Lobula Plate Research Articles

Octopaminergic modulation of contrast sensitivity

Sensory systems adapt to prolonged stimulation by decreasing their response to continuous stimuli. Whereas visual motion adaptation has traditionally been studied in immobilized animals, recent work indicates that the animal's behavioral state influences the response properties of higher-order motion vision-sensitive neurons. During insect flight octopamine is released, and pharmacological octopaminergic activation can induce a fictive locomotor state. In the insect optic ganglia, lobula plate tangential cells (LPTCs) spatially pool input from local elementary motion detectors (EMDs) that correlate luminosity changes from two spatially discrete inputs after delaying the signal from one. The LPTC velocity optimum thereby depends on the spatial separation of the inputs and on the EMD's delay properties. Recently it was shown that behavioral activity increases the LPTC velocity optimum, with modeling suggesting this to originate in the EMD's temporal delay filters. However, behavior induces an additional post-EMD effect: the LPTC membrane conductance increases in flying flies. To physiologically investigate the degree to which activity causes presynaptic and postsynaptic effects, we conducted intracellular recordings of Eristalis horizontal system (HS) neurons. We constructed contrast response functions before and after adaptation at different temporal frequencies, with and without the octopamine receptor agonist chlordimeform (CDM). We extracted three motion adaptation components, where two are likely to be generated presynaptically of the LPTCs, and one within them. We found that CDM affected the early, EMD-associated contrast gain reduction, temporal frequency dependently. However, a CDM-induced change of the HS membrane conductance disappeared during and after visual stimulation. This suggests that physical activity mainly affects motion adaptation presynaptically of LPTCs, whereas post-EMD effects have a minimal effect.

Modulation of visual processing by flight behavior in Drosophila

Event Abstract Back to Event Modulation of visual processing by flight behavior in Drosophila Gaby Maimon1* 1 The Rockefeller University, Lab of Integrative Brain Function, United States A good theory of a visual system would explain why information about the outside world is reflected by the activity of neurons in just the observed format and not some other format. Data for such theories typically come from presenting stimuli to quiescent animals. In nature, however, animals tend to move around their environment, which has two important consequences. First, natural stimuli will tend to have rich dynamics, prescribed by the statistics of how the retina and animal move. How such natural dynamics affect sensory-neuron responses has received some attention. Second, and perhaps less studied, is the possibility that the act of locomotion itself may change the state of the visual system, such that visual-neuron activity becomes more suited for guiding locomotion and less suited for the needs of a stationary animal. To study the influence of locomotion on visual processing –– and other higher brain processes down the road –– we developed an electrophysiological recording system in Drosophila. We obtain whole-cell patch clamp recordings from genetically identified neurons simultaneously with tethered-flight behavior in the same animal. In our initial experiments, we targeted the vertical-system visual neurons (VS cells) of the lobula plate. Our data revealed that VS cells show at least two, prominent, physiological modulations during flight: a tonic shift in the baseline membrane voltage and a strong boost of visually driven activity. Both these modulations are likely to change the chemically or electrically mediated synaptic output at the VS-cell terminals. When the animal stops flying, visual responses remain boosted for many seconds whereas the baseline membrane voltage returns to its pre-flight level immediately, suggesting that these two effects have separable underlying mechanisms. Functionally, one possibility is that stronger visual responses might allow for more accurate or faster locomotion-specific behavior. Because electrical signaling is energetically costly, the brain may also save energy by keeping cellular responses at a minimum when neurons are not actively being used to drive behavior. These results demonstrate the importance of studying visual neurons, and formulating visual-system theories, not only in the context of naturalistic sensory input but also in the context of ethologically relevant locomotor output. Keywords: Drosophila, flight, patch clamp, Vision, Neurophysiology, modulation Conference: Tenth International Congress of Neuroethology, College Park. Maryland USA, United States, 5 Aug - 10 Aug, 2012. Presentation Type: Invited Symposium (only for people who have been invited to a particular symposium) Topic: Neuromodulation Citation: Maimon G (2012). Modulation of visual processing by flight behavior in Drosophila. Conference Abstract: Tenth International Congress of Neuroethology. doi: 10.3389/conf.fnbeh.2012.27.00067 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 04 Jul 2012; Published Online: 07 Jul 2012. * Correspondence: Dr. Gaby Maimon, The Rockefeller University, Lab of Integrative Brain Function, New York, New York, 10065, United States, maimon@rockefeller.edu Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Gaby Maimon Google Gaby Maimon Google Scholar Gaby Maimon PubMed Gaby Maimon Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Bio-inspired visual ego-rotation sensor for MAVs

