Behavioral Neuroscience, lecture on electric fish
USD Department of Biology
Behavioral Neuroscience
Electric Fish and JAR Behavior
Afferent Path of Electroreception
Control of Electric Organ Discharge
Efferent Output producing EOD
Jamming Avoidance Response
Electric field Glutamate
Electric Fish Figures
Electric Fish Circuitry
Acronyms/Abbreviations    end
VI. Efferent ouput to produce EOD  			

	A. Pacemaker nucleus (Pn) in medulla
		1. modulatory inputs lead to distinct behavioral patterns
			a. alter neuronal firing rate
				i. Glu input from diencephalon (PPn) to pacemaker cells
					(1) AMPA/NMDA
				ii. Glu input from mesencephalon (SPPn)to relay cells
					(1) NMDA 
			b. pacemaker + relay fire 1:1 with EO electrocytes
		2. pacemaker cells
			a. connect with each other and relay cells
				i. mixed synapses (chemical and electrical)
			b. synchronous firing
		3. relay cells
			a. project to spinal motor neurons
			b. Glutamate output
	B. Motor neurons in spinal cord
		1. specialized electromotor neurons
			a. innervate electrocytes of electric organ
				i. innervate just one side
				ii. Eigenmannia and all other electric fish
					(1) except Apteronotus
			b. comprise electric organ itself (Apteronotus)
				i. branched nerve endings
		2. stimulated to fire by AMPA/NMDA receptors
			a. Glu from Pn relay cells
		3. ACh = acetylcholine output
			a. similar to neuromuscular junction
	C. Electric Organ
		1. Size and shape depend on environment and function
			a. Weakly electric fish 
				i. native to highly conductive water 
					(1) short, broad electric organs
				ii. less conductive water 
					(1) long, thin organs
			b. Extends bilaterally from tip of tail anteriorly
		2. Electrocytes 
			a. Modified striatal muscle fibers  
				i. maintained/driven by ion channels
				  (similar to Na+/K+ ATPase)
			b. Flat disc-like multinucleate cells 
				i. stacked similar to a battery
					(1) parallel rows along length of organ
					(2)  stacks of electrocytes 
						  discharge capacity 
				ii. embedded in gelatinous matrix
				iii. sheathed in high resistance connective tissue
					(1) channels flow of current along the
						    electric organ
				iv. specifuc morphology of the electrocytes
					varies greatly between species 
			c. May contain one or two electrically excitable faces
		3. Activated by nicotinic ACh receptors
			a. neuromuscular junction on one side of disc
			b. Function by pumping Na+ and K+ through channels
	D. Electric Organ Discharge Physiology
		1. Pattern of discharge produces "pulse" or "wave"
			a. Pulse: EOD emitted at arbitrary intervals
				i. low and irregular repetition rate
			b. Wave: EOD emitted at patterned and regular intervals
				i. pacemaker necessary
				ii. brief signals at consistent interval
		2. Monophasic pulse
			a. electrocyte has one electrically excitable face (posterior)
				i. other face inexcitable
			b. membrane depolarized by ACh-Nicotinic synaptic input
				i. Na+ enters excitable face
				ii. K+ exits inexcitable face 
			c. net depolarization from electrocytes 
				i. mulitple membrane depolarizations
				ii. serial arrangement of electrocytes
				iii. yields summed the small electric potentials
				iv. much higher external potentials
			d. flows anteriorly along electric organ into water
				i. channeled by connctive tissue
					(1) rostrally  anterior cathode pole of EO
					(2) out into water
			e. electric field signal re-enters fishes tail
				i. anode pole
		2. Biphasic pulse
			a. electrocytes have two excitable faces
			b. first AP travels anteriorly toward head
			c. depolarization of second membrane sends AP toward tail
			d. alternate current flow: head then tail  EOD biphasic shape
		3. Wave-type EOD
			a. electrocytes similar to pulse EOD
				i. First, discharge one side of electric organ
				ii. then discharge the other side
					(1) discharge in series
			b. sinusoidal wave caused EOD pulses generated at regular rate
			c. frequency determined by pacemaker nucleus
			d. pulse frequency varies inversely with pulse length

VII. Jamming Avoidance Response