Untitled Flashcards Set
Neural Tube Origins & Organization
1. Give an example of how transcription factors and/or morphogens affect the anterior/posterior and dorsal/ventral patterning of the CNS.
Morphogens: chemical inducers/proteins that determine cell fate
Synthesised and secreted at signaling centres at tissue boundaries of embryonic cells
Regulate expression through concentration-dependency in diffusion
How? Establish and regulate gradient of transcription factors to allow cell specialisation
Neurons destined for anterior neocortex express increased levels of Pax6
Neurons destined for posterior neocortex express increased levels of Emx2
Methodology:
Differences in transcription factors ->
Differences in gene expression & protein production ->
Signals to attract neural precursor cells to appropriate destinations
Dorsal-ventral patterning: sonic hedgehog
Important for neural tube closure & establishing identity of ventral motor neurons in spinal cord
Expressed highly in notochord and floor plate
Increased Shh: Increased ventral-like motor neurons
Decreased Shh: increased dorsal-like interneurons
Blocking Shh:
Prevents transcription -> inhibits differentiation of motor neurons
Effects depend on:
Patched (Ptc1) & smoothened (Smo) surface receptors
Transcription factor Gli1 which promotes motor neuron differentiation
Shh absent:
Ptc1 inhibits Smo
This prevents Gli1 from translocating to nucleus
Decreases motor neurons
Shh present:
Binds to Ptc1
Translocation of Smo to cilium
Gli translocation to nucleus
Increased transcription and differentiation of motor neurons
2. How do BMPs contribute to neuroectoderm formation? Where do BMPs come from (what secretes them)? How are BMPs inhibited? List examples of these inhibitors and where they come from.
Bone Morphogenic proteins:
Growth factors secreted by ectoderm in cell-to-cell signaling that leads ectoderm to become skin
BMP inhibitors:
Secreted by DBL that block BMPs and permit the default state of the dorsal ectoderm to become nervous tissue
Examples:
Noggin
Chordin
follistatin
3. Describe the stages of neurulation.
Neural plate formed 2 weeks post fertilisation
Centre will drop inwards forming the neural groove
2 edges of neural plate thicken and fold forming neural folds
2 edges of fold fuse at centre creating neural tube
4. What are the three primary germ layers? What do they give rise to? Be able to label figures of neurulation and gastrulation processes
Prosencephalon/forebrain
Telencephalon
Olfactory bulb, cerebral cortex, hippocampus, basal ganglia
diencephalon
Retina, thalamus, hypothalamus
mesencephalon/midbrain
Mesencephalon
Midbrain
rhombencephalon/hindbrain
Metencephalon
Cerebrum, pons
Myelencephalon
medulla
Developing Cortex: proliferation, migration, & differentiation
1. Draw interkinetic nuclear migration (IKM). What happens at each of the 5 stages?
Cell attached to ventricular surface and pial surface
Cell migrates towards pial surface
DNA replication happens here
Nucleus contains 2 DNA copies, migrates towards ventricular surface
Cell retracts its process from pial surface
Mitosis occurs, symmetrical division occurs to produce new RGS progenitor cells which will go on to divide
2. What does it mean that the cortex develops inside-out? What method was used to prove this? How does this method work (in detail)?
Deepest cortex layers formed first, superficial layers then added on top
Neurons born earlier will reside more interior
Subplate:
Temporary structure important for proper wiring of cortical neurons
Site of initial thalamic synaptic connections
The layer disappears after development is complete
Neurons migrate away
Thalamic afferents go to cortical plates to specify type of neuron
Cortical plate
Next earliest cells born migrate to the cortical plate, then settle in the deepest layer (layer 6)
Cells that are born later migrate to more superficial layers
Tritiated thymidine experiment
Inject gestating rheus monkey w/ radio-active thymadine to attach to alanine
The earlier the thymadine is given in cortical neuro-development, the deeper in the cortex it should be (layer 6)
This helps label birth dates of cells
3. What are 2 chemical signals are secreted by the marginal zone that influence a) migration, and b) differentiation?
Reelin: stop signal produced by cajal-retzius cells in marginal zone
Signal: stop migrating neurons
Semaphorin 3A: protein secreted by marginal zone
Important for differentiating the characteristic architecture of cortical neurons
Signal:
Repels growing axons from pial surface
Attracts growing dendrites towards pial surface
Wiring the Brain: Initial connections, rearrangement, & critical periods
1. Compare and contrast Sperry’s (chemoaffinity) and Hebb’s (exuberant connections) hypotheses on axon pathway formation.
