Elementary Motion Detectors Research Articles

Fully Memristive Elementary Motion Detectors for a Maneuver Prediction.

Insects can efficiently perform object motion detection via a specialized neural circuit, called an elementary motion detector (EMD). In contrast, conventional machine vision systems require significant computational resources for dynamic motion processing. Here, a fully memristive EMD (M-EMD) is presented that implements the Hassenstein-Reichardt (HR) correlator, a biological model of the EMD. The M-EMD consists of a simple Wye (Y) configuration, including a static resistor, a dynamic memristor, and a Mott memristor. The resistor and dynamic memristor introduce different signal delays, enabling spatio-temporal signal integration in the subsequent Mott memristor, resulting in a direction-selective response. In addition, a neuromorphic system is developed employing the M-EMDs to predict a lane-changing maneuver by vehicles on the road. The system achieved a high accuracy (> 87%) in predicting future lane-changing maneuvers on the Next Generation Simulation (NGSIM) dataset while reducing the computational cost by 92.9% compared to the conventional neuromorphic system without the M-EMD, suggesting its strong potential for edge-level computing.

Divisive normalization processors in the early visual system of the Drosophila brain

Divisive normalization is a model of canonical computation of brain circuits. We demonstrate that two cascaded divisive normalization processors (DNPs), carrying out intensity/contrast gain control and elementary motion detection, respectively, can model the robust motion detection realized by the early visual system of the fruit fly. We first introduce a model of elementary motion detection and rewrite its underlying phase-based motion detection algorithm as a feedforward divisive normalization processor. We then cascade the DNP modeling the photoreceptor/amacrine cell layer with the motion detection DNP. We extensively evaluate the DNP for motion detection in dynamic environments where light intensity varies by orders of magnitude. The results are compared to other bio-inspired motion detectors as well as state-of-the-art optic flow algorithms under natural conditions. Our results demonstrate the potential of DNPs as canonical building blocks modeling the analog processing of early visual systems. The model highlights analog processing for accurately detecting visual motion, in both vertebrates and invertebrates. The results presented here shed new light on employing DNP-based algorithms in computer vision.

Disynaptic inhibition shapes tuning of OFF-motion detectors in Drosophila

The circuitry underlying the detection of visual motion in Drosophila melanogaster is one of the best studied networks in neuroscience. Lately, electron microscopy reconstructions, algorithmic models, and functional studies have proposed a common motif for the cellular circuitry of an elementary motion detector based on both supralinear enhancement for preferred direction and sublinear suppression for null-direction motion. In T5 cells, however, all columnar input neurons (Tm1, Tm2, Tm4, and Tm9) are excitatory. So, how is null-direction suppression realized there? Using two-photon calcium imaging in combination with thermogenetics, optogenetics,apoptotics, and pharmacology, we discovered that it is via CT1, the GABAergic large-field amacrine cell, where the different processes have previously been shown to act in an electrically isolated way. Within each column, CT1 receives excitatory input from Tm9 and Tm1 and provides the sign-inverted, now inhibitory input signal onto T5. Ablating CT1 or knocking down GABA-receptor subunit Rdl significantly broadened the directional tuning of T5 cells. It thus appears that the signal of Tm1 and Tm9 is used both as an excitatory input for preferred direction enhancement and, through a sign inversion within the Tm1/Tm9-CT1 microcircuit, as an inhibitory input for null-direction suppression.

Nonlinear, neuronal adaptation in insect vision models improves target discrimination within repetitively moving backgrounds

Neurons which respond selectively to small moving targets, even against a cluttered background, have been identified in several insect species. To investigate what underlies these robust and highly selective responses, researchers have probed the neuronal circuitry in target-detecting, visual pathways. Observations in flies reveal nonlinear adaptation over time, composed of a fast onset and gradual decay. This adaptive processing is seen in both of the independent, parallel pathways encoding either luminance increments (ON channel) or decrements (OFF channel). The functional significance of this adaptive phenomenon has not been determined from physiological studies, though the asymmetrical time course suggests a role in suppressing responses to repetitive stimuli. We tested this possibility by comparing an implementation of fast adaptation against alternatives, using a model of insect ‘elementary small target motion detectors’. We conducted target-detecting simulations on various natural backgrounds, that were shifted via several movement profiles (and target velocities). Using performance metrics, we confirmed that the fast adaptation observed in neuronal systems enhances target detection against a repetitively moving background. Such background movement would be encountered via natural ego-motion as the insect travels through the world. These findings show that this form of nonlinear, fast-adaptation (suitably implementable via cellular biophysics) plays a role analogous to background subtraction techniques in conventional computer vision.

