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PART 3: MOTOR STRATEGIES #15: ESCAPE BEHAVIOR IN CRAYFISH. CH7: escape behavior in crayfish behavior features & functional anatomy neuronal architecture adaptive modulation summary: chapter 7. BEHAVIOR & FUNCTIONAL ANATOMY. walking is normal mode of locomotion - PowerPoint PPT Presentation
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CH7: escape behavior in crayfish
behavior features & functional anatomy
neuronal architecture
adaptive modulation
summary: chapter 7
PART 3: MOTOR STRATEGIES#15: ESCAPE BEHAVIOR IN CRAYFISH
walking is normal mode of locomotion integrated motor escape response tail flip tail propulsion using flexor & extensor muscles
BEHAVIOR & FUNCTIONAL ANATOMY
nongiant slower
medial giant: anterior stimulus move back rapid
lateral giant: tail stimulus move up & back rapid
3 types of tail flip response
BEHAVIOR & FUNCTIONAL ANATOMY
tail flip can be elicited by electrical stimulus tactile stimulus
responses are comparable triggers initiate complex motor sequences
BEHAVIOR & FUNCTIONAL ANATOMY
typical invertebrate CNS plan (ganglia + connectives) brain SOG complex 5 thoracic ganglia 6 abdominal ganglia... contain tail flip circuitry
ganglia communicate & are coordinated via connectives peripheral comm. via roots
1: swimmerets 2: extensors 3: flexors (motor only)
NEURONAL ARCHITECTURE
2 pairs of prominent giant axons lateral giant interneurons (LGI)
cell bodies & dendrites in each abd. segment electrical synapses (septate / segmental) axons project next segment lateral giant escape
medial giant intern. (MGI) cell bodies & dendrites in brain ~ single fast neuron medial giant escape
NEURONAL ARCHITECTURE
giant interneurons motor giant neurons (MoGs) MoGs flexor muscles sensory input to:
head MGI
all MoGs tail LGI
1-3 MoGs focus on LGls
NEURONAL ARCHITECTURE
LGI tail flip circuitry sensory input: ~1000 hairs with sensory neurons sensory interneurons: LGIs & brain
A: phasic C: tonic
LGIs
NEURONAL ARCHITECTURE
LGI tail flip circuitry sensory input: ~1000 hairs with sensory neurons sensory interneurons: LGIs & brain
A: phasic C: tonic
LGIs MoGs
NEURONAL ARCHITECTURE
LGI tail flip circuitry sensory input: ~1000 hairs with sensory neurons sensory interneurons: LGIs & brain
A: phasic C: tonic
LGIs MoGs flexor muscles:
5 / segment + other input
NEURONAL ARCHITECTURE
chemical synapses (slow) at input & output electrical synapses (fast) elsewhere sensory LGI
directly () short latency indirectly () long latency
NEURONAL ARCHITECTURE
chemical synapses (slow) at input & output electrical synapses (fast) elsewhere sensory LGI
directly () short latency indirectly () long latency
sensory influence fast flexor motor neurons LGI MoGs & segmental giant (SG)... very fast !
NEURONAL ARCHITECTURE
LGIs SG (electrical)
SGs fast flexor motor neurons (electrical)
NEURONAL ARCHITECTURE
LGI neurons at center of circuit
convergence of sensory input LGI
divergence of LGI output motor
NEURONAL ARCHITECTURE
3 components of “flipping out” behavior
rapid flexion of abdomen
re-extension of abdomen
swimming
independent behavior modules
NEURONAL ARCHITECTURE
LGIs only involved in flexion
2 abdominal sensory input channels
biphasic LGI spike (EPSP)
indirect chemical
direct electrical
NEURONAL ARCHITECTURE
rapid flexion response to abrupt tail stimulus because sensory - interneuron chemical synapses depress
with prolonged stimuli electrical synapses LGI
have high threshold & short
time constants sensory input presynaptic
LGI inhibition
NEURONAL ARCHITECTURE
2 pathways from LGI (elect) MoG (chem) flexor muscles SG (elect) FFs (chem) flexor muscles
FFs threshold below that of signal from SG... no delay in signal
NEURONAL ARCHITECTURE
LGI fast speed from large diameter axons electrical synapses
LGI sufficient & necessary for tail flip response ?
NEURONAL ARCHITECTURE
necessary: sever MoG* stimulate tail flip hyperpolarize LGI measure severed MoG output
LGI sufficient & necessary for tail flip response...
“command neurons”
sufficient: inject current tail flip
NEURONAL ARCHITECTURE
LGI makes all-or-nothing decision to escape ? what about upstream sensory decision ? ... graded, not all-or-none synaptic input together... explains why there is no partial tail flip
NEURONAL ARCHITECTURE
no single LGI satisfied criteria they are in series, linked abdominal segments act as functional unit
command neuron firing or stimulation elicits complex behavior... eg, coordinated / rhythmic appendage movement
criteria: neuron should demonstrate activity necessary & sufficient to elicit behavior normal response to sensory stimulus normal pattern of activitation
NEURONAL ARCHITECTURE
LGI inhibitory signals: “command-derived inhibition” ensures that additional flexor responses do not occur
NEURONAL ARCHITECTURE
LGI inhibitory signals: “command-derived inhibition” ensures that additional flexor responses do not occur
LGI spikes inhibit further LGI & MGI spikes sensory, LGIs, MoGs & muscles inhibited
NEURONAL ARCHITECTURE
further inhibition of
extension
slow flexor and slow extensor systems
widespread inhibitory influence
critical timing (details... )
every level of tail flip circuitry
NEURONAL ARCHITECTURE
read and be sure you understand text sections on
re-extension
swimming
problems... journal questions
NEURONAL ARCHITECTURE
other influences on tail flip responses ?
does not always work
modulated by
restraint-induced inhibition
motivation (feeding)
learning
ADAPTIVE MODULATION
blocked by nerve cord transection
decreased facilitation of reflex
increased inhibition at higher
levels
voluntary tail flip remains
restraint-induced inhibition
ADAPTIVE MODULATION
cut nerve cord
abolishes feeding-
induced increase
must be eating to
influence response
motivational modulation of escape behavior
feeding raises threshold of tail flip response
ADAPTIVE MODULATION
feeding modulates LGI
firing only
degree of inhibition
relative to stimulus
“competition”
ADAPTIVE MODULATION
modulation of escape behavior by learning repetition... what is important & what is not habituation: reduced response with repeated stimuli self-induced habituation by water movement ? prevented by command-derived inhibition
ADAPTIVE MODULATION
anterior tactile stimulus tail flip response mediated by lateral giant interneurons (LGI) sensory hair inputs LGIs sufficient & necessary for response
widespread activation of flexor system command neurons, trigger escape response command-derived inhibition, cancels competing
response, enables subsequent elements
SUMMARY
command-derived inhibition, cancels competing response, enables subsequent elements
reextension from sensory feedback (reafference), via stretch receptors (muscle receptors, MROs) & sensory hairs on tailfan
swimming from central pattern generator activated by sensory input with prolonged delay
modulated by various influences... restraint, feeding, learning
SUMMARY
NO CLASS on T.3.20
SECTION 3 REVIEW on R.3.22
2nd MIDTERM EXAM:
written, 15% of final grade
ASSIGNED (web page) @ 6 pm T.3.27
DUE (eMail) @ 3 pm R.3.29
NEUROBIOLOGY CALENDAR