Chemoaffinity hypothesis: form precedes function
Chemical markers/signals lead azons to find their targets
Exuberant connections hypothesis: functions form connections
Axons are initially connected to multiple targets and are slowly refined based on how well they work/their function
2. Describe the 2 experiments performed by Roger Sperry that proved the chemoaffinity hypothesis.
Newt experiment
Surgically rotate a Newt’s eye 180 degrees
Severed the optic nerve and let it regrow
Tested to see whether the nerve will regrow to its original connection or if it will form a new connection based on the eye rotation
Placed the newt in a body of water and placed bait floating above it
Said that if newt swam up then its sight is no longer upside down and the optic nerve formed new connections to fix it (hebb was right)
But if the newt swam down then that means that the optic nerve grew back to where it was originally connected (sperry was right)
The newt eventually swam downwards, proving the chemoaffinity hypothesis correct
Strip assay experiment: established chemoaffinity presence, how do they work?
Background info:
Temporal retinas usually synapse onto the anterior tectum
Nasal retinas usually synapse onto the posterior tectum
Conducted an experiment where he layed out stripes of alternating anterior and posterior tectum, then brought axons from both the nasal and temporal retinas and wanted to see how they would grow based on stripes
Saw that in terms of the nasal retina, it grew equally onto both stripes with no evident preference
But with the temporal retina, he saw that it only grew on the anterior tectum and not posterior, why? Well there are multiple hypothesis
The anterior tectum produces an attracting signal
The posterior tectum produces a repellant signal
Both
In order to answer why, sperry heat-inactivated the tectum stripes one at a time starting with the anterior
He saw that there was no difference in growth
Then he heat-inactivated the posterior tectum and saw that the axons then spread equally across the stripes
This indicated to him that the posterior tectum had a repellent signal
The chemorepellant is ephrin A5
The anterior tectum had low concentrations of it
The posterior tectum had high concentrations of it
Nasal retina cells had low levels of ephrin A5 receptors, therefore they could not detect its presence and grew equally
3. Explain the main findings of the Verhage paper.
Munc 18 is a crucial protein for synaptic transmission but not development
The cortex can develop normally without it, but once synaptic transmission needs to start occurring and it can’t, the synaptic silence causes degeneration of the cortex
4. Compare and contrast experience-dependent vs. activity-dependent synaptic rearrangement. Provide examples of each
Activity dependent rearrangement: requires correlated activity
Spontaneous correlated activity between ipsilateral retina and postsynaptic LGN
Experience dependent: requires correlated activity and sensory experience
Development of barrel fields happens postnatally based on experience
5. Describe how segregation of LGN layers and ocular dominance columns forms. Relate this to experience-dependent and activity-dependent mechanisms
Ocular dominance columns are columns of the V1 that respond to the ipsilateral/contralateral eye
Retinal ganglion cells make connections with the LGN thalamic inputs to project to certain layers of the LGN
LGN then projects to the visual cortex
6. What are the cellular/synaptic mechanisms that cause the shift in ocular dominance columns following monocular deprivation during the critical period?
Hubel & weasel cat experiment
Briefly deprived a cat from sight in one eye from birth to 2.5 months, then measured its visual abilities at 38 months
Saw that the cat only uses the non-deprived eye for vision, the deprived eye does not have any activated cells and there is no evidence of binocular vision
This proves that there was a shift in the ocular dominance columns based on the visual inputs received postnatally
Critical period: 0-2.5 months for ocular dominance columns in cats
7. Define strabismus. How are critical periods important to this?
Uncorrelated eye movement (do not move in unison)
Not pointed in the same direction
About 5% of newborns are born with this condition, critical period is crucial because if not fixed during it then it will be impossible to fix due to experience dependent rearrangement bc the V1 is now used to uncorrelated activity
Intro to Learning & Memory and Anatomy of Memory
1. Differentiate between declarative and non-declarative memory (and their sub-categories) and the brain regions involved.