A bioinspired angular velocity decoding neural network model for visually guided flights

Efficient and robust motion perception systems are important pre-requisites for achieving visually guided flights in future micro air vehicles. As a source of inspiration, the visual neural networks of flying insects such as honeybee and Drosophila provide ideal examples on which to base artificial motion perception models. In this paper, we have used this approach to develop a novel method that solves the fundamental problem of estimating angular velocity for visually guided flights. Compared with previous models, our elementary motion detector (EMD) based model uses a separate texture estimation pathway to effectively decode angular velocity, and demonstrates considerable independence from the spatial frequency and contrast of the gratings. Using the Unity development platform the model is further tested for tunnel centering and terrain following paradigms in order to reproduce the visually guided flight behaviors of honeybees. In a series of controlled trials, the virtual bee utilizes the proposed angular velocity control schemes to accurately navigate through a patterned tunnel, maintaining a suitable distance from the undulating textured terrain. The results are consistent with both neuron spike recordings and behavioral path recordings of real honeybees, thereby demonstrating the model’s potential for implementation in micro air vehicles which have only visual sensors.

Mechanism for analogous illusory motion perception in flies and humans

Visual motion detection is one of the most important computations performed by visual circuits. Yet, we perceive vivid illusory motion in stationary, periodic luminance gradients that contain no true motion. This illusion is shared by diverse vertebrate species, but theories proposed to explain this illusion have remained difficult to test. Here, we demonstrate that in the fruit fly Drosophila, the illusory motion percept is generated by unbalanced contributions of direction-selective neurons' responses to stationary edges. First, we found that flies, like humans, perceive sustained motion in the stationary gradients. The percept was abolished when the elementary motion detector neurons T4 and T5 were silenced. In vivo calcium imaging revealed that T4 and T5 neurons encode the location and polarity of stationary edges. Furthermore, our proposed mechanistic model allowed us to predictably manipulate both the magnitude and direction of the fly's illusory percept by selectively silencing either T4 or T5 neurons. Interestingly, human brains possess the same mechanistic ingredients that drive our model in flies. When we adapted human observers to moving light edges or dark edges, we could manipulate the magnitude and direction of their percepts as well, suggesting that mechanisms similar to the fly's may also underlie this illusion in humans. By taking a comparative approach that exploits Drosophila neurogenetics, our results provide a causal, mechanistic account for a long-known visual illusion. These results argue that this illusion arises from architectures for motion detection that are shared across phyla.

Event-Based Eccentric Motion Detection Exploiting Time Difference Encoding.

Attentional selectivity tends to follow events considered as interesting stimuli. Indeed, the motion of visual stimuli present in the environment attract our attention and allow us to react and interact with our surroundings. Extracting relevant motion information from the environment presents a challenge with regards to the high information content of the visual input. In this work we propose a novel integration between an eccentric down-sampling of the visual field, taking inspiration from the varying size of receptive fields (RFs) in the mammalian retina, and the Spiking Elementary Motion Detector (sEMD) model. We characterize the system functionality with simulated data and real world data collected with bio-inspired event driven cameras, successfully implementing motion detection along the four cardinal directions and diagonally.

Drosophila-inspired 3D moving object detection based on point clouds

3D moving object detection is one of the most critical tasks in dynamic scene analysis. In this paper, we propose a novel Drosophila-inspired 3D moving object detection method using Lidar sensors. According to the theory of elementary motion detector, we have developed a motion detector based on the shallow visual neural pathway of Drosophila. This detector is sensitive to the movement of objects and can well suppress background noise. Designing neural circuits with different connection modes, the approach searches for motion areas in a coarse-to-fine fashion and extracts point clouds of each motion area to form moving object proposals. An improved 3D object detection network is then used to estimate the point clouds of each proposal and efficiently generates the 3D bounding boxes and the object categories. We evaluate the proposed approach on the widely-used KITTI benchmark, and state-of-the-art performance was obtained by using the proposed approach on the task of motion detection.

