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Exam 2
Motor - Hippocampus
338
Other
Graduate
11/13/2020

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Term
what is the goal of neural engineering?
Definition
provide electrical interfaces to the nervous system to ameliorate disease states
Term
what does an array of electrodes that is inserted into the cochlea of a deaf person allow to happen?
Definition
allows the person to hear due to the microphone that picks up sound & transduces it to a receiver inside the skull which is converted into a series of electrical pulses that stimulates various electrodes at different spots along the basilar membrane
Term
are we close to having retinal prosthetics?
Definition
no (still some time away from being clinically viable)
Term
idea behind retinal prosthetics
Definition
replace the activity of the damaged retina to allow for visual stimuli to be experienced
Term
is deep brain stimulation (DBS) commonly used for dystonia & Parkinson's Disease?
Definition
yes (has been well proved in over 100,000 people!)
Term
are brain-computer interfaces (BCI) to restore motor function clinically viable?
Definition
not yet!
Term
goal of BCI to restore motor function
Definition
to ameliorate paralysis
Term
cursor on screen can be controlled by electrodes implanted in ______ so that we can map the direction of movement associated with the different neural firings
Definition
primary motor cortex
Term
what can Jan (paralyzed woman from Pitt) do with her BCI?
Definition
control a robotic arm to be able to even feed herself!
Term
6 reasons reaching is difficult
Definition
1. it takes years to learn
2. redundant degrees of freedom
3. muscles are complex
4. spinal reflexes
5. inputs & outputs are in different formants
6. you have to pre-plan movements
Term
Morasso task to study the properties of arm movements
Definition
subjects were asked to move an object on a table from one location to another
Term
Morasso results from studying the properties of arm movements
Definition
there are many regularities in the reaches
Term
what was the shape of the hand velocity data profile from Morasso's reaching experiments?
Definition
"bell-shaped velocity profile"
Term
Fitts' Law as it pertains to reaching regularities
Definition
there is a speed/accuracy tradeoff
Term
"2/3 power law" as it pertains to reaching regularities
Definition
speed is relative to curvature
Term
how does handwriting represent the scale & effector invariance of arm movements?
Definition
writing with different hands or different utensils look similar
Term
are repeated arm movements similar or different?
Definition
remarkably similar
Term
are arm movements smooth or coarse?
Definition
smooth
Term
Flash & Hogan model for explaining why hand movements were smooth & straight with the bell-shaped velocity
Definition
we perform movements in a manner that aim to decrease jerk (jerk = derivative of acceleration)
Term
who proposed the signal-dependent noise theory of motor control?
Definition
Wolpert & Harris
Term
who proposed the optimal feedback control theory of motor control?
Definition
Todorov & Jordan & Scott
Term
who proposed the internal models theory of motor control?
Definition
Ito
Term
3 (safe) assumptions made by the signal-dependent noise theory of motor control
Definition
1. the goal of motor control is to make accurate movements
2. neural control signals are Poisson: the noise scales with the signal
3. noise accumulates over the duration of the movement
Term
signal-dependent theory of motor control
Definition
for a given movement duration, the neural command minimizes error OR for a given error tolerance, the neural command maximizes the speed
Term
3 parts to the optimal feedback control theory of motor control
Definition
1. a multiple-input, multiple-output system may have redundancies
2. when such a system is noisy, the redundant space can be exploited
3. "Don't correct errors that don't hurt you"
Term
which motor control theory is described by this example system:
1. goal: X1 + X2 = 2 --> this is not difficult! you set both inputs to equal 1, then the problem is solved
2. suppose the system is perturbed & you need to get back to your goal state --> moving back to your solution manifold is the minimum intervention solution
3. even if it is not exactly at 1,1 --> it may be easier to settle at 1.8,0.2 to still have the end goal be the same even though it is not exactly where it was at the perturbation
Definition
optimal feedback control
Term
"minimal intervention principle" (Todorov & Jordan)
Definition
the nervous system tolerates & does not correct variability that does not impact task performance
Term
what principle calls for "do not correct variability that does not matter"
Definition
"minimal intervention principle"
Term
benefit of OFC (optimal feedback control)
Definition
smaller command signals
Term
hallmark of an OFC (optimal feedback control) solution
Definition
structured variability
Term
what is the thinking behind the internal models theory of motor control?
Definition
-if feedback were fast, you could just start moving & correct as you go
-instead, the brain builds an internal representation of the body & the environment
Term
3 non-ideal aspects of muscles
Definition
1. force depends on length & velocity
2. sluggish
3. they only work in one direction
Term
the same neural command to a muscle can be interpreted differently depending on the ______!
Definition
current length/velocity of the muscle
Term
what body part is made up of sliding filaments & cross bridges?
Definition
muscles
Term
force of muscle depends on ______
Definition
length
Term
experiments conducted on ______ demonstrated that muscle force depends on length
Definition
single muscle fibers
Term
muscle spindle
Definition
in parallel with muscle fibers; detect muscle length
Term
golgi tendon organ
Definition
in series with muscle fibers; detect muscle force
Term
muscle spindles regulate ______
Definition
posture
Term
what reflex is described in this example: if someone pours water into your cup, you can react to the cup getting heavier & lift it back up by tightening muscles more as opposed to just dropping the cup if you could not adjust your muscles
Definition
"myotatic reflex"
Term
how did Evarts learn that M1 codes muscles?
Definition
spike-triggered averaging to identify neurons that project directly to the spinal cord; EMG recorded activity of wrist muscles & electrode was placed in M1 --> could see result from M1 AP --> EMG muscle spike
Term
how was this question tested: does this M1 cell care about the movement or the muscle?
Definition
monkey would manipulate the object which could be weighted with various weight; looked at EMG from wrist flexor & wrist extensor; recorded from pyramidal tract neuron
Term
how does Georgeopoulos & Schwartz learn that M1 codes movement?
Definition
center-out reach task
Term
Georgeopoulos & Schwartz center-out reach task conclusion
Definition
a population of neurons cares about the direction of had movement & does not need to care about the force needed for the hand to get there
Term
how did Strick learn that M1 codes both movement & muscle?
