A neurobiological and computational analysis of target discrimination in visual clutter by the insect visual system.

Some insects have the capability to detect and track small moving objects, often against cluttered moving backgrounds. Determining how this task is performed is an intriguing challenge, both from a physiological and computational perspective. Previous research has characterized higher-order neurons within the fly brain known as 'small target motion detectors‘ (STMD) that respond selectively to targets, even within complex moving surrounds. Interestingly, these cells still respond robustly when the velocity of the target is matched to the velocity of the background (i.e. with no relative motion cues). We performed intracellular recordings from intermediate-order neurons in the fly visual system (the medulla). These full-wave rectifying, transient cells (RTC) reveal independent adaptation to luminance changes of opposite signs (suggesting separate 'on‘ and 'off‘ channels) and fast adaptive temporal mechanisms (as seen in some previously described cell types). We show, via electrophysiological experiments, that the RTC is temporally responsive to rapidly changing stimuli and is well suited to serving an important function in a proposed target-detecting pathway. To model this target discrimination, we use high dynamic range (HDR) natural images to represent 'real-world‘ luminance values that serve as inputs to a biomimetic representation of photoreceptor processing. Adaptive spatiotemporal high-pass filtering (1st-order interneurons) shapes the transient 'edge-like‘ responses, useful for feature discrimination. Following this, a model for the RTC implements a nonlinear facilitation between the rapidly adapting, and independent polarity contrast channels, each with centre-surround antagonism. The recombination of the channels results in increased discrimination of small targets, of approximately the size of a single pixel, without the need for relative motion cues. This method of feature discrimination contrasts with traditional target and background motion-field computations. We show that our RTC-based target detection model is well matched to properties described for the higher-order STMD neurons, such as contrast sensitivity, height tuning and velocity tuning. The model output shows that the spatiotemporal profile of small targets is sufficiently rare within natural scene imagery to allow our highly nonlinear 'matched filter‘ to successfully detect many targets from the background. The model produces robust target discrimination across a biologically plausible range of target sizes and a range of velocities. We show that the model for small target motion detection is highly correlated to the velocity of the stimulus but not other background statistics, such as local brightness or local contrast, which normally influence target detection tasks. From an engineering perspective, we examine model elaborations for improved target discrimination via inhibitory interactions from correlation-type motion detectors, using a form of antagonism between our feature correlator and the more typical motion correlator. We also observe that a changing optimal threshold is highly correlated to the value of observer ego-motion. We present an elaborated target detection model that allows for implementation of a static optimal threshold, by scaling the target discrimination mechanism with a model-derived velocity estimation of ego-motion. Finally, we investigate the physiological relevance of this target discrimination model. We show that via very subtle image manipulation of the visual stimulus, our model accurately predicts dramatic changes in observed electrophysiological responses from STMD neurons.Thesis (Ph.D.) - University of Adelaide, School of Molecular and Biomedical Science, 200

Review

Detecting the direction of image motion is a fundamental component of visual computation, essential for survival of the animal. However, at the level of individual photoreceptors, the direction in which the image is shifting is not explicitly represented. Rather, directional motion information needs to be extracted from the photoreceptor array by comparing the signals of neighboring units over time. The exact nature of this process as implemented in the visual system of the fruit fly Drosophila melanogaster has been studied in great detail, and much progress has recently been made in determining the neural circuits giving rise to directional motion information. The results reveal the following: (1) motion information is computed in parallel ON and OFF pathways. (2) Within each pathway, T4 (ON) and T5 (OFF) cells are the first neurons to represent the direction of motion. Four subtypes of T4 and T5 cells exist, each sensitive to one of the four cardinal directions. (3) The core process of direction selectivity as implemented on the dendrites of T4 and T5 cells comprises both an enhancement of signals for motion along their preferred direction as well as a suppression of signals for motion along the opposite direction. This combined strategy ensures a high degree of direction selectivity right at the first stage where the direction of motion is computed. (4) At the subsequent processing stage, tangential cells spatially integrate direct excitation from ON and OFF-selective T4 and T5 cells and indirect inhibition from bi-stratified LPi cells activated by neighboring T4/T5 terminals, thus generating flow-field-selective responses