Descending neurons supplying the neck and flight motor of Diptera: physiological and anatomical characteristics

In Diptera, dorsal neuropils of the pro-, meso-, and metathoracic ganglia supply motor neurons to neck and flight muscles. Motor circuits are supplied by more than 50 pairs of descending neurons (DNs) whose dendritic trees in the brain are restricted to dorsal neuropils of the deutocerebrum where they are grouped together into discrete clusters. Each cluster is visited by wide-field motion-sensitive neurons and by morphologically small-field retinotopic elements. This organization suggests that flight descending neurons should respond to complex stimuli reflecting panoramic movement and small-field motion. Intracellular recordings, combined with dye filling, confirm this. Certain descending neurons responding to visual flow fields terminate bilaterally in superficial pterothoracic neuropils, at the level of indirect (power) flight muscle motor neurons. Other DNs terminate laterally, and provide segmental collaterals to areas containing neck and direct (steering) flight muscle motor neurons. Such DNs are activated by wide-field directional stimuli corresponding to pitch, roll, or yaw, and to small-field stimuli. Appropriate directional mechanosensory stimuli also activate dorsal descending neurons. The significance of dorsal descending neurons for the control of flight is discussed and compared with studies on course deviation neurons in other insects. It is suggested that, in Diptera, dorsal descending neurons may separately be involved in the control of velocity, stabilization, and steering manoeuvres.