Abstract
Precise spike timing can be critical in sensory systems. In a few specific motor systems, we now know millisecond-scale timing of neural spikes is functionally important for behavior. However, we know little about the extent of timing codes across the whole motor program of an animal. Taking advantage of the relatively few motor units that control the wings of a hawk moth, we captured a comprehensive, spike-resolved motor program in tethered flight. We simultaneously record nearly every action potential from all muscles and the resulting forces. We find that timing encodes more information than rate in every motor unit. Motor units use consistent encoding, blending precise spike timing and rate information in a 3:1 ratio, despite their varying functions. Finally, we show that each muscle is coordinated with all other muscles through spike timings while spike rates are independent. Spike timing codes are ubiquitous, consistent, and essential for coordination.