Flies are capable of extraordinary flight maneuvers at very high speeds largely due to their highly elaborate visual system. In this work we present a fly-inspired FPGA based sensor system able to visually sense rotations around different body axes, for use on board micro aerial vehicles (MAVs). Rotation sensing is performed analogously to the fly's VS cell network using zero-crossing detection. An additional key feature of our system is the ease of adding new functionalities akin to the different tasks attributed to the fly's lobula plate tangential cell network, such as object avoidance or collision detection. Our implementation consists of a modified eneo SC-MVC01 SmartCam module and a custom built circuit board, weighing less than 200 g and consuming less than 4 W while featuring 57,600 individual two-dimensional elementary motion detectors, a 185° field of view and a frame rate of 350 frames per second. This makes our sensor system compact in terms of size, weight and power requirements for easy incorporation into MAV platforms, while autonomously performing all sensing and processing on-board and in real time.

Linking vision and action in Drosophila

Event Abstract Back to Event Linking vision and action in Drosophila Vivek Jayaraman1* 1 Janelia Farm Research Campus, Howard Hughes Medical Institute, United States My lab’s goal is to extract principles of neural circuit operation that underlie complex behavior in Drosophila melanogaster. Powerful genetic tools available in the fly allow us to target selected subsets of neurons and manipulate and record their activity using genetically encoded calcium indicators and electrophysiology. Over the past few years, we have developed preparations that permit physiological recordings from identified neurons in the central brain of tethered walking and flying flies during open- and closed-loop behavior. In our initial experiments we demonstrated that calcium responses of the motion-sensitive horizontal system lobula plate tangential cells in the early visual system are correlated with the tethered fly’s optomotor behavior. We also found that these neurons modulate their response gain and temporal frequency (motion-speed) tuning depending on behavioral state. I will briefly review these results, and describe more recent findings from our exploration of a deeper brain region called the central complex. Studies across a range of insects, including locusts, cockroaches and Drosophila, suggest that the central complex is an adaptive sensorimotor processing center, with a role in sensory-guided orientation, navigation, place-learning and motor coordination. We are using a combination of experimental and computational techniques in an effort to characterize sensorimotor representations and circuit dynamics in the region in the context of behavior. Keywords: Behavior, Drosophila, gain modulation, motor, sensory, state dependence Conference: Tenth International Congress of Neuroethology, College Park. Maryland USA, United States, 5 Aug - 10 Aug, 2012. Presentation Type: Invited Symposium (only for people who have been invited to a particular symposium) Topic: Sensorimotor Integration Citation: Jayaraman V (2012). Linking vision and action in Drosophila. Conference Abstract: Tenth International Congress of Neuroethology. doi: 10.3389/conf.fnbeh.2012.27.00214 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 29 Apr 2012; Published Online: 07 Jul 2012. * Correspondence: Dr. Vivek Jayaraman, Janelia Farm Research Campus, Howard Hughes Medical Institute, Ashburn, VA, 20147, United States, vivek@janelia.hhmi.org Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Vivek Jayaraman Google Vivek Jayaraman Google Scholar Vivek Jayaraman PubMed Vivek Jayaraman Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Pattern-dependent response modulations in motion-sensitive visual interneurons--a model study.

