Understanding the mechanism of facilitation in hoverfly TSDNs.

Many animals use visual motion cues to track and pursue small, fast-moving targets, such as prey or conspecifics. In target-pursuing insects, including dragonflies and hoverflies, Small Target Motion Detector (STMD) neurons are found in the optic lobes and are believed to be presynaptic to Target Selective Descending Neurons (TSDNs) that project to motor command centres. While STMDs respond robustly to target motion - even when displayed against moving backgrounds - TSDN target responses are modulated by background motion. Depending on whether the background motion is syn- or contra-directional to the target motion, the response of the TSDNs is either suppressed or facilitated (amplified). This suggests that TSDNs not only receive input from STMDs but also from neurons sensitive to background motion, but this neural circuit is not clearly understood. To explore the underlying neural mechanisms, we developed three candidate TSDN circuit models - which combine input from bio-plausible STMDs and optic flow-sensitive Lobula Plate Tangential Cells (LPTCs) in different ways - and fitted them to published electrophysiology data from hoverfly TSDNs. We then tested the best-fitting models against new electrophysiological data using different background patterns. We found that the overall best model suggests simple inhibition from LPTCs with the same preferred direction as the STMDs feeding into the TSDN. This parsimonious mechanism can explain the facilitation and suppression of TSDN responses to small targets, and may inform similar studies in other animals.