Definition
designed an apparatus that could disentangle the wrist movement & movement of the cursor on the screen; sometimes the monkey had to be pro (wrist-up), mid (wrist-sideways), or sup (wrist-down)
Term
Strick M1 coding experiment results
Definition
1. some neurons were muscle-like
2. some neurons were extrinsic-like
3. some neurons were kind of both
Term
the eye-to-hand reference frame transformation consists of: ______
Definition
visual receptive fields combine with proprioception to yield limb-centered tuning which drives the muscles!
Term
series of experiments demonstrated that neurons in parietal cortex responded for reaching movements with how reach movements related to ______
Definition
gaze direction
Term
Churchland & Shenoy believed that M1 codes ______
Definition
dynamics
Term
what is meant by "variability is conserved" if M1 codes dynamics?
Definition
if one neuron is firing a little higher, an associated neuron will fire a little higher too! to keep their population response equivalent
Term
M1 coding for ______ is described by this statement: common motifs of responses are the same depending on the action being made (i.e. reach to left or reach to right)
Definition
dynamics
Term
what is the firs step in looking at structured variability across a neural population according to Churchland & Shenoy?
Definition
throw away the mean response
Term
goal of neural prosthetics
Definition
help someone who has a spinal cord injury so basically, they can still think about actions & make motor plans --> neural prosthetics can reanimate the muscles in response to the neural signals
Term
Collinger does a lot of _______ work
Definition
neural prosthetics
Term
Schwartz neural prosthetics
Definition
can get a monkey BCI to grasp different objects
Term
what are used in BCI systems to allow for looking at the activity of multiple neurons recorded simultaneously during a single trial --> use these signals to control a cursor on a computer screen?
Definition
multielectrode array recordings
Term
most BCI work is done using ______
Definition
monkey neural activity & mapping that activity to move a cursor on a screen
Term
3 steps in calibrate the BCI for "observation-based calibration"
Definition
1. we move the cursor right or left
2. neurons are active
3. map those activity patterns to cursor kinematics
Term
how can a monkey control a BCI?
Definition
if the monkey can generate those same patterns that aroused when he watched the cursor on the screen, he can control the movement with his neural activity to hit the target
Term
three decode algorithms for BCI
Definition
1. population vector
2. linear filter
3. kalman filter
Term
what task was used to determine the population vector algorithm of decoding neural activity?
Definition
center-out reach task
Term
3 parts of the population vector algorithm for BCI decoding
Definition
1. neurons are broadly tuned
2. many neurons active during each reach
3. "population code" for movement direction
Term
4 steps for the population vector algorithm for BCI decoding
Definition
1. fit each neuron with a cosine
2. estimate the preferred direction
3. weight each preferred direction by the cell's firing weight
4. sum all the weighted vectors
Term
intuition behind linear filter for BCI decoding
Definition
each spike contributes a little pulse of movement...add those together
Term
simple way to describe linear filter for BCI decoding
Definition
it is just movement direction = neural activity with some involvement of noise & other free parameters
Term
Bayesian statistics is an effort to ameliorate two different kinds of predictors: ______
Definition
1. prior knowledge
2. observed evidence
Term
the following is an example of what kind of statistics: if you are asked if it is going to rain tomorrow, your prior knowledge would be what time of year it is (e.g. October vs. July) & your observation would be what the weather looks like overnight (e.g. dark & storm vs. warm & clear)
Definition
Bayesian
Term
how do you do Bayesian statistics with reaches for BCI (kalman filter decoding)?
Definition
using the trajectory model (prior knowledge of arm position/trajectory) & observation (neural) model, we compute at each time point:
-arm estimate: a function of its previous state & the current neural activity
-arm's dynamics + neural activity
Term
what is the motivating question behind latent variable modeling?
Definition
what does the activity of a neuron really tell us?
Term
dysfunction at the _______ can result in motor (PD, HD), cognitive (ADHD, Tourette's), & affective (OCD) symptoms
Definition
basal ganglia
Term
basal ganglia output is directed at what two areas of cerebral cortex?
Definition
1. 'motor'
2. 'non-motor'
Term
is the input-output system of the basal ganglia highly topographic?
Definition
yes
Term
does the basal ganglia communicate with the cerebellum?
Definition
yes
Term
is the dorsal or ventral striatum the source of a major 'open loop' input to the cerebral cortex?
Definition
ventral
Term
the basal ganglia collects information to direct ______
Definition
motor output
Term
multiple regions of cerebral cortex project into the ______ (the input structure of the BG)
Definition
neostriatum
Term
output nucleus of BG
Definition
GPi
Term
cerebellar loop with the cerebral cortex
Definition
cerebral cortex --> cerebellar cortex --> influence the major output nuclei of the cerebellum --> major output to ventral-lateral nucleus of thalamus --> drove motor cortex
Term
BG motor circuit with the cerebral cortex
Definition
M1 --> striatum --> GPi/SNr --> VL thalamus --> back to M1
Term
five 'circuits' that incorporate the BG
Definition
1. skeleto-motor
2. oculo-motor
3. dorsolateral prefrontal
4. lateral orbitofrontal
5. anterior cingulate
Term
what is the big picture idea concerning the five circuits that incorporate the BG?
Definition
multiple circuits from distinct areas of the cerebral cortex go to distinct portions of the striatum to control these overall distinct functional/anatomical loops
Term
how were the five circuits that incorporate the BG tested to see if they actually existed?
Definition
use rabies virus to do retrograde transport to test out the circuits
Term
big cool thing about using rabies for anatomical tracing
Definition
it can be transported up through second order neurons!
Term
basal ganglia have the capacity to influence cortical areas involved in what 4 domains?
Definition
1. movement
2. cognition
3. affect
4. visuo-spatial perception
Term
are the basal ganglia just motor structures?
Definition
NO!
Term
topographic layout of the striatum
Definition
limbic, executive, motor (with individual leg/arm/face regions), & then limbic again
Term
what is another name for the caudate putamen?