Even if a stimulus pattern moves at a constant velocity across the receptive field of motion-sensitive neurons, such as lobula plate tangential cells (LPTCs) of flies, the response amplitude modulates over time. The amplitude of these response modulations is related to local pattern properties of the moving retinal image. On the one hand, pattern-dependent response modulations have previously been interpreted as 'pattern-noise', because they deteriorate the neuron's ability to provide unambiguous velocity information. On the other hand, these modulations might also provide the system with valuable information about the textural properties of the environment. We analyzed the influence of the size and shape of receptive fields by simulations of four versions of LPTC models consisting of arrays of elementary motion detectors of the correlation type (EMDs). These models have previously been suggested to account for many aspects of LPTC response properties. Pattern-dependent response modulations decrease with an increasing number of EMDs included in the receptive field of the LPTC models, since spatial changes within the visual field are smoothed out by the summation of spatially displaced EMD responses. This effect depends on the shape of the receptive field, being the more pronounced - for a given total size - the more elongated the receptive field is along the direction of motion. Large elongated receptive fields improve the quality of velocity signals. However, if motion signals need to be localized the velocity coding is only poor but the signal provides – potentially useful – local pattern information. These modelling results suggest that motion vision by correlation type movement detectors is subject to uncertainty: you cannot obtain both an unambiguous and a localized velocity signal from the output of a single cell. Hence, the size and shape of receptive fields of motion sensitive neurons should be matched to their potential computational task.

Flight Activity Alters Velocity Tuning of Fly Motion-Sensitive Neurons

Sensory neurons are mostly studied in fixed animals, but their response properties might change when the animal is free to move. Indeed, recent studies found differences between responses of sensory neurons in resting versus moving insects. Since the dynamic range of visual motion stimuli strongly depends on the speed at which an animal is moving, we investigated whether the visual system adapts to such changes in stimulus dynamics as induced by self-motion. Lobula plate tangential cells of flies lend themselves well to study this question because they are known to code for ego-motion based on optic-flow. We recorded the responses of the lobula plate tangential cell H1 to a visual pattern moving at different velocities under three different conditions: fixed flies before and after application of the octopamine agonist chlordimeform (CDM) and tethered flying flies. CDM has been previously shown to induce arousal in flies. We found that flying as well as the application of CDM significantly broadens the velocity tuning of H1 toward higher velocities.

Neural correlates of illusory motion perception in Drosophila

When the contrast of an image flickers as it moves, humans perceive an illusory reversal in the direction of motion. This classic illusion, called reverse-phi motion, has been well-characterized using psychophysics, and several models have been proposed to account for its effects. Here, we show that Drosophila melanogaster also respond behaviorally to the reverse-phi illusion and that the illusion is present in dendritic calcium signals of motion-sensitive neurons in the fly lobula plate. These results closely match the predictions of the predominant model of fly motion detection. However, high flicker rates cause an inversion of the reverse-phi behavioral response that is also present in calcium signals of lobula plate tangential cell dendrites but not predicted by the model. The fly's behavioral and neural responses to the reverse-phi illusion reveal unexpected interactions between motion and flicker signals in the fly visual system and suggest that a similar correlation-based mechanism underlies visual motion detection across the animal kingdom.

Candidate Glutamatergic Neurons in the Visual System of Drosophila

The visual system of Drosophila contains approximately 60,000 neurons that are organized in parallel, retinotopically arranged columns. A large number of these neurons have been characterized in great anatomical detail. However, studies providing direct evidence for synaptic signaling and the neurotransmitter used by individual neurons are relatively sparse. Here we present a first layout of neurons in the Drosophila visual system that likely release glutamate as their major neurotransmitter. We identified 33 different types of neurons of the lamina, medulla, lobula and lobula plate. Based on the previous Golgi-staining analysis, the identified neurons are further classified into 16 major subgroups representing lamina monopolar (L), transmedullary (Tm), transmedullary Y (TmY), Y, medulla intrinsic (Mi, Mt, Pm, Dm, Mi Am), bushy T (T), translobula plate (Tlp), lobula intrinsic (Lcn, Lt, Li), lobula plate tangential (LPTCs) and lobula plate intrinsic (LPi) cell types. In addition, we found 11 cell types that were not described by the previous Golgi analysis. This classification of candidate glutamatergic neurons fosters the future neurogenetic dissection of information processing in circuits of the fly visual system.

