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A fundamental task performed by many visual systems is to distinguish apparent motion caused by eye movements from real motion occurring within the environment. During saccadic eye movements, this task is achieved by inhibitory signals of central and retinal origin that suppress the output of motion-detecting neurons. To investigate the retinally-generated component of this suppression, we used a computational model of a locust looming-detecting pathway that experiences saccadic suppression. This model received input from the camera of a mobile robot that performed simple saccade-like movements, allowing the model's response to simplified real stimuli to be tested. Retinally-generated saccadic suppression resulted from two inhibitory mechanisms within the looming-detector's input architecture. One mechanism fed inhibition forward through the network, inhibiting the looming-detector's initial response to movement. The second spread inhibition laterally within the network, suppressing the looming-detector's maintained response to movement. These mechanisms prevent a looming-detector model response to whole-field visual stimuli. In the locust, this mechanism of saccadic suppression may operate in addition to centrally-generated suppression. Because lateral inhibition is a common feature of early visual processing in many organisms, we discuss whether the mechanism of retinally-generated saccadic suppression found in the locust looming-detector model may also operate in these species.

The lobula giant movement detector (LGMD1 and -2) neurons in the locust visual system are parts of motion-sensitive pathways that detect objects approaching on a collision course. The dendritic processes of the LGMD1 and -2 in the lobula are localised to discrete regions, allowing the dendrites of each neuron to be distinguished uniquely. As was described previously for the LGMD1, the afferent processes onto the LGMD2 synapse directly with each other, and these synapses are immediately adjacent to their outputs onto the LGMD2. Here we present immunocytochemical evidence, using antibodies against choline-protein conjugates and a polyclonal antiserum against choline acetyltransferase (ChAT; Chemicon Ab 143), that the LGMD1 and -2 and the retinotopic units presynaptic to them contain acetylcholine (ACh). It is proposed that these retinotopic units excite the LGMD1 or -2 but inhibit each other. It is well established that ACh has both excitatory and inhibitory effects and may provide the substrate for a critical race in the LGMD1 or -2, between excitation caused by edges moving out over successive photoreceptors, and inhibition spreading laterally resulting in the selective response to objects approaching on a collision course. In the optic lobe, ACh was also found to be localised in discrete layers of the medulla and in the outer chiasm between the lamina and medulla. In the brain, the antennal lobes contained neurons that reacted positively for ACh. Silver- or haematoxylin and eosin-stained sections through the optic lobe confirmed the identities of the positively immunostained neurons., (Copyright 2000 Wiley-Liss, Inc.)

The visual systems of insects perform complex processing using remarkably compact neural circuits, yet these circuits are often studied using simplified stimuli which fail to reveal their behaviour in more complex visual environments. We address this issue by testing models of these circuits in real-world visual environments using a mobile robot. In this paper we focus on the lobula giant movement detector (LGMD) system of the locust which responds selectively to objects which approach the animal on a collision course and is thought to trigger escape behaviours. We show that a neural network model of the LGMD system shares the preference for approaching objects and detects obstacles over a range of speeds. Our results highlight aspects of the basic response properties of the biological system which have important implications for the behavioural role of the LGMD.

The image of a rapidly approaching object has to elicit a quick response. An animal needs to know that the object is approaching on a collision course and how imminent a collision is. The relevant information can be computed from the way that the image of the object grows on the retina of one eye. Firm data about the types of neurones that react to such looming stimuli and trigger avoidance reactions come from recent studies on the pigeon and the locust. The neurones responsible are tightly tuned to detect objects that are approaching on a direct collision course. In the pigeon these neurones signal the time remaining before collision whereas in the locust they have a crucial role in the simple strategy this animal uses to detect an object approaching on a collision course.

The lobula giant movement detector (LGMD) neuron in the locust visual system is part of a motion-sensitive pathway that detects objects approaching on a collision course. Here we show that the retinotopic units presynaptic to the LGMD make synapses directly with each other and these synapses are immediately adjacent to their outputs onto the LGMD. Synapses occur along the fine dendrites of the LGMD in the distal lobula, often in large numbers and completely covering the LGMD processes. Gamma aminobutyric acid (GABA) was eliminated as a possible neurotransmitter at these synapses when immunogold-tagged monoclonal GABA antibody did not specifically label the afferent processes. We used a histochemical method to demonstrate that acetylcholine esterase, the enzyme that hydrolyses acetylcholine at cholinergic synapses, was present in the synaptic clefts between the retinotopic units and along the membrane of the LGMD. It is well established that acetylcholine has both excitatory and inhibitory effects and we propose that these retinotopic units excite the LGMD, but inhibit each other; and that the synapses form the substrate for a critical race between excitation caused by edges moving out over successive photoreceptors, and inhibition spreading laterally. This results in the selective response to objects approaching on a collision course.

The locust descending contralateral motion detector (DCMD) responds to movements anywhere within a wide visual field, but responds most strongly to the images of approaching objects. It has been claimed that the response peaks before the end of an approach movement, providing a signal that anticipates collision. However, we find that when the locust eye is presented with appropriate computer-generated images of approaching objects, the response builds up until after movement has stopped. Premature peaking in the response is due to failure to stimulate the eye with sufficiently small and frequent jumps in image edges. We conclude that the DCMD signals impending collision by tracking edge motion throughout object approach.

The LGMD is the major source of visual input from the compound eye to the ipsilateral DCMD. Inactivating the LGMD or hyperpolarizing it, so it no longer spikes, abolishes the response of the DCMD to the visual stimulus. Synaptic transmission between the LGMD and DCMD neurones is chemical. A spike in the LGMD terminals induces a postsynaptic potential in the DCMD dendrites, with a transmission delay of 1 ms. There is a conductance increase in the DCMD during an LGMD-mediated PSP. The conductance increase occurs at membrane potentials when the current/voltage relationship of the DCMD membrane is linear, and at several different membrane potentials. The LGMD-mediated PSP within the dendritic region of the DCMD has a rise time of 1.3 ms, a half-time for decay of 2.2 ms and a total duration of 8.3 ms. In the cell body it has a rise time of 3.3 ms, a half-time for decay of 8 ms and a total duration of 21.3 ms. The amplitude of the LGMD-mediated PSP depends on the membrane potential of the DCMD. The PSP amplitude is increased by membrane hyperpolarization and decreased by membrane depolarizations. At a membrane potential 30 mV more positive than resting potential the extrapolated size of the PSP is zero. The synaptic efficiency of the LGMD-DCMD connection is usually 1.2. (formula; see text) There is a threshold of 13 mV in the LGMD before synaptic transmission occurs. Currents less than 13 mV are not transmitted in either direction across the synapse although they do reach the synaptic region if they are injected at the extremities of the neurones within the brain. Length constants for the LGMD are 0.36 mm between points c and d in the protocerebrum and 0.63 mm between point b in the optic lobe and point d in the protocerebrum. The length constant measured between the dendrite region of the DCMD and its cell body is 1.34 mm. DCMD spikes and PSPs follow spikes in the LGMD at a constant latency at frequencies up to 400 Hz. Usually a spike in the LGMD induces a spike in the DCMD.