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functional roles of the basal ganglia
initiation, modulation, and termination of output to motor cortex
eye movements, learning of routine behaviors (habits), reward associated behaviors, may play a role in complex thought and reasoning
basal ganglia
important component of the motor system, essential for learning and execution of automatic action
caudate nucleus, putamen, globus pallidus
corticostriatal pathway
collective term for all cortical projections to the striatum
cerebral cortex —> striatum
input zone
striatum: caudate, putamen
striatum
gets its name because of the striped appearance from white matter fibers running between the caudate and putamen
medium spiny neurons
90% of neurons in striatum
divergence of inputs from corticostriatal axons allows a single one to integrate the influence of many cortical cells
output zone
globus pallidus externus and globus pallidus internus/substantia nigra pars reticulata
globus pallidus externus
tonically active GABAergic neurons that project to the subthalamic nucleus
tonically active
neurons fire constantly until inhibited
globus pallidus internus/substantia nigra pars reticulata
tonically active GABAergic neurons that project to the thalamus
subthalamic nucleus
located ventral to the thalamus
a glutamatergic nucleus receiving projections from the GPe and projects to the GPi/SNr
glutamatergic nucleus
contains cell bodies of glutamatergic neurons
substantia nigra
a midbrain structure
parkinson’s disease
substantia nigra pars compacta (SNc)
dopaminergic cells closely compacted together; modulate striatum
substantia nigra pars reticulata (SNr)
GABAergic cells (tonically active) that are the other major output zone of the basal ganglia
direct pathway
allows for initiation of movement when cortical activation of striatal medium spiny neurons that project to GPi/SNpr is sufficient to disinhibit the VA/VL complex of the thalamus
cerebral cortex —> striatum —> globus pallidus internus —> thalamus —> frontal cortex
excitatory
indirect pathway
antagonizes the activity of the direct pathway, resulting in termination of movement by increasing activity of the GPi/SNpr and enhancing the inhibition of the thalamus
cerebral cortex —> striatum —> globus pallidus externus —> subthalamic nucleus —> globus pallidus internus —> thalamus —> frontal cortex
inhibitory
disinhibition
inhibition of an inhibitory neuron; results in removal of inhibition and an increase in the degree of excitation
excitatory D1 dopamine receptors
when dopamine is released from the SNc, there is an excitation of these striatal neurons
inhibitory D2 dopamine receptors
when dopamine is released from the SNc, there is an inhibition of these striatal neurons
hyperdirect pathway
inhibitory (emergency brake)
parkinson’s disease
loss of dopaminergic neurons in substantia nigra
resting tremor
huntington’s disease
degeneration of GABA-ergic neurons in the caudate nucleus and putamen affecting external globus pallidus
hemiballismus
basal ganglia syndrome resulting from the damage to the subthalamic nucleus
typically restricted to one side
hypokinetic movement disorder
inability to initiate movements at a normal rate of speed
akinesia
no movement
bradykinesia
slow movements
rigidity
increased muscle tone
dyskinesia
involuntary movement
chorea
to dance
ballism
wild flailing
learning
process by which experiences change our nervous system and hence our behavior
it allows us to respond to behavioral changes and act appropriately (appropriate behavior in appropriate situation)
durable and usually adaptive change in an animal behavior traceable to a specific experience in the individual’s life
memories
these long-term changes in the nervous system following learning
experiences
change the way we percieve, perform, think, plan and behave by physically changing the structure of appropriate parts of the nervous system
encoding
learning produces changes in the nervous system by encoding the new information
consolidation
strengthening changes associated with the initial information that is learned
storage
after consolidation, the memory is stored and maintained via these persistent changes in the nervous system
retrieval
process of accessing and using the information stored by the neural changes
part of the learning process
perceptual learning
learning to recognize a particular stimulus (new stimulus, changes/variation in familiar stimulus)
primary function is the ability to identify and categorize objects and situations (learning to recognize things, not what to do when they are present)
all sensory systems are capable
involves changes in the appropriate sensory association cortex
stimulus-response learning
learning to automatically make a particular response in the presence of a particular stimulus
involves the establishment of connections between circuits involved in perception and those involved in movement
the response behavior can be a simple automatic reflex or a complicated sequence of movements learned previously
ex. classical and instrumental conditioning
constant start position
motor learning
learning to a produce a behavior
modifications to motor circuit
how we respond to environment; not independent of perceptual learning
different response to different stimuli
relational learning
learning how stimuli are related
ex. complex sensory associations, spatial learning, episodic learning
organized in time
variable start position
classical conditioning
a learning procedure in which a neutral stimulus becomes associated with a naturally occurring stimulus to elicit a conditioned response
ex. Pavlov’s dog
unconditional stimulus (US)
when a stimulus that initially produces no particular response is followed several times by a repeated stimulus
unconditional response (UR)
produces a defensive or appetitive response
conditioned stimulus (CS)
the first stimulus
conditioned response (CR)
when a conditional stimulus itself evokes the response
eyeblink conditioning paradigm
training: tone + puff of air in eye
end result: tone produces eyeblink
Hebb rule
the hypothesis proposed by Donald Hebb in 1949 that the cellular basis of learning involves strengthening of a synapse that is repeatedly active when the postsynaptic neuron fires
limbic system
set of interconnected brain structures whose primary functions are: learning, memory, motivation, and emotion
it is a FUNCTIONAL system, with structures located both in the telencephalon and diencephalon
limbic cortex, hippocampus, amygdala, hypothalamus
amygdala
