Many muscles of the locust are supplied by dorsal unpaired median neurons (DUM neurons) that release octopamine and alter the contractions caused by spikes in motor neurons. To determine when these neuromodulatory neurons are normally activated during behaviour, intracellular recordings were made simultaneously from them and from identified motor neurons during the specific motor pattern that underlies kicking. A kick consists of a rapid and powerful extension of the tibia of one or both hind legs that is produced by a defined motor pattern. Only 3 identified DUM neurons of the 20 in the metathoracic ganglion spike during a kick, and they supply muscles involved in generating the kick. Their spikes occur in a distinctive and repeatable pattern that is closely linked to the pattern of spikes in the flexor and extensor tibiae motor neurons. When the extensor and flexor muscles cocontract, these three DUM neurons produce a burst of spikes at frequencies that can rise to 25 Hz, and with the number of spikes (3-15) related to the duration of this phase of the motor pattern. The spikes stop when the flexor muscle is inhibited and therefore before the tibia is extended rapidly. The other DUM neurons which supply muscles that are not directly involved in kicking are either inhibited or spike only sporadically. 2. The activation of a specific subset of DUM neurons during kicking may thus be timed to influence the action of the muscles that participate in this movement and appear to be controlled by the same circuits that determine the actions of the participating motor neurons. These modulatory neurons thus have specific individual actions in the control of movement.
