across the menstrual cycle
Reduced estrogen levels produce increased number of spines on pyramidal cells throughout neocortex, but decreased spine density in hippocampus

What are glucocorticoids? What is their function?
Steroid hormones secreted by adrenal glands that regulate metabolism, inflammation, and immune system
Crucial for mobilizing energy sources (Eg. proteins, carbohydrates) during stress response
How is the hypothalamic-pituitary-adrenal (HPA) axis regulated?
Medial prefrontal cortex (mPFC) and hippocampus have important roles in negative feedback regulation of HPA axis
Increased glucocorticoid levels affect dendritic morphology in mPFC and hippocampus
Chronic stress exposure may be neurotoxic
What is a traumatic brain injury (TBI)?
Damage to brain resulting from external force that causes impartments in function
Main causes: falls, accidents, violence
What are three ways to recover from a TBI?
Compensation
Reorganization
Neurogenesis
What is compensation?
Way to recover from a TBI that involves learning to solve problems in new ways
What is reorganization?
Way to recover from a TBI the involves forming new neural connections and “doing more with less”
What is neurogenesis?
Way of recovering from a TBI that involves stimulating the generation of new neurons to produce new circuits