Neural Action Fields for Optic Flow Based Navigation: A Simulation Study of the Fly Lobula Plate Network

Optic flow based navigation is a fundamental way of visual course control described in many different species including man. In the fly, an essential part of optic flow analysis is performed in the lobula plate, a retinotopic map of motion in the environment. There, the so-called lobula plate tangential cells possess large receptive fields with different preferred directions in different parts of the visual field. Previous studies demonstrated an extensive connectivity between different tangential cells, providing, in principle, the structural basis for their large and complex receptive fields. We present a network simulation of the tangential cells, comprising most of the neurons studied so far (22 on each hemisphere) with all the known connectivity between them. On their dendrite, model neurons receive input from a retinotopic array of Reichardt-type motion detectors. Model neurons exhibit receptive fields much like their natural counterparts, demonstrating that the connectivity between the lobula plate tangential cells indeed can account for their complex receptive field structure. We describe the tuning of a model neuron to particular types of ego-motion (rotation as well as translation around/along a given body axis) by its ‘action field’. As we show for model neurons of the vertical system (VS-cells), each of them displays a different type of action field, i.e., responds maximally when the fly is rotating around a particular body axis. However, the tuning width of the rotational action fields is relatively broad, comparable to the one with dendritic input only. The additional intra-lobula-plate connectivity mainly reduces their translational action field amplitude, i.e., their sensitivity to translational movements along any body axis of the fly.

Recording from Two Neurons: Second-Order Stimulus Reconstruction from Spike Trains and Population Coding

We study the reconstruction of visual stimuli from spike trains, representing the reconstructed stimulus by a Volterra series up to second order. We illustrate this procedure in a prominent example of spiking neurons, recording simultaneously from the two H1 neurons located in the lobula plate of the fly Chrysomya megacephala. The fly views two types of stimuli, corresponding to rotational and translational displacements. Second-order reconstructions require the manipulation of potentially very large matrices, which obstructs the use of this approach when there are many neurons. We avoid the computation and inversion of these matrices using a convenient set of basis functions to expand our variables in. This requires approximating the spike train four-point functions by combinations of two-point functions similar to relations, which would be true for gaussian stochastic processes. In our test case, this approximation does not reduce the quality of the reconstruction. The overall contribution to stimulus reconstruction of the second-order kernels, measured by the mean squared error, is only about 5% of the first-order contribution. Yet at specific stimulus-dependent instants, the addition of second-order kernels represents up to 100% improvement, but only for rotational stimuli. We present a perturbative scheme to facilitate the application of our method to weakly correlated neurons.

Wide-Field Motion Integration in Fly VS Cells: Insights from an Inverse Approach

Fly lobula plate tangential cells are known to perform wide-field motion integration. It is assumed that the shape of these neurons, and in particular the shape of the subclass of VS cells, is responsible for this type of computation. We employed an inverse approach to investigate the morphology-function relationship underlying wide-field motion integration in VS cells. In the inverse approach detailed, model neurons are optimized to perform a predefined computation: here, wide-field motion integration. We embedded the model neurons to be optimized in a biologically plausible model of fly motion detection to provide realistic inputs, and subsequently optimized model neuron with and without active conductances (gNa, gK, gK(Na)) along their dendrites to perform this computation. We found that both passive and active optimized model neurons perform well as wide-field motion integrators. In addition, all optimized morphologies share the same blueprint as real VS cells. In addition, we also found a recurring blueprint for the distribution of gK and gNa in the active models. Moreover, we demonstrate how this morphology and distribution of conductances contribute to wide-field motion integration. As such, by using the inverse approach we can predict the still unknown distribution of gK and gNa and their role in motion integration in VS cells.

Control theoretic interpretation of directional motion preferences in optic flow processing interneurons

In this article, we formalize the processing of optic flow in identified fly lobula plate tangential cells and develop a control theoretic framework that suggests how the signals of these cells may be combined and used to achieve reflex-like navigation behavior. We show that this feedback gain synthesis task can be cast as a combined static state estimation and linear feedback control problem. Our framework allows us to analyze and determine the relationship between optic flow measurements and actuator commands, which greatly simplifies the implementation of biologically inspired control architectures on terrestrial and aerial robotic platforms.