involved in emotional experiences and responses
hippocampus
involved in some forms of learning and memory
hypothalamus
critically important for normal physiological functioning
classically conditioned emotional response
amygdala plays important part
established by pairing a neutral stimulus (tone) with an unconditioned stimulus
the tone becomes conditioned stimulus and response to it becomes conditioned response
lateral nucleus
neurons transmitting auditory information
neurons transmitting somatosensory information
central nucleus
where weak and strong synapses on neurons project to
conditioning
animal is learning to respond
extinction
animal is learning not to respond
dietary specialist
concentrates exclusively on one or very few safe foods are unable to acquire taste aversion after consumption of an unusual-tasting food
selection
favors costly investements in learning only when there is environmental unpredictability that has reproductive relevance for individuals
insectivorous bats
ability to distinguish a taste from a different effect
instrumental conditioning
a learning procedure whereby the effects of a particular behavior in a particular situation increase (reinforce) or decrease (punish) the probability of the behavior (also called operant conditioning)
reinforcing stimulus
an appetitive stimulus that follows a particular behavior and thus makes the behavior become more frequent
punishing stimulus
an aversive stimulus that follows a particular behavior and thus makes the behavior become less frequent
functions of reinforcement system
to detect reinforcing stimuli
to strengthen the connections between the neurons that detect the discriminative stimulus (perception) and the neurons that produce the instrumental response
ventral tegmental area (VTA)
group of dopaminergic neurons in the ventral midbrain whose axons form the mesolimbic and mesocortical systems and are important in reinforcement
nucleus accumbens
nucleus of the basal forebrain near the septum; receives dopamine from neurons of the VTA and is thought to be involved in reinforcement and attention
two major pathways between the senosry and the motor association cotex
direct transcortical pathway
connection via basal ganglia and thalamus
both involved in instrumental conditoning
transcortical pathway
acquisition of complex behavior:
following instructions/manual (humans)
observing and imitating behavior (animals and humans)
after several repetitions basal ganglia learn the process and take over most of it
hippocampal formation
a forebrain structure of the temporal lobe, constituting an important part of the limbic system; includes the hippocampus proper (Ammon’s horm), dentate gyrus, and subiculum
entorhinal cortex
a region of the limbic cortex that provides the major source of input to the hippocampal formation
dentate gyrus
part of the hippocampal formation; recieves inputs from the entorhinal cortex and projects to the field CA3 of the hippocampus
granule cells
small neurons located in the dentate gyrus; they send axons to the field CA3 of the hippocampus
perforant path
the system of axons that travel from cells in the entorhinal cortex to the dentate gyrus of the hippocampal formation
field CA1
part of the hippocampus; recieves inputs from field CA3 and projects to the hippocampal formation via the subiculum
field CA3
part of the hippocampus; recieves input from the dentate gyrus and projects to the field CA1
pyramidal cells
a category of large neurons with a pyramid shape; found in the cerebral cortex and Ammon’s horn of the hippocampal formation
population EPSP
an evoked potential that represents the EPSPs (excitatory postsyntaptic potentials) of a population of neurons
long-term potentiation (LTP)
a long-term increase in the excitability of a neuron to a particular synaptic input caused by repeated high-frequency activity
how LTP is induced
by stimulating the axons in the perforant path with a burst of approximately 100 pulses of electrical stimulation, delivered within few seconds
the size of the first population EPSP
indicates the strength of the synaptic connection before LTP has taken place
evidence for LTP
is obtained by periodically delivering single pulses to the perforant path nd recording the increased responses in the dentate gyrus
record from cells within the dentate gyrus
subsequent perforant pathway stimulation results in progressive increases in population EPSP’s
how a synapse will be strengthened
pre and postsynaptic firing occurs at about the same time
one stimulus
non-associative LTP
multi-stimuli
associative LTP
associative LTP
a long-term potentiation in which concurrent stimulation of weak and strong synapses to a given neuron strengthens the weak ones
follows the Hebb rule
LTP - bottom line
requires 2 things: activation of synapses and depolarization of postsynaptic neuron
synaptic plasticity
ability of the synapse to change in strength in response to either use or disuse of transmission over synaptic pathways
usually associated with learning
long-term depression
long-term decrease in the excitability of a neuron to a particular synaptic input caused by stimulation of the terminal button while the postsynaptic membrane is hyperpolarized or only slightly depolarized
reverse process to LTP
synapses keep decreasing
sensory memory
a brief period of time during which the sensation of environment stimuli is initially remembered
length ranges from fractions of a second (most senses) to a few seconds (echoic memory)
occurs in each of the senses
echoic memory
answer a question without hearing it again
short-term memory (STM)
contains meaningful or salient enough information transferred from sensory memory
limited capacity
brief duration (seconds to minutes)
can be lost without rehearsal
long-term memory (LTM)
contains information from short-term memory that is consolidated
relatively permanent storage (minutes, hours, days, decades)
strengthened with increased retrieval
unlimited capacity
unlimited capacity
can remember things from 100+ years ago
working memory
is related to short-term memory, but it lasts slightly longer and is involved in the manipulation of information
is equal to STM
chunking
combining a few elements into one element
used in everyday life
consolidation
process by which rehearsal of information in STM results in transfer to LTM
amnesia
refers to a failure to remember
can be temporary or permanent
3 types
anterograde amnesia
difficulty in forming new memories for events that occur after a brain trauma
cannot later remember events that occur after brain damage