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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 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 Neural response Neural circuit – anatomy Neural circuit - schematic Sense organs  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 Abdominal ganglia Transverse section LG MG neurite somata neuropil LG to MoG  Electrical synapse LG  Motoneuron path  Indirect  Chemical  Motoneuron filed with procion yellow LG → SG → MN  Segmental giant LG  MG interaction by rectifying electrical synapse between LG and SG  SG provides chemical excitation of flexor motor neurons  SG acts as amplifier  Prevents FF Motoneuron  9 Fast Flexor motor neurons  Individually identifiable  All excited by LG via SG  Rectifying synapse  MG and LG separated LG & motoneurons Summary so far  Excitatory pathway sense cell to muscle contraction 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 FFMN FI 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 Drosophila Rapid activation of GF 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 Cockroach  Air movement hairs give directionality  Escape correct way!  Giant fibres 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
 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
									 
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                             
                                            