Bioinspired Approaches for Autonomous Small-Object Detection and Avoidance

Small-object detection and avoidance in unknown environments is a significant challenge to overcome for small autonomous vehicles that are generally highly agile and restricted in payload and computational processing power. Typical machine-vision and range measurement-based solutions suffer either from restricted fields-of-view or significant computational complexity and are, hence, not easily portable to small platforms. In order to overcome these drawbacks, in this paper, two novel bioinspired approaches are proposed to extract information about small-field objects contained in planar optic flow. The first approach, which is analogous to the small-field extraction process hypothesized to occur in the lobula plate of the fly visual system, is based on the Fourier residual analysis of instantaneous optic flow. Alternatively, the flow-of-flow method is the engineering analogue of the small-field extraction process thought to occur in the fruit-fly's medulla, and extracts high-frequency content of optic flow by means of an elementary motion detector array. Both approaches extract instantaneous relative range and bearing of small-field obstacles from planar optic flow in a local environment characterized by small and wide-field obstacles, which is then combined with an artificial potential function-based low-order steering control law. The proposed sensing and control scheme is experimentally validated with a quadrotor vehicle that is able to effectively navigate an unknown environment laden with small-field clutter. This bioinspired approach is computationally efficient, which renders extraction of vehicle velocity and local environment structure superfluous, and thus, serves as a robust, reflexive solution to the problem of small-object detection, and avoidance for small autonomous robots.

Dynamic nonlinearities enable direction opponency in Drosophila elementary motion detectors.

Direction-selective neurons respond to visual motion in a preferred direction. They are direction-opponent if they are also inhibited by motion in the opposite direction. In flies and vertebrates, direction opponency has been observed in second-order direction-selective neurons, which achieve this opponency by subtracting signals from first-order direction-selective cells with opposite directional tunings. Here, we report direction opponency in Drosophila that emerges in first-order direction-selective neurons, the elementary motion detectors T4 and T5. This opponency persists when synaptic output from these cells is blocked, suggesting that it arises from feedforward, not feedback, computations. These observations exclude a broad class of linear-nonlinear models that have been proposed to describe direction-selective computations. However, they are consistent with models that include dynamic nonlinearities. Simulations of opponent models suggest that direction opponency in first-order motion detectors improves motion discriminability by suppressing noise generated by the local structure of natural scenes.

The Neuronal Basis of an Illusory Motion Percept Is Explained by Decorrelation of Parallel Motion Pathways

Both vertebrates and invertebrates perceive illusory motion, known as "reverse-phi," in visual stimuli that contain sequential luminance increments and decrements. However, increment (ON) and decrement (OFF) signals are initially processed by separate visual neurons, and parallel elementary motion detectors downstream respond selectively to the motion of light or dark edges, often termed ON- and OFF-edges. It remains unknown how and where ON and OFF signals combine to generate reverse-phi motion signals. Here, we show that each of Drosophila's elementary motion detectors encodes motion by combining both ON and OFF signals. Their pattern of responses reflects combinations of increments and decrements that co-occur in natural motion, serving to decorrelate their outputs. These results suggest that the general principle of signal decorrelation drives the functional specialization of parallel motion detection channels, including their selectivity for moving light or dark edges.

Linear Summation Underlies Direction Selectivity in Drosophila

While linear mechanisms lay the foundations of feature selectivity in many brain areas, direction selectivity in the elementary motion detector (EMD) of the fly has become a paradigm of nonlinear neuronal computation. We have bridged this divide by demonstrating that linear spatial summation cangenerate direction selectivity in the fruit fly Drosophila. Using linear systems analysis and two-photon imaging of a genetically encoded voltage indicator, we measure the emergence of direction-selective (DS) voltage signals in the Drosophila OFF pathway. Our study is a direct, quantitative investigation of the algorithm underlying directional signals, with the striking finding that linear spatial summation is sufficient for the emergence of direction selectivity. A linear stage of the fly EMD strongly resembles similar computations in vertebrate visual cortex, demands a reappraisal of the role of upstream nonlinearities, and implicates the voltage-to-calcium transformation in the refinement of feature selectivity in this system. VIDEO ABSTRACT.

Spiking Elementary Motion Detector in Neuromorphic Systems.