Declarative: facts and events
Medial temporal lobe, diencephalon
Semantic: facts
Episodic: events
Non-declarative
Procedural memory
Operant conditioning, striatum
Learning skills and habits that need repetition
Skeletal musculature
Classical, cerebellum
Muscle memory
Emotional response
Classical, amygdala
2. Describe the deficits of patient H.M. From this knowledge, what can conclude regarding systems consolidation of memory (which model is correct)?
Lesion caused amygdala & entorhinal cortex damage
Most of EC & hippocampus missing, perirhinal cortex not intact and medial mammillary nucleus shrunken
Had normal procedural memory as it doesn’t require hippocampus
But had no memory of the specific experiences when he was taught
Had normal working memory as it doesn’t require hippocampus
Could remember a list of 6 numbers but would forget them if distracted
HM suffered from anterograde amnesia (inability to form new declarative memories) & temporal retrograde amnesia (memories right before lesion)
3. Explain how the “what” (meaning, a coherent concept) of memory is stored. Define distributed memory and Hebb’s cell assembly and relate this to the “what” of memory.
4. Draw the circuit involved in sensitization and habituation of the gill withdrawal reflex in the Aplysia.
a. What is the mechanism of habituation? (i.e. what cellular changes cause reduced response)
Touching the siphon -> activates sensory neurons -> form excitatory synapses w/ motor neurons
Transmission between sensory & motor neurons is depressed
This leads to decreased ability of siphon stimuli to evoke gill contraction
Depression is presynaptic and is likely due to decrease in number of synaptic vesicles available for release & inactivation of voltage-dependent calcium channels
b. What is the mechanism of short-term and long-lasting sensitization? (i.e. what cellular changes lead to increased response? What is the role of S-type K-channels?)
Short-term sensitisation: activation of serotonin release onto motor neurons
Prior to tail shock, siphon stimulation evokes AP in sensory nerves
AP has a short duration due to quick membrane repolarisation by:
Standard voltage gated K+ channels
S-type K+ channels (always open)
Tail shocks activate sensory neurons in tail
Sensory neurons excite modulatory interneurons that release serotonin (5HT) onto presynaptic terminal of sensory axons
Serotonin enhances glutamate release from sensory axons
Increased glutamate release causes larger EPSP in motor neuron which causes a larger response (sensitisation)
More in depth details
Serotonin released on sensory neuron, binds to metabotropic serotonin receptors
Binding of serotonin stimulates cAMP production
cAMP binds to PKA which liberates catalytic subunits of pKA
Catalytic subunits phosphorylate S-type K+ channel
Phosphorylation causes a conformational change causing them to close
When they close, they can’t help repolarise the membrane
This prolongs AP, longer depolarisation, increased glutamate release
Long-term maintenance:
larger/more tail shocks increases 5HT release on sensory neuron
This increases PKA which shuts down s-type K+ channels but also activates CREB via phosphorylation
Transcription factor
Phosphorylation of CREB
pCREB binds to DNA, induces ubiquitin hydrolase
Ubiquitin hydrolase: prevents ubiquitin from degrading proteins
Prevents pKA degradation
This causes inc. channels close which prolongs AP and increases glutamate release
Activates other transcription factors
Results in long-term increase in synapse and synaptic growth
5. What are place cells? Grid cells? Where are these cells located? How do these cells differ?
Place cells
In hippocampus
Have receptive fields that fire at certain areas/places
Grid cells
In entorhinal cortex
Fires based on where you are in grid pattern in space
This forms a comprehensive map of where one is in space
Each grid cell fires in multiple location in space forming the grid pattern of the spatial receptive field
