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L31. Neural Circuits for Escape and Startle

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L31. Neural Circuits for Escape and Startle Motor Behavior Motor control of escape and startle as examples of: Coordination of: multiple muscle groups L31. Neural Circuits for appendages Escape and Startle limbs Speed …and what it says about decision Stereotyped, repetitive acts making in the Nervous system Decisions Oct. 19, 2011 BioNB 4240 Cornell University © Carl D. Hopkins 1qs c 2 Decision making: a motor hierarchy 3 4 Command Neuron Concept • Keis Wiersma & Ikeda (1964) – interneurons in crayfish tail – electrical stimulation of single cell evoked coordinated movement (swimmeret movement, posture control, escape) – pattern of stimulation was unimportant 5 6 1 Command neuron in cricket brain Escape • Rapid: Latency can be less than 10-20 ms • All or none (decisive!) • Complex: many actions coordinated. • Dominant: once initiated, blocks all other behavioral acts. • Illustrates decision making properties of command neurons. 7 8 Escape: the nervous system of the crayfish Procambarus clarkii 9 10 Flexion Escape Control by Giant Anatomy of Escape Interneurons FLEXOR Muscles EXTENSOR Muscles Giant interneuron fires Implanted electrodes record spikes from ventral nerve cord. Giant motor neuron fires Recorded activity during escape response: giant interneuron fires; flexor muscle in abdominal segments fire. Fast flexor muscles contract Transverse section of the connectives shows two giant interneurons: lateral giant, medial giant. 11 12 2 Lateral Giant Escape Lateral Giant Escape mechanical stim. to tail mechanical stim. to tail rostral segments flex. rostral segments flex. abrupt, single flexion drawing tail abrupt, single flexion drawing tail upward. Somersault. upward. Somersault. Medial giant escape. Medial giant escape. anterior stimulation (visual, anterior stimulation (visual, mechanical) mechanical) All segments flex All segments flex tail flip, propels animal tail flip, propels animal backward. backward. Krasne & Wine (1972) Krasne & Wine (1972) 13 14 Extended Escape Swimming LGI vs. MGI Escape NON-GIANT ESCAPE Extended swimming: flexion, extension at 20 Hz. Multiple contractions. Does not involve lateral or medial giants (except for first flip) • weaker and slower than MGI escape or LGI escape. • multiple forms: linear flips, twisting flips, pitching flips (following medial giant: linear flips only; following lateral giant: pitching flips) 15 16 Speed of Response Physiology of Escape Latency of flexion response is 10 ms Electrical synapses (fast) large neurons (fast conduction velocity) large synapses (strong EPSP) Electrical stimulation of LGI evokes spike in motor giant. Delay: short; electrical synapse Recording from LGI in response electrical stimulation at sensory receptor neuron. Note two 17 synaptic events: alpha and beta 18 3 LGI Escape Circuit LGI Escape Circuit Mechanosensory Input: ~1000 sensory hairs on tail. Sensory fibers (tail afferents) are directionally sensitive. Tuned. 80 Hz input (no response to small, low Freq. wave action). Sensory interneurons: receive inputs from sensory afferents. Afferent output to lateral giants via alpha pathway (direct, electrotonic); beta (indirect, chemical) 19 20 Lateral Giant Stimulating and recording set up. Recording from nerve Short latency EPSPs from sensory hairs (both direct and indirect). cord; Electrical synapses are below threshold: many required to stimulate giant. Motor nerve; Sensory stimulation. Lateral Giant makes decision to act. Recording from Lateral LGI fires: tail flip (LGI is sufficient) giant. Lateral giant is essential to action (necessary): Intracellular current stimulation. 1) hyperpolarization of the lateral giant will block the response to normal stimuli. 2) the amount of hyperpolarization needed to block the tail flip is exactly B) No hyperpolarization. the same as that needed to block LGI from spiking. C) hyperpolarized. 3) In hyperpolarized cell, the EPSP gets increasingly larger and larger with increased stimulus, but does not fire a spike. No behavior. LGI is the decision cell in this circuit. 21 22 Graded EPSP with Kupfermann and Weiss increasing stimulus strength shows that input to LGI is gradual. Threshold is sharp. Neuron must be both necessary and sufficient for behavior. LGI is decision making cell (all or none). 23 24 4 LGI neurons causes widespread inhibition LGI as Executive Neuron following command discharge Once the command is given to act, additional control must be exerted in order to inhibit competing behavioral acts. This is the executive function of a Command Neuron. extracellular from flexor motor nerve extensor inhibitor EPSP fast extensor motor neuron IPSP Inhibition of extensor muscles: (3 pathways): - excite extensor muscle inhibitors - inhibit motor neurons on fast extensor muscles - inhibit muscle receptor organs that normally respond to extension stretch. 25 26 EIs Executive control: inhibition with Flow diagram for escape responses in crayfish different delays: Command-driven inhibition occurs at a, b, c, d, e Relative times of inhibition after the LGI fires, for the points, a-e in the circuit 27 28 Modification of Escape Other Examples of Escape • Restraint • Power dive in noctuid moth • Feeding • wind-driven escape in cockroach • Learning (habituation) • Acoustic startle reflex • C-start reflex in fish 29 30 5 Escape in Fish 31 32 Simmons and Young (2010) Nerve cells and animal behaviour. Cambridge Univ. Press. 33 Simmons and Young (2010) Nerve cells and animal behaviour. Cambridge Univ. Press. 34 Korn and Faber (2005) Neuron 47(1):13-28 Korn and Faber (2005) Neuron 47(1):13-28 35 36 6 Lessons from escape behavior • Escape behavior is highly adaptive (essential), rapid, evoked by stimulus. An extreme case of decision making (very decisive). • In crayfish and in teleost fish, mediated by giant interneuron(s). • Different forms of behavior depending upon which cell is activated. • Command neurons: necessary and sufficient to produce entire behavior • Behavior is complex: involves sequence of acts, excitation of some cells, inhibition of others. Executive action of command neurons. • Command neuron’s function is modulated by many factors, including learning. 37 7 .
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