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Neural circuits. Lecture 3. Cellular neuroscience. Nerve cells with ion channels and synapses How do neurons interact? How is activity patterned? How is appropriate activity selected? How is sensory input used? How is motor output coordinated and generated?. Why Crayfish?.
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Neural circuits Lecture 3
Cellular neuroscience • Nerve cells with ion channels and synapses • How do neurons interact? • How is activity patterned? • How is appropriate activity selected? • How is sensory input used? • How is motor output coordinated and generated?
Why Crayfish? • Why escape behaviour? • Simple behaviour • Short duration startle response • simple nervous system • Abdominal ganglia with about 400 neurons
2 Escape behaviours • Anterior tap • Goes back • All segments bend • Tail tap • Goes up • Segments 1-3 bend • Differences in physiology match differences in adaptive behaviour
Abdominal tap • Ventral nerve cord • Contains lateral giant • LG Stimulated by tap • LG • Causes motoneurons • Then muscles to be active
Tactile hairs activated by water movement Sense organs
Excite Sensory interneuron Direct path (a) Bi-synaptic path (b) Multiplicity – lowers threshold But with safety factor Sense organs
Abdominal ganglia MG Transverse section LG neurite neuropil somata
LG to MoG • Electrical synapse
LG Motoneuron path • Indirect • Chemical • Motoneuron filed with procion yellow
LG →SG →MN • Segmentalgiant • Prevents LG MG interaction by rectifying electrical synapse between LG and SG • SG provides chemical excitation of flexor motor neurons • SG acts as amplifier
FF Motoneuron • 9 Fast Flexor motor neurons • Individually identifiable • All excited by LG via SG • Rectifying synapse • MG and LG separated
Summary so far • Excitatory pathwaysense cell to musclecontraction
Preventing second escape • Turn off hair cell afferents • CDI neurons produce delay and postsynaptic inhibition of the SI
Preventing second escape • Turn off hair cell afferents • CDI neurons produce delay and postsynaptic inhibition of the SI • CDI neurons produce delay and also presynaptic inhibition of the receptors
Inhibition of Posture • MRO normally excites extensor motoneuron and flexor inhibitor • MRO turned off twice • Accessory cell • Fast extensor
End of escape • Inhibition of the flexion system LG spike FI FFMN
Major features of net • Need sensory coincidence to fire LG • Ensures safety if single cell accidentally fires • Lowers behavioural threshold below single neuron threshold (law of averages) • Fast • Multiple, parallel pathways • Combination of electrical feed-forward and chemical excitation • Chemical allows amplification of signal • Chemical allows modulation of pathway
Other systems • Locust & Drosophila jump • Cockroach running • Fish C-start
Photoactivation of GF • Flies cannot see http://www.sciencedirect.com/science/MiamiMultiMediaURL/B6WSN-4FWM4P4-J/B6WSN-4FWM4P4-J-4/7051/d542b7199c07d3f274131cb29e173241/Movie_S2..mov
Cockroach • Arthropod – escapes from toads, etc • Responds to air movement
Air movement hairs give directionality Escape correct way! Giant fibres Cockroach
Teleost fish • Mauthner cell • Large hindbrain, descending cell • Responds acoustically
Feed forward pathway • Receptor – interneuron or • Receptor – Mauthner ?
C-start startle response • But note Mauthner cell only used in some fast starts, • other homologous cells exist in other neuromeres
Conclusions • Apparently simple behaviour has complex neural circuit • Giant fibers for fast response • Feed-forward pathways • Safety features so only escape when needed • Chemical systems • Amplification • Modulation • Inhibition