Apparent motion of the surroundings on an agent's retina can be used to navigate through cluttered environments, avoid collisions with obstacles, or track targets of interest. The pattern of apparent motion of objects, (i.e., the optic flow), contains spatial information about the surrounding environment. For a small, fast-moving agent, as used in search and rescue missions, it is crucial to estimate the distance to close-by objects to avoid collisions quickly. This estimation cannot be done by conventional methods, such as frame-based optic flow estimation, given the size, power, and latency constraints of the necessary hardware. A practical alternative makes use of event-based vision sensors. Contrary to the frame-based approach, they produce so-called events only when there are changes in the visual scene. We propose a novel asynchronous circuit, the spiking elementary motion detector (sEMD), composed of a single silicon neuron and synapse, to detect elementary motion from an event-based vision sensor. The sEMD encodes the time an object's image needs to travel across the retina into a burst of spikes. The number of spikes within the burst is proportional to the speed of events across the retina. A fast but imprecise estimate of the time-to-travel can already be obtained from the first two spikes of a burst and refined by subsequent interspike intervals. The latter encoding scheme is possible due to an adaptive nonlinear synaptic efficacy scaling. We show that the sEMD can be used to compute a collision avoidance direction in the context of robotic navigation in a cluttered outdoor environment and compared the collision avoidance direction to a frame-based algorithm. The proposed computational principle constitutes a generic spiking temporal correlation detector that can be applied to other sensory modalities (e.g., sound localization), and it provides a novel perspective to gating information in spiking neural networks.

Spatial Encoding of Translational Optic Flow in Planar Scenes by Elementary Motion Detector Arrays

Elementary Motion Detectors (EMD) are well-established models of visual motion estimation in insects. The response of EMDs are tuned to specific temporal and spatial frequencies of the input stimuli, which matches the behavioural response of insects to wide-field image rotation, called the optomotor response. However, other behaviours, such as speed and position control, cannot be fully accounted for by EMDs because these behaviours are largely unaffected by image properties and appear to be controlled by the ratio between the flight speed and the distance to an object, defined here as relative nearness. We present a method that resolves this inconsistency by extracting an unambiguous estimate of relative nearness from the output of an EMD array. Our method is suitable for estimation of relative nearness in planar scenes such as when flying above the ground or beside large flat objects. We demonstrate closed loop control of the lateral position and forward velocity of a simulated agent flying in a corridor. This finding may explain how insects can measure relative nearness and control their flight despite the frequency tuning of EMDs. Our method also provides engineers with a relative nearness estimation technique that benefits from the low computational cost of EMDs.

White noise analysis for the correlation-type elementary motion detectors with half-wave rectifiers

The motion detection mechanism of insects has been attracted attention of many researchers. Several motion-detection models have been proposed on the basis of insect visual system studies. Here, we examine two models, the Hassenstein–Reichardt (HR) model and the two-detector (2D) model. We analytically obtain the mean and variance of the stationary responses of the HR and the 2D models to white noise, and we derive the signal-to-fluctuation-noise ratio (SFNR) to evaluate encoding abilities of the two models. Especially when analyzing the 2D model, we calculate higher-order cumulants of a rectified Gaussian. The results show that the 2D model robustly works almost as well as the HR model in several sets of parameters estimated on the basis of experimental data.

FPGA-Based EMD Assist Block for Motion Detection in Critical Environments

This study aims to carry out the implementation in FPGA (Field Programmable Gate Array) of a processing system capable of detecting and tracking moving vehicles without losing the system's reliability and efficiency of real-time application. Hence, for simple technique of tracking reach good results is proposed for removing shadows, an auxiliary processing based on insect vision system EMD (Elementary Motion Detector). The principle of this processing is based on cross-correlation and has been implemented in some work to get the direction of moving objects, however, this work is modified and implemented to auxiliary in removing shadows of objects detected using color, spatial and temporal information. The proposed auxiliary processing proves to be efficient in a video stream where the two most critical vehicles are selected with respect the cast shadow. The results of the implementation proposal are presented in terms of resource consumption and maximum operating frequency in a Spartan-6 FPGA-based development board. The experimental results are obtained in co-simulation system and show the effectiveness of the proposed system.