Walking Modulates Speed Sensitivity in Drosophila Motion Vision

Changes in behavioral state modify neural activity in many systems. In some vertebrates such modulation has been observed and interpreted in the context of attention and sensorimotor coordinate transformations. Here we report state-dependent activity modulations during walking in a visual-motor pathway of Drosophila. We used two-photon imaging to monitor intracellular calcium activity in motion-sensitive lobula plate tangential cells (LPTCs) in head-fixed Drosophila walking on an air-supported ball. Cells of the horizontal system (HS)--a subgroup of LPTCs--showed stronger calcium transients in response to visual motion when flies were walking rather than resting. The amplified responses were also correlated with walking speed. Moreover, HS neurons showed a relatively higher gain in response strength at higher temporal frequencies, and their optimum temporal frequency was shifted toward higher motion speeds. Walking-dependent modulation of HS neurons in the Drosophila visual system may constitute a mechanism to facilitate processing of higher image speeds in behavioral contexts where these speeds of visual motion are relevant for course stabilization.

Two-photon calcium imaging from head-fixed Drosophila during optomotor walking behavior.

Drosophila melanogster is a model organism rich in genetic tools to manipulate and identify neural circuits involved in specific behaviors. Here we present a novel technique for two-photon calcium imaging in the central brain of head-fixed Drosophila walking on an air-supported ball. The ball’s motion is tracked at high resolution and can be treated as a proxy for the fly’s own movements. We used the genetically encoded calcium sensor, GCaMP3.0, to record from important elements of the motion-processing pathway, the horizontal-system (HS) lobula plate tangential cells (LPTCs) in the fly optic lobe. We presented motion stimuli to the tethered fly and found that calcium transients in HS-neurons correlated with robust optomotor behavior during walking. Our technique allows an entirely new set of questions to be addressed by monitoring behavior and physiology in identified neurons in a powerful genetic model organism with an extensive repertoire of walking behaviors.

Fly Motion Vision

Fly motion vision and resultant compensatory optomotor responses are a classic example for neural computation. Here we review our current understanding of processing of optic flow as generated by an animal's self-motion. Optic flow processing is accomplished in a series of steps: First, the time-varying photoreceptor signals are fed into a two-dimensional array of Reichardt-type elementary motion detectors (EMDs). EMDs compute, in parallel, local motion vectors at each sampling point in space. Second, the output signals of many EMDs are spatially integrated on the dendrites of large-field tangential cells in the lobula plate. In the third step, tangential cells form extensive interactions with each other, giving rise to their large and complex receptive fields. Thus, tangential cells can act as matched filters tuned to optic flow during particular flight maneuvers. They finally distribute their information onto postsynaptic descending neurons, which either instruct the motor centers of the thoracic ganglion for flight and locomotion control or act themselves as motor neurons that control neck muscles for head movements.

Localized direction selective responses in the dendrites of visual interneurons of the fly

BackgroundThe various tasks of visual systems, including course control, collision avoidance and the detection of small objects, require at the neuronal level the dendritic integration and subsequent processing of many spatially distributed visual motion inputs. While much is known about the pooled output in these systems, as in the medial superior temporal cortex of monkeys or in the lobula plate of the insect visual system, the motion tuning of the elements that provide the input has yet received little attention. In order to visualize the motion tuning of these inputs we examined the dendritic activation patterns of neurons that are selective for the characteristic patterns of wide-field motion, the lobula-plate tangential cells (LPTCs) of the blowfly. These neurons are known to sample direction-selective motion information from large parts of the visual field and combine these signals into axonal and dendro-dendritic outputs.ResultsFluorescence imaging of intracellular calcium concentration allowed us to take a direct look at the local dendritic activity and the resulting local preferred directions in LPTC dendrites during activation by wide-field motion in different directions. These 'calcium response fields' resembled a retinotopic dendritic map of local preferred directions in the receptive field, the layout of which is a distinguishing feature of different LPTCs.ConclusionsOur study reveals how neurons acquire selectivity for distinct visual motion patterns by dendritic integration of the local inputs with different preferred directions. With their spatial layout of directional responses, the dendrites of the LPTCs we investigated thus served as matched filters for wide-field motion patterns.