Definition
striatum
Term
how does PD affect striatal function?
Definition
when the SNc degenerates (particularly the ventral part), there is a DA depletion from the motor component of the striatum
Term
does the SNc project to the whole striatum?
Definition
YES!
Term
one pro & one con for using L-DOPA as a treatment for PD
Definition
pro: it helps replenish DA

con: it may cause too much DA in parts of the striatum that do not need it!
Term
HD is deemed symptomatic when ______
Definition
the patients begin to show motor symptoms
Term
looking at pre-HD pathology, there are issues with _______ areas of striatum
Definition
limbic
Term
as HD progresses, it spreads to involve ______ & ______ function
Definition
motor & executive
Term
what was the goal of experiments where they injected bicuculline (GABAergic antagonist) into various sections of the BG?
Definition
shut off certain cells in the various striatum regions (limbic, motor, executive, executive striatum specifically) --> gave credence to the hypotheses earlier of the different segments within the GPe
Term
why are bicuculline injections reversible dysfunction?
Definition
it is quickly metabolized
Term
executive territory GPe bicuculline injection --> ?
Definition
movements are normal BUT monkey does not stop moving!
Term
executive territory striatum bicuculline injection --> ?
Definition
monkey makes very few movements & if he makes one it is terribly slow
Term
motor territory arm representation GPe bicuculline injection --> ?
Definition
monkey demonstrated dyskinesia (abnormal movements)
Term
limbic territory GPe bicuculline injection --> ?
Definition
nail biting continuous for 2 hours!
Term
how did the experimenters ensure that the monkey tests where different areas of BG influence different cortical areas to influence behavior in different ways was true anatomically (not just that it behaviorally made sense)?
Definition
did a rabies injection test
Term
limbic bicuculline injection in GPe --> ?
Definition
stereotypy following injection
Term
associative bicuculline injection in GPe --> ?
Definition
hyperactivity following injection
Term
sensorimotor bicuculline injection in GPe --> ?
Definition
dyskinesia following injection
Term
is there a topography of function & of output in the GPe?
Definition
yes
Term
how was it discovered that there is cerebellar output to the basal ganglia?
Definition
injected rabies into striatum --> first order neurons in thalamus --> further retrograde transneuronal transport to second-order neurons in deep cerebellar nuclei!
Term
how was it discovered that there is basal ganglia output to the cerebellum?
Definition
injected rabies into cerebellum --> first order retrograde neurons in pons --> further retrograde transport to second order STN in BG!
Term
output stage of BG (STN) --> disynaptically to _______
Definition
cerebellar input
Term
output stage of cerebellum (deep nuclei) --> disynaptically to ______
Definition
BG input
Term
how did brain scans from PD patients link cerebellar & basal ganglia activation?
Definition
they show that the cerebellum is very much lit up on the scans!
Term
______ is "characterized by reduced metabolic activity in prefrontal & parietal cortex, associated with relative increases in the cerebellum & dentate nuclei"
Definition
PD
Term
how do tics in Tourette syndrome show that cerebellar & basal ganglia activation are linked?
Definition
Tourette = classic BG disorder, but scans show cerebellum as activated as well!
Term
how does appetitive classical conditioning show that cerebellar & basal ganglia activation are linked?
Definition
cerebellum was scanned as a control, but it turned out to be as activated as the BG during the conditioning!
Term
we think of cerebellum as learning through ______ & BG as learning through ______
Definition
cerebellum = synaptic plasticity

BG = practice/reward
Term
once you go back third order before the GPi/SNr, you have what three possible inputs?
Definition
1. direct pathway (putamen)
2. indirect pathway (GPe)
3. STN
Term
injections in the STN to test input & output from M1 --> ?
Definition
saw that the two regions overlapped!
Term
did injections in the putamen to test input & output from M1 --> ?
Definition
saw that the two regions overlapped!
Term
do ventral regions of the putamen get input from M1? does it project to M1?
Definition
do not get input from M1 BUT it projects to M1
Term
what is this an example of: PD patient can barely walk but can pretty much perfectly ride a bike!
Definition
paradoxical akinesia
Term
proposed that ______ is what is allowing a PD patient who can barely walk to be able to ride his bike
Definition
open loop ventral pathway (maybe because it was a lifelong love of his)
Term
where is the human hippocampus located?
Definition
in the temporal lobe
Term
why does the name hippocampus mean "seahorse"?
Definition
because the structure, in humans, looks like a seahorse
Term
does the rat hippocampus look like a seahorse?
Definition
no - it looks more like a banana-shape
Term
4 parts of the rat hippocampal formation
Definition
1. dentate gyrus
2. CA3
3. CA1
4. entorhinal cortex
Term
3 parts of the hippocampus proper
Definition
1. DG: dentate gyrus ("tooth-like bump")
2. hilus (reciprocally connected to DG)
3. CA3/CA1: Cornu Amonis (Ammon's horn)
Term
hippocampal basic circuit: EC (entorhinal cortex) --> via layer III perforant path --> ?
Definition
CA1
Term
hippocampal basic circuit: EC --> via layer II perforant path --> ? (2)
Definition
1. DG
2. CA3
Term
hippocampal basic circuit: DG --> via mossy fibers --> ?
Definition
CA3
Term
hippocampal basic circuit: CA3 --> via Schaffer collaterals --> ?
Definition
CA1
Term
hippocampal basic circuit: CA3 --> via recurrent collaterals --> ?
Definition
CA3
Term
what is one of the largest & most powerful synapses in the brain?
Definition
mossy fiber synapse
Term
3 parts of the hippocampal system
Definition
1. EC: entorhinal cortex
2. hippocampus proper (DG/hilus, CA3, CA1)
3. sub: subiculum
Term
the hippocampus is connected via _____ & ______ pathways to cortical areas & subcortical areas
Definition
EC & fimbria-fornix
Term
is the hippocampus well connected to other brain areas or more isolated?