The Temporal Tuning of the Drosophila Motion Detectors Is Determined by the Dynamics of Their Input Elements

Detecting the direction of motion contained in the visual scene is crucial for many behaviors. However, because single photoreceptors only signal local luminance changes, motion detection requires a comparison of signals from neighboring photoreceptors across time in downstream neuronal circuits. For signals to coincide on readout neurons that thus become motion and direction selective, different input lines need to be delayed with respect to each other. Classical models of motion detection rely on non-linear interactions between two inputs after different temporal filtering. However, recent studies have suggested the requirement for at least three, not only two, input signals. Here, we comprehensively characterize the spatiotemporal response properties of all columnar input elements to the elementary motion detectors in the fruit fly, T4 and T5 cells, via two-photon calcium imaging. Between these input neurons, we find large differences in temporal dynamics. Based on this, computer simulations show that only a small subset of possible arrangements of these input elements maps onto a recently proposed algorithmic three-input model in a way that generates a highly direction-selective motion detector, suggesting plausible network architectures. Moreover, modulating the motion detection system by octopamine-receptor activation, we find the temporal tuning of T4 and T5 cells to be shifted toward higher frequencies, and this shift can be fully explained by the concomitant speeding of the input elements.

Temporally Diverse Excitation Generates Direction-Selective Responses in ON- and OFF-Type Retinal Starburst Amacrine Cells

The complexity of sensory receptive fields increases from one synaptic stage to the next. In many cases, increased complexity is achieved through spatiotemporal interactions between convergent excitatory and inhibitory inputs. Here, we present evidence that direction selectivity (DS), a complex emergent receptive field property of retinal starburst amacrine cells (SACs), is generated by spatiotemporal interactions between functionally diverse excitatory inputs. Electrophysiological whole-cell recordings from ON and OFF SACs show distinct temporal differences in excitation following proximal compared with distal stimulation of their receptive fields. Distal excitation is both faster and more transient, ruling out passive filtering by the dendrites and indicating a task-specific specialization. Model simulations demonstrate that this specific organization of excitation generates robust DS responses in SACs, consistent with elementary motion detector models. These results indicate that selective integration of spatiotemporally patterned excitation is a computational mechanism for motion detection in the mammalian retina.

Transgenic line for the identification of cholinergic release sites in Drosophila melanogaster.

ABSTRACTThe identification of neurotransmitter type used by a neuron is important for the functional dissection of neuronal circuits. In the model organism Drosophila melanogaster, several methods for discerning the neurotransmitter systems are available. Here, we expanded the toolbox for the identification of cholinergic neurons by generating a new line FRT-STOP-FRT-VAChT::HA that is a conditional tagged knock-in of the vesicular acetylcholine transporter (VAChT) gene in its endogenous locus. Importantly, in comparison to already available tools for the detection of cholinergic neurons, the FRT-STOP-FRT-VAChT::HA allele also allows for identification of the subcellular localization of the cholinergic presynaptic release sites in a cell-specific manner. We used the newly generated FRT-STOP-FRT-VAChT::HA line to characterize the Mi1 and Tm3 neurons in the fly visual system and found that VAChT is present in the axons of both cell types, suggesting that Mi1 and Tm3 neurons provide cholinergic input to the elementary motion detectors, the T4 neurons.

Peripheral Processing Facilitates Optic Flow-Based Depth Perception.

Flying insects, such as flies or bees, rely on consistent information regarding the depth structure of the environment when performing their flight maneuvers in cluttered natural environments. These behaviors include avoiding collisions, approaching targets or spatial navigation. Insects are thought to obtain depth information visually from the retinal image displacements (“optic flow”) during translational ego-motion. Optic flow in the insect visual system is processed by a mechanism that can be modeled by correlation-type elementary motion detectors (EMDs). However, it is still an open question how spatial information can be extracted reliably from the responses of the highly contrast- and pattern-dependent EMD responses, especially if the vast range of light intensities encountered in natural environments is taken into account. This question will be addressed here by systematically modeling the peripheral visual system of flies, including various adaptive mechanisms. Different model variants of the peripheral visual system were stimulated with image sequences that mimic the panoramic visual input during translational ego-motion in various natural environments, and the resulting peripheral signals were fed into an array of EMDs. We characterized the influence of each peripheral computational unit on the representation of spatial information in the EMD responses. Our model simulations reveal that information about the overall light level needs to be eliminated from the EMD input as is accomplished under light-adapted conditions in the insect peripheral visual system. The response characteristics of large monopolar cells (LMCs) resemble that of a band-pass filter, which reduces the contrast dependency of EMDs strongly, effectively enhancing the representation of the nearness of objects and, especially, of their contours. We furthermore show that local brightness adaptation of photoreceptors allows for spatial vision under a wide range of dynamic light conditions.