# Understanding the mechanism of facilitation in hoverfly TSDNs

## TL;DR

This paper explores how hoverflies combine visual cues to track small targets, revealing a neural mechanism that explains how background motion affects target pursuit.

## Contribution

The study proposes a novel computational model of TSDN circuits that explains facilitation and suppression of target responses using inhibition from LPTCs.

## Key findings

- The best model suggests simple inhibition from LPTCs with the same direction preference as STMDs.
- This mechanism explains TSDN facilitation and suppression in response to small targets.
- The model does not require learning, suggesting innate sensorimotor behaviors in hoverflies.

## Abstract

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.

Many human sports, including tennis, football, and basketball, rely on the ability to visually detect and respond to the motion of a small, rapidly moving object. Indeed, some sports stars seem to have an uncanny ability to predict the future location of a ball. Similarly, many animals also need to detect the motion of small objects, as these may represent an approaching predator, prey, or a conspecific that needs to be avoided or pursued. Insects, in particular, which have small brains with very modest power requirements, appear to solve the problem of visually detecting small targets effortlessly. Male hoverflies naturally engage in complex pursuit behaviour, and since they have specialised neurons for target tracking, hoverflies are ideal for understanding the underlying neural circuitry. Here, we combine neural recordings from hoverflies with computational modelling to show how neurons that respond selectively to target motion are combined with neurons that respond to widefield optic-flow to generate behaviourally relevant sensorimotor responses. Importantly, our model does not involve learning and hence suggests that these behaviours can be innate.