Definition
very well connected (it is "looking at" what a lot of the brain is doing)
Term
2 input/output pathways for hippocampus
Definition
1. alveus/fimbria/fornix
2. EC
Term
alveus/fimbria/fornix communicates with ______ & _____
Definition
septal nuclei & mammillary bodies
Term
why was it thought that the hippocampus may form new episodic memories?
Definition
anterograde amnesia (HM & others) patients had hippocampal damage
Term
why was it thought that hippocampus may be a cognitive map?
Definition
place cells in rats; spatial attention cells in monkeys
Term
why was it thought that hippocampus may be involved in configural association?
Definition
lesioned rats are impaired on tasks requiring them to recognize cue configurations
Term
HM's bilateral partial hippocampectomy was thought to be responsible for his _____
Definition
anterograde amnesia
Term
the _____ is more easily damaged than other brain areas, e.g. more sensitive to anoxia
Definition
hippocampus
Term
F Vargha-Khadem (1997): people with hippocampal damage (due to anoxic episodes, seizures, or a drug overdose) exhibit ______ but ______
Definition
disabling memory problems (e.g. list recall) but otherwise intellectually "normal" (speech, language, literacy, factual knowledge)
Term
Alzheimer's patients show hippocampal damage & exhibit _____ deficits
Definition
memory
Term
loss of hippocampus --> _____
Definition
memory issues
Term
David Marr's theory of hippocampus
Definition
hippocampus encodes episodes by associating inputs from different sensory modalities into a unified whole
Term
the architecture of hippocampus supports rapid storing of _______
Definition
a day's worth of memories (say 10,000 items)
Term
_____ can perform associative retrieval given a partial memory (e.g. a particular odor might remind us of an incident in our past)
Definition
hippocampus
Term
hippocampus is short-term storage; memories are transferred to _____ during sleep
Definition
cortex
Term
Marr's (1971) hippocampal model
Definition
-binary patterns stored as fixed-point attractors
-circuitry does pattern completion for associative recall
-recurrent connections in CA3 help with completion
-conjunctive subsampling plus threshold gives orthogonalization for better pattern separation
Term
how is spatial working memory tested?
Definition
8-arm radial maze with food cups at each arm end; all food cups are baited at the beginning of each trial; during each trial, rats must remember which arms have already been visited
Term
Naeve et al 1997 spatial working memory task & results
Definition
8-arm radial maze with food cups at each arm end; rats must remember which arms have already been visited because a second arm visit provides no reward --> rats with hippocampal lesions are severely impaired at this task
Term
what task was used to determine whether the hippocampus was involved in sequence learning? (2 versions)
Definition
cups filled with sand have food buried under them; each cup is scented with a unique odor; rat learns a sequence of odors in which he gets presented those cups

experiment 1: rat is given two cups & can dig in one or the other; the cup whose odorant came earlier in the sequence has food, the other does not

experiment 2: instead of a sequential order task, it is a scent recognition task in which one cup has an odor from the sequence & the other cup has a novel odor; the cup with the novel odor has the food
Term
how do hippocampal lesioned rats perform on the sequential learning task?
Definition
barely above chance
Term
how do hippocampal lesioned rats perform on the odor recognition task?
Definition
fine!
Term
what did the sequential odor task demonstrate about the hippocampus?
Definition
there is a specific issue with sequence ordered discrimination in hippocampal lesions!
Term
______ quickly stores episodic memories via a one-shot Hebbian learning rule
Definition
hippocampus
Term
_______ creates more sophisticated memory representations, which requires a gradient-descent learning rule & multiple passes through the training data
Definition
cortex
Term
______ trains _______ by replaying stored memories so that the gradient descent learning rule can do its job
Definition
hippocampus trains cortex
Term
McClelland et al (1995) argues that the transfer from hippocampus --> cortex is necessary for memories because _______
Definition
multiple passes through training data is necessary for real pattern learning
Term
what theory says that hippocampus maintains a representation of space?
Definition
"cognitive map"
Term
O'Keefe = discoverer of ______
Definition
place cells
Term
Moser & Moser = discovered _______
Definition
grid cells
Term
how were place cells discovered in rats by O'Keefe & Dostrovsky (1971)?
Definition
noticed continuous firing fields with gaussian falloff; place fields over the physical space, forming a "cognitive map" of the environment
Term
_______ observed that certain hippocampal cells would fire high activity when the rat was in a specific area in the open field arena (technically called a Muller cylinder)
Definition
O'Keefe
Term
place fields are present instantly in a new environment, but take ______ minutes to fully develop
Definition
10-20
Term
can place cells be controlled by distal visual cues?
Definition
yes
Term
are place cells dependent on visual input?
Definition
no (persist in the dark)
Term
only about _______ of place cells have fields in a typical small environment
Definition
1/3
Term
do place cells have related fields in different environments?
Definition
no
Term
is the topography of physical map in space reflected in topography of place cells?
Definition
no!
Term
what is the shape of place fields in a cylindrical & square arena?
Definition
roughly gaussian
Term
what are the peak firing rats of place fields?
Definition
5-10 Hz
Term
are place fields in a cylindrical & square arena related?
Definition
no
Term
Mittelstaedt & Mittelstaedt (1980): gerbil pup retrieval
Definition
took a pup out of the nest box & moved to another location; in the dark, the mother would wander around until she found him to bring him back; once found, her path back to the nest was nearly a straight beeline back to the nest!
Term
how did Mittelstaedt & Mittelstaedt avoid this compound in their gerbil pup retrieval path integration in rodents task: was there another sensory cue? such as the smell of the nest box or hearing her other pups?
Definition
the authors would move the cup with the mom & the found pup ever so slightly so she would not notice
Term
_______ cells learn & maintain the correspondence between local view representations & path integrator coordinates
Definition
place cells
Term
Redish & Touretsky model of rodent navigation
Definition
local view information (from visual system) --> place cells

local view information --> heat direction information --> path integrator --> place cells

local view information --> head direction information --> local view system
Term
can rats do path integration in the dark?
Definition
only for a certain period of time!