6. What are the phases of classical conditioning? What is involved in each?
Acquisition: higher value of US & more salient CS results in better conditioning
Extinction: presenting CS without US
Spontaneous recovery: CR reappears after some time
Cellular Mechanisms of Learning & Memory
1. List the 3 requirements for LTP. Describe experimental evidence that proves these things as requirements for LTP.
pre/post synaptic neurons must be active at the same time
Cooperativity: helps form associations between experiences
Presence of NMDARs
Act as coincidence detectors
Mg-block requires sufficient depolarisation to unblock
Are also permeable to calcium
Presence of calcium
Glutamate release -> AMPARs allow for sufficient depolarisation to unblock NMDARs
Calcium enters via NMDARs, binds and activates camKII
2. Describe the two processes that maintain synaptic homeostasis
a. Metaplasticity
Sets threshold for LTP depending on synaptic history
If LTP is induced before, threshold increases making it harder to continue
NR2B NMDARs: Promote LTP (dec. threshold)
NR2A NMDARs: promote LTD (inc. threshold)
b. Synaptic Scaling
Absolute synaptic effectiveness is adjusted to preserve homeostasis synaptic weight
CAMK IV induces the insertion/removal of NMDARs/AMPARs to balance activity
3. What is spatial summation? How does it contribute to cooperativity of memory encoding? How is this different from temporal summation?
Spacial summation: many pre-synaptic inputs are simultaneously stimulated
Temporal summation: 1 cell sends multiple signals to 1 dendrite
4. What are synaptic tags? What is their importance/function? What molecule is a suspected synaptic tag and what proteins might it be tagging?
A weak stimulation that induces early LTP provides a tag to the synapse that allows them to capture newly synthesised proteins induced by strong stimulating synapse
Transient (2-3 hours)
Newly synthesised proteins consolidate LTP at weak synapse to convert it to late LTP
CAMKII is suspected to be a synaptic tag
Proteins
Densin: cell adhesion molecule (allows CamKI to bind to other proteins)
ABP: adds anchoring sites for AMPARsCad: enlarges both pre and post synapses
5. Describe the actions of CamKII as it relates to LTP (early and late).
Early LTP
CamKII is an important 2nd messenger in LTP
Phosphorylated serine 831 in AMPARs to increase conductance
Increases conductance, entrance of Na+, and allows for depolarisation in less time
Increases number of AMPARs in the membrane
Phosphorylates stargazin on AMPARs to immobilise them in the membrane
Stargazin: protein that allows AMPARs to be anchored to the membrane
Permanently locate them there and reduce likelihood to get recycled
Late LTP: required for long-lasting spine enlargement
Calpain: protease that breaks down cytoskeleton of spine
AVC: provide proteins to rebuild
RC: contain AMPARs
Polyribosomes: site of active translation of proteins
Working Memory & Reconsolidation
1. What is reconsolidation? How does this relate to creating false memories? What influences how well something is remembered?
Reconsolidation: recalling a memory after it had been stored, rendering it an unstable state were it is prone ot perturbations
Influences on how well we remember something:
Sleep deprivation
Attention
Emotional saliency
Whether its been exposed to you before
Motivation
1. Compare and contrast the 3 theories of motivation discussed in class. Describe an example of each theory in real life. (Drive reduction theory, Anticipatory Contrast, Response Deprivation).
Drive-reduction theory: motivation drives us towards homeostasis
Set point compares to internal state and produces a behavioural output
Hedonic impact: “liking” the pleasure gained from a reward
Incentive salience: “wanting” motivation that drives to obtain a reward
Anticipatory contrast: the value of a reward is dependent on our previous experience and expectations
Running speed of mice varies based on expectation and outcome
Response deprivation: any behaviour can serve as a reinforcer or punisher if access to behaviour is limited
Behavioural bliss point: all behaviours have a preferred frequency
2. Does dopamine mediate liking or wanting? What is the experimental evidence for this?
Dopamine encodes incentive motivation to obtain a reward
3. Locate the midbrain dopamine neurons on a figure and know where they project to. How do dopamine neurons fire during reward learning, and how does this relate to reward prediction error (E.g., Figure 16.17)?
Rewards often occur after reward-predicting stimuli
Reward prediction errors occur when there is a mismatch between what is expected and what is received
Dopamine neurons fire to a predictive cue