The neuromodulatory system plays a critical role in sensorimotor system function and animal behavior. Its influence on axons, however, remains enigmatic although axons possess receptors for a plethora of modulators, and pathologies of the neuromodulatory system impair neuronal communication. The most dramatic neuromodulatory effect on axons is ectopic spiking, a process common to many systems and neurons during which action potentials are elicited in the axon trunk and travel antidromically towards the site of sensory transduction. We argue that ectopic action potentials modify sensory encoding by invading the primary spike initiation zone in the periphery. This is a particularly intriguing concept, since it allows the modulatory system to alter sensory information processing. We demonstrate that aminergic modulation of a proprioceptive axon that elicits spontaneous ectopic action potentials changes spike frequency, which determines the burst behavior of the proprioceptor. Increasing ectopic spike frequency delayed the peripheral burst, caused reductions in spike number and burst duration, and changes in sensory firing frequency. Computational models show these effects depend on slow ionic conductances to modulate membrane excitability. Thus, axonal neuromodulation provides a means to rapidly influence sensory encoding without directly or locally affecting the sites of stimulus reception and spike initiation.
Part of the book: Sensory Nervous System