Term
5 places where place cells can be found
Definition
1. EC
2. DG
3. CA3
4. CA1
5. subiculum
Term
2 places where head direction cells can be found
Definition
1. presubiculum
2. retrosplenial cortex
Term
Samsonovich & McNaughton model
Definition
a hippocampal model based on attractor bump theory
Term
attractor bump theory: bump moves around on the sheet by changes in activity on _______
Definition
its edges
Term
Samsonovich & McNaughton's "charts" proposal
Definition
you are born with a certain number of charts in the hippocampus & when you are in a new place, an unused map is "grabbed" to use for that place
Term
can the hippocampus represent multiple maps simultaneously?
Definition
yes (in wiring patterns & can activate one of the maps depending on where the rat is)
Term
why can multiple maps co-exist in an attractor network?
Definition
because activity patterns are sparse, the weight matrix is also sparse
Term
simulation of how multiple maps can co-exist in an attractor network (hippocampus place cell maps)
Definition
in simulation: neurons that are nearby neighbors have strong connections, far away neurons have weak connections; when in a new map, the neurons locations are shuffled, & connection weights are re-distributed

in real life: obviously, they do not move around BUT we can think of connection weights as changing for various maps!
Term
how did Oler & Markus (2000) test task-dependent hippocampal remapping?
Definition
recorded from DG, CA3, & CA1 while animals (rats) ran wither on a figure-8 or plus maze
Term
results of Oler & Markus (2000) task-dependent hippocampal remapping task
Definition
-some cells only had a firing field in task A
-some cells only had a firing field in task B
-some cells fired for a given place (e.g. on the bottom right arm) regardless of task
Term
do all neuron fields remap depending on which task was being performed?
Definition
some but not all
Term
do rats remap place cells immediately upon changing tasks?
Definition
not necessarily (onset may be delayed, rate may be gradual, extend may be partial or complete)
Term
Bostock et al (1991): delayed abrupt complete remapping task & results
Definition
-train in cylinder with white card, then alternate exposure to white & black cards
-most rats did not remap upon first exposure to black card
-but once a rat remapped, it continued to do so
Term
how did Masters & Skaggs test if remapping of hippocampal place cells matters?
Definition
rats would get a brain stim reward at a specific location in the white card task & a different location in the black card task; one rat quickly remapped & learned the task, one never did either, a third rat did not remap until day 11, when it suddenly "got" the task
Term
Johnson & Redish (2007) hippocampal replay experiment
Definition
hippocampus plays sequence forward, from present position to the goal, when animal is paused at a T-intersection & deciding which way to turn
Term
Foster & Wilson (2006) hippocampal replay experiment
Definition
hippocampus replays sequences backwards when animal is paused
Term
can we recover position from firing rate pattern of place cells?
Definition
yes!
Term
what kind of experiments gave a role for hippocampus in spatial cognition?
Definition
hippocampal replay
Term
where did McNaughton place the path integrator in the brain?
Definition
hippocampus
Term
Why did Redish & Touretsky say that the path integrator cannot be in the hippocampus?
Definition
multiple maps make it too hard to update position
Term
Where did Fyhn et al (2004) find the path integrator?
Definition
medial entorhinal cortex ("grid" cells)
Term
Sutherland & Rudy (1988) Morris water maze task
Definition
rats with fornix lesions can still navigate to a visual platform, but they are impaired at learning to find the hidden platform
Term
why is the Morris water maze filled with milky (opaque), cold water
Definition
it prevents rats from being able to see that platform!
Term
eye movements place the image of things that interest us on the _______
Definition
fovea (the part of the retina with the highest activity)
Term
eye movement allow us to track a moving object, to keep the image of that item on the _______
Definition
fovea
Term
if an image appears to a certain site, eye movements called _______ rotate both eyes to that image now falls on the fovea of each
Definition
saccades
Term
if you look (i.e. direct the fovea) from a far object to a near one, _______ eye movements are taking place
Definition
vergence
Term
when an object moves, the image can be kept still on the fovea by mean of a _______ eye movement (e.g. when tracking a moving ball or your finger)
Definition
pursuit
Term
if we move our head, the _______ eye movement is elicited whose function is to keep the image still on the whole retina, not just the fovea
Definition
vestibulo-ocular reflex (VOR)
Term
the _______ is activated when the image of the world slips on a large portion of the retina & produce a sense of self motion (e.g. you sometimes feel like you are moving when sitting in a car that is stopped & a car beside you starts to move)
Definition
optokinetic response (nystagmus)
Term
how many extraocular muscles operate as three agonist/antagonist pairs to move each eye?
Definition
six
Term
lateral/medial recti - _______ movements
Definition
horizontal
Term
superior/inferior recti - _______ movements; small contribution to torsion
Definition
vertical
Term
superior oblique/inferior oblique - _______ & to a smaller extent, vertical movements
Definition
torsion (cyclorotation of the orbit)
Term
lateral rectus: turns _______
Definition
temporally (i.e. away from the nose)
Term
superior rectus: turns _______
Definition
upwards (& intorts a little - towards the nose)
Term
superior oblique: moves eye ________
Definition
intorts (i.e. rotates towards the nose) & moves down a little bit
Term
2 ways we move our eyes
Definition
muscles & motoneurons
Term
innervation differences between lateral rectus & medial rectus
Definition
lateral = abducens

medial = oculomotor
Term
does the abducens nerve cross the midline?
Definition
yes!
Term
which disorder is described by this: patients can look straight ahead fine or in one direction fine, but when the patient wants to look to the other direction (i.e. left) the abducens nerve for the lateral rectus on that side isn't working so that eye won't move (i.e. left eye won't move)
Definition
sixth nerve (abducens) palsy
Term
Edinger-Westphal nucleus
Definition
parasympathetic (pupillary) response of the eye
Term
central caudal nucleus
Definition
controls the eyelid
Term
eye movement control hierarchy
Definition
cerebral centers (frontal eye fields, parieto-occipital eye fields) --> higher order premotor centers (superior colliculus, cerebellar flocculus) --> lower order premotor centers (PPRF, MRF, vestibular nuclei) --> ocular motor nucleus --> ocular motor nerves --> extraocular muscles
Term
Ghandi's three monkeys experiment
Definition
you look at a gaze pattern of a monkey viewing an image of three monkeys; when the dot is stationary, the animal is fixating; when the dot is moving, the animal is making saccades across the image
Term
as the size of an eye movement increases, the _______ of the eye movement increases pretty much linearly
Definition
duration
Term
do both cortical & subcortical regions contribute to the control of saccades?
Definition
yes
Term
______ neurons discharge at a tonic rate during fixation, burst during ipsiversive eye movements, & decrease or cease activity during contraversive eye movements
Definition
abducens
Term
the eye position (during fixation) is directly proportional to the discharge rate of ______ neurons
Definition
abducens
Term
tonic firing of abducens motoneurons --> ?
Definition
holding the eye in that position that corresponds with the level of firing of the active neurons
Term
paramedian pontine reticular formation (PPRF) burst activity scales with the size of _______
Definition
eye movement (increasing horizontal movement --> increases in firing)
Term
______ control the phasic (increases/decreases) in abducens neuron firing associated with eye movements
Definition
excitatory burst neurons
Term
2 parts of the eye circuit that serve as the neural integrators
Definition
nucleus prepositus hypoglossi & medial vestibular complex
Term
______ integrates the information of changes in PPRF firing associated with the movement & uses that information to determine how high or low the firing rate should be for the abducens neurons when they hold their tonic rate associated with a specific fixation point
Definition
neural integrator
Term
where do we find omnipause neurons (OPNs)
Definition
paramedian pontine reticular formation
Term
omnipause neurons
Definition
monosynaptically inhibit EBNs; tonic discharge rate during fixation & cease activity during saccades, functioning in anti-phase with EBNs
Term
_______ "gate" the saccade generation pathway
Definition
omnipause neurons
Term
where does the pulse signal come from to generate saccades (horizontal vs vertical/torsional)?
Definition
horizontal = PPRF

vertical & torsional = riMLF
Term
where does the step signal come from to generate saccades (horizontal vs vertical/torsional) come from?
Definition
horizontal = PPH (nucleus prepositus hypoglossi)

vertical & torsional = INC
Term
visual field sections are mapped onto _______
Definition
visual cortex
Term
does the superior colliculus have topographical organization?
Definition
yes
Term
where is the superior colliculus located?
Definition
top of the brainstem
Term
superior colliculus has things close to the _______ more highly represented
Definition
fovea
Term
the SC is a laminar structure separated functionally into _______, _______, & ______ layers
Definition
superficial, intermediate, & deep
Term
difference between what excites superficial vs. intermediate/deep layers of the superior colliculus
Definition
superficial = respond to the presentation of a visual target

intermediate/deep = elicit motor with or without a visual response
Term
is the sensory response to SC limited to visual stimuli?
Definition
no
Term
is the motor output of SC limited to saccades?
Definition
no
Term
how does SC neuron activity change throughout a task in which a visual cue precedes a motor output saccade?
Definition
increase in activity slightly after target presentation ("visual" burst) --> drops down to baseline --> big increase in activity at saccade ("motor" burst)
Term
how does FEF neuron activity change throughout a task in which a visual cue precedes a motor output saccade?
Definition
increase in activity slightly after target presentation ("visual" burst) --> remains at increased firing level --> further slight increase at saccade ("motor" burst)
Term
3 types of activity neurons in SC
Definition
1. both strong visual & saccade-related responses
2. weak visual response but high saccade-related responses
3. bursts at neither but change activity in between visual & saccade times
Term
each neuron in the ______ layers of the SC discharges during saccades of a restricted amplitude & direction
Definition
intermediate
Term
how does the topographic map of movement fields in the intermediate & deep layers of the SC coincide with the visual response fields of the superficial layers?
Definition
overlap quite nicely
Term
will a given SC neuron fire for saccades of many different amplitudes & directions?
Definition
yes!!
Term
who expressed deep skepticism regarding the idea that memory traces (engrams) were in a particular region of the cerebral cortex?
Definition
Karl Lashley
Term
two principles proposed by Karl Lashley regarding where memories are stored
Definition
Equipotentiality: different regions contributed equivalently to the storage of a memory trace - because damage to any region causes a loss

mass action: memories depend on the collective action of numerous regions - because the greater the cortical territory destroyed, the graver the loss
Term
Lashley's task to study memory engrams
Definition
training rats to run a maze & then kept track of how many errors were made in the maze & how that related to cortical lesions
Term
2 results following HM's bilateral hippocampectomy
Definition
1. reduction in seizure frequency
2. almost complete loss of recent memory
Term
hippocampus is part of the _______ cortex
Definition
limbic
Term
one way hippocampus is not cortical & one way that it is
Definition
-it is not cortical in the sense that it is not 6-layered

-topographically it is part of the cortical gray matter sheet at its edge
Term
______ lies next to the entorhinal cortex which lies next to the perirhinal cortex
Definition
hippocampus
Term
3 steps through which hippocampus communicates with subcortical structures
Definition
1. via a bundle of fibers that go under corpus callosum
2. up to basal forebrain
3. hits the septum
Term
hippocampus communicates with the rest of the cortex via fibers that emanate from/travel to the _______
Definition
entorhinal cortex
Term
what type of amnesia did HM have?
Definition
anterograde (could not remember new information)
Term
what type of things could HM have memories for?
Definition
distinct events that happened prior to his operation (especially good for distant memories)
Term
anterograde amnesia
Definition
the inability to form new memories despite the ability to remember old ones
Term
Jay McClelland et al consolidation theory
Definition
episodic memories are stored first in the hippocampus in the form of a sparse & temporary representation & gradually transferred to the neocortex in the form of a distributed & permanent representation
Term
despite being sparse, hippocampus would still eventually fill up --> therefore, memories eventually move out from the hippocampus to the _______ in order that they first came in
Definition
neocortex
Term
in the neocortex, stored knowledge is represented in a _______ fashion
Definition
distributed
Term
as the _______ process takes place with information being gradually transferred to neocortex, the information that is transferred is no longer held in the hippocampus to allow it to have space for new "snapshots"
Definition
consolidation
Term
2 times when memory consolidation may occur
Definition
1. during sleep
2. constant rehearsal during the day
Term
_______ is there to provide a medium for the initial storage of memories in a form that avoids interference with the knowledge already acquired in the neocortical system
Definition
hippocampus
Term
why does incorporation from hippocampus to neocortex take a long time?
Definition
to allow new knowledge to be interleaved with ongoing exposure to exemplars of the existing knowledge structure, so that eventually the new knowledge may be incorporated into the structured system already contained in the neocortex
Term
who touted this theory: episodic memories reside permanently in the hippocampus & the gradual transfer of information from the hippocampus to the neocortex gives rise to a neocortical representation underlying the semantic knowledge rather than episodic memory
Definition
Mascovich
Term
transformation theory
Definition
memories are "consolidated" in neocortex are fundamentally different from those that were initially dependent on the hippocampus
Term
Wincour et al interpretation of HM's description of events from the past are impoverished in details even for events that occurred early in life
Definition
recall is not truly episodic
Term
what could imply the existence of an impairment extending beyond memory in amnesics affecting the ability to imagine events in context
Definition
even when describing imagined events, amnesiacs provide accounts relatively impoverished in internal detail
Term
hippocampus may play a role not only in taking snapshots of events in contexts ("episodes") BUT also in ________
Definition
imagining them!
Term
results of this experiment: asking HM to identify celebrities who became famous either before or after his surgery (before 1950s, during 1950s, or after 1950s)?
Definition
he did as well as controls in recognizing famous people from before 1950s --> was slightly impaired for celebrities during 1950s --> was even more impaired for celebrities that became famous after 1950s
Term
semantic knowledge is stored in _______
Definition
neocortex!
Term
evidence that hippocampal lesions may cause an impairment of semantic learning
Definition
HM could not learn the meanings of words new to him
Term
task that demonstrated that HM could not learn the meanings of words new to him
Definition
perform a series of trials where individuals would either have to define word, identify a synonym, use it in a sentence etc. --> HM simply never mastered the words
Term
what task demonstrated that amnesics possess a capacity for active rehearsing using a limited-capacity buffer
Definition
after viewing 10 words in succession over the course of half a minute, amnesic patients are as good as normal controls at recalling the final two words but their performance plummets for earlier words
Term
what would happen with amnesics if you interrupt word-recall with an intermediate task?
Definition
they are not be good at the most recent words anymore
Term
in a delayed match-to-sample task with a variable delay between the sample & the probes, how did HM do?
Definition
can retain categorical easily rehearsed information (trigram identity) over an indefinitely long delay in the absence of distraction but quickly loses difficult-to-rehearse parametric information (ellipse aspect ratio)
Term
what can we take away from the fact that HM can do fine on easily rehearsed categorical working memory information but quickly loses difficult-to-rehearse information?
Definition
active rehearsing using working memory can be used if that capacity was not stretched
Term
example experiment that showed hippocampal activity fires both when experiencing something & when recalling it?
Definition
patient's hippocampal activity fired much more highly when they saw the Simpson's --> when talking about the Simpson's, neural activity surged again!
Term
what demonstrated that classical conditioning does not depend on the hippocampus?
Definition
amnesics exhibit relatively normal Pavlovian conditioning
Term
task that demonstrated that amnesics exhibit relatively normal Pavlovian conditioning
Definition
patient would sit in front of a speaker/monitor & would hear either tones of high/low frequency & see either red/blue squares on a monitor; the subjects would also be next to an extremely loud boat horn; the subjects would also be recorded from skin electrodes for stress responses (sweating AKA skin conductance)
Term
can amnesics remember details about tasks?
Definition
no (even when they can perform the tasks fine)
Term
conclusion from the study that amnesics can do classical conditioning but cannot remember the task when asked details about it?
Definition
amnesics are capable of a form of learning that does not require episodic memory
Term
results from this task: individuals are shown very incomplete pictures (basically dot fragments) that are gradually filled in until they are the complete image (ex. an airplane); then, after waiting some time, you present reduced images & ask the person what they represent
Definition
there is a carry-over from previous sessions that allow the individuals to identify reduced picture even though they could not initially do so
Term
can HM do priming tasks?
Definition
yes (although he did not remember having taken the test before)
Term
how does HM do with mirror-drawing skill?
Definition
relatively normal acquisition & retention
Term
if you ask HM (or other amnesics) about details about the mirror-drawing task --> ?
Definition
they are poor at remembering details about the sessions during which they acquired the skill
Term
what task allowed this conclusion to be made: even skills altogether lacking a motor component appear not to depend on the hippocampus
Definition
amnesics show relatively normal acquisition & retention of the skill of mirror-drawing
Term
probabilistic learning
Definition
trying to figure out the "rules" of the environment
Term
what allowed this conclusion to be made: acquisition of knowledge about the relation between cues & outcomes when the relations are probabilistic appears not to depend on the hippocampus
Definition
amnesics show relatively normal statistical learning in the weather prediction task
Term
weather prediction task
Definition
you are shown cards with various designs on them & learn how the designs to various % chance of sun or rain; you are asked to predict sun or rain when a given card was shown
Term
what happens when you ask amnesics for details about a task they just performed (e.g. the weather prediction task)?
Definition
they cannot do this
Term
facts (semantic knowledge) & events (episodic memory) are dependent on the extended _______ system in the sense that damage to the system impairs newer (anterograde) acquisition in these domains
Definition
hippocampal
Term
Rizzolatti et al important motor areas
Definition
M1, SMA, PMd, PMv, central cingulate motor area, rostral cingulate motor area, pre-SMA, FEF, SEF
Term
M1 is about equal to Brodmann's area ______
Definition
4
Term
SMA + PMd + PMv + pre-SMA + SEF are about equal to Brodmann's area ______
Definition
6
Term
FEF is about equal to Brodmann's area ______
Definition
2
Term
how does motor cortex in humans relate to monkeys?
Definition
it is roughly the same
Term
are connecting motor pathways organized topographically?
Definition
yes
Term
M1 plays a critical role in _______
Definition
voluntary control of movement
Term
who did electrical stimulation on awake patients in various parts of the cerebral cortex & could determine the behavioral effects?
Definition
Penfield & Rasmussen
Term
Penfield's stimulation of M1 --> ?
Definition
extension of toes, movement of foot, movement of knee, & movement of shoulder
Term
______ published the classic homunculus as we know it today showing the different parts of the body represented by the motor cortex & how much cortex is dedicated to different parts
Definition
Penfield
Term
what type of experiment allowed for relation of activity in one small region of area M1 to activation of muscles as measured by EMG in monkey
Definition
spike-triggered averaging - recording electrical activation of individual muscles following action potentials fired by a cortical neuron - has revealed that even a single neuron's activity is correlated with contractions of multiple muscles
Term
do neurons influence a single muscle in a one-to-one ratio?
Definition
no (but sometimes can innervate related muscles)
Term
Lemon spike-triggered averaging experiment
Definition
activated an area of neurons as opposed to single neuron --> led to greater responses of the movement associated with the original neuron & then a few more movements come online
Term
what happens when a train of stimulation is given to M1 & activity in EMG muscles are recorded?
Definition
do not get nice spikes in EMG but tetanic wider plateau-like read outs
Term
how is a given digit represented in motor cortex?
Definition
represented several times over in different parts of motor cortex
Term
Georgeopolous
Definition
did a series of experiments to determine the "population vector" coding theory of motor neuron direction preference/contributions to movement
Term
how does neural firing rate allow for population vectors to be made for movement coding from M1?
Definition
neurons fire at a maximal rate for its "preferred" direction & slope off to either size in a bell-shaped curve
Term
does this study tell us whether neural activity really means movement direction or muscle direction: Georgeopoulos using vector averaging for estimating the direction of intended movement from population coding of multiple M1 neurons?
Definition
no
Term
Strick et al task to disentangle direction of movement vs muscle flexion/extension
Definition
monkey moves a lever up, down, right, left, etc. with the hand in different positions (prone [palm down], supine [palm up], or mid)
Term
did Strick et al task show that individual muscles are affected by extrinsic directions?
Definition
yes
Term
conclusion from these results: the firing of some neurons in M1 encodes extrinsic direction of motion (right, up, left, down) rather than anatomical direction (extension, flexion, ulna, radial) when these are dissociated by having monkey move a lever with hand prone or supine
Definition
although it is possible to activate certain muscles by activating certain neurons in the cortex, it is not really the case that it is a one-to-one ratio for neurons to muscles
Term
in the neurons where neuron preferred direction matches preferred muscle direction --> it is probably involved in _______ of movement
Definition
initiation
Term
in the neurons where neuron preferred direction is opposite preferred muscle direction --> it is probably involved in _______ movement
Definition
braking
Term
in the neurons where neuron preferred direction is orthogonal to preferred muscle direction --> it is probably involved in _______
Definition
stabilizing
Term
where is the supplementary motor area (SMA)
Definition
medial face of cerebral hemisphere
Term
how was SMA first identified?
Definition
electrical stimulation studies in various species (including humans)
Term
who discovered the role of SMA stimulation from one of their patients?
Definition
Penfield & Rasmussen
Term
is there a representation of the body in the SMA?
Definition
yes
Term
how do lesions of human SMA compare to lesions of M1?
Definition
much more subtle effect
Term
2 tests that showed what SMA lesions can lead to
Definition
1. patients have trouble making alternate movements of two hands (e.g. clenching hands in alternation)
2. serial movements (e.g. touching your fingers in a sequence with your thumb)
Term
what is the result of SMA lesions?
Definition
the ability to make complex patterns of movement (not gross motor impairments)
Term
single neurons in monkey SMA give clear signs of involvement in ________
Definition
bimanual coordination
Term
Tanji et al study on SMA & sequence learning showed --> ?
Definition
neurons in SMA fired differently depending on the sequence of movements (not necessarily the movement itself) & some neurons would fire between two specific movements, regardless of the rest of the sequence
Term
What did Strick et al show us about PMv?
Definition
neurons are sensitive to movement direction as described by an extrinsic system (not the intrinsic muscle-like pattern)
Term
some neurons in area _______ have visual receptive fields defined relative to the head, like neurons in parietal area VIP
Definition
PMv
Term
as the eyes move, the receptive fields move on the ______ to stay fixed relative to the position of the head
Definition
retina
Term
result of this study: recorded from PMv neuron that had a receptive field on right side of monkey's face & responded to visual stimuli moving towards right side of monkey's face --> would show stimulus 1 (towards right side of face) or stimulus 2 (towards left side of face)
Definition
neuron would respond to stimulus 1 (moving towards right side of face) regardless of where it was looking & would not respond to stimulus 2 (moving towards left side of face) regardless of where the monkey was looking
Term
result of this study: monkey faces screen with multiple reach targets; on each trial, he fixated either on a right dot or a left dot; when looking at a dot, he could reach for one of four targets around the dot (so the middle target overlapped); recorded in PMv
Definition
neural firing in PMv (with associations to MIP) would be strongest for a reach for a particular direction (regardless of where it was on the screen); neuron was sensitive for reach direction in retinal coordinates rather than body coordinates
Term
result of this experiment: recording from PMv; had monkeys look at and/or grasp various objects
Definition
neurons fired strongly when grasping an object in the light, grasping it in the dark, or just looking at it in th elight
Term
who originally described mirror cells in PMv?
Definition
Rizzolatti et al
Term
mirror cells
Definition
neurons that fire during a given grasping action by the monkey or when the monkey sees someone else perform that action - may embody "motor semantics"
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