Sensor Fusion in Identified Visual Interneurons

Animal locomotion often depends upon stabilization reflexes that use sensory feedback to maintain trajectories and orientation. Such stabilizing reflexes are critically important for the blowfly, whose aerodynamic instability permits outstanding maneuverability but increases the demands placed on flight control. Flies use several sensory systems to drive reflex responses, and recent studies have provided access to the circuitry responsible for combining and employing these sensory inputs. We report that lobula plate VS neurons combine inputs from two optical sensors, the ocelli and the compound eyes. Both systems deliver essential information on in-flight rotations, but our neuronal recordings reveal that the ocelli encode this information in three axes, whereas the compound eyes encode in nine. The difference in dimensionality is reconciled by tuning each VS neuron to the ocellar axis closest to its compound eye axis. We suggest that this simple projection combines the speed of the ocelli with the accuracy of the compound eyes without compromising either. Our findings also support the suggestion that the coordinates of sensory information processing are aligned with axes controlling the natural modes of the fly's flight to improve the efficiency with which sensory signals are transformed into appropriate motor commands.

Processing of Horizontal Optic Flow in Three Visual Interneurons of theDrosophilaBrain

Motion vision is essential for navigating through the environment. Due to its genetic amenability, the fruit fly Drosophila has been serving for a lengthy period as a model organism for studying optomotor behavior as elicited by large-field horizontal motion. However, the neurons underlying the control of this behavior have not been studied in Drosophila so far. Here we report the first whole cell recordings from three cells of the horizontal system (HSN, HSE, and HSS) in the lobula plate of Drosophila. All three HS cells are tuned to large-field horizontal motion in a direction-selective way; they become excited by front-to-back motion and inhibited by back-to-front motion in the ipsilateral field of view. The response properties of HS cells such as contrast and velocity dependence are in accordance with the correlation-type model of motion detection. Neurobiotin injection suggests extensive coupling among ipsilateral HS cells and additional coupling to tangential cells that have their dendrites in the contralateral hemisphere of the brain. This connectivity scheme accounts for the complex layout of their receptive fields and explains their sensitivity both to ipsilateral and to contralateral motion. Thus the main response properties of Drosophila HS cells are strikingly similar to the responses of their counterparts in the blowfly Calliphora, although we found substantial differences with respect to their dendritic structure and connectivity. This long-awaited functional characterization of HS cells in Drosophila provides the basis for the future dissection of optomotor behavior and the underlying neural circuitry by combining genetics, physiology, and behavior.

Physiology in Drosophila motion-sensitive neurons during walking and flight behavior

Event Abstract Back to Event Physiology in Drosophila motion-sensitive neurons during walking and flight behavior Johannes D. Seelig1*, Eugenia M. Chiappe1, Gus K. Lott1, Michael B. Reiser2 and Vivek Jayaraman1 1 HHMI, Janelia Farm Research Campus, United States 2 Janelia Farm Research Campus , United States Drosophila melanogaster is a genetic model organism with many experimental advantages, including the ability to genetically manipulate specific sub-populations of neurons. Selected neurons in the fruit fly central brain can also be targeted for in vivo electrophysiology and two-photon calcium imaging. This powerful combination of physiology and genetic tools-such as cell-type-specific labeling, activity sensing, light-activation and silencing-is increasingly being applied to questions in systems neuroscience [1]. Our goal is to understand circuit computations underlying sensory-motor transformation in the fly brain. This requires recording not just neural activity but also the fly's behavior. Towards this end, we will present two novel experimental setups that we have developed: (i) Two-photon calcium imaging while the fly is walking on an air-supported 'Buchner' ball [2, 3] in a virtual arena [4]. (ii) Two-photon calcium imaging when the tethered fly is flying in a virtual arena. In order to assess the quality of behavior in our imaging preparation, we focused on a well-known behavior, the optomotor response. This turning response to visual motion can be induced by presenting a large field vertical grating moving horizontally in front of the fly. We show that tethered walking and flying flies reliably perform optomotor behavior during two-photon imaging. To assess the quality of the calcium imaging, we investigated calcium responses in lobula plate tangential cells (LPTCs). This small group of neurons has been exhaustively studied in fixed preparations in larger flies, and their responses to large-field stimuli (such as those that induce the optomotor response) have been carefully characterized [5]. In blowflies, LPTCs respond to large field motion stimuli with strong calcium responses, making these cells ideal candidates to test the capabilities of the behaving Drosophila preparation. Recording from Horizontal System (HS) LPTCs, we find reliable calcium responses in behaving fruit flies using a new genetically encoded calcium indicator, GCaMP3.0 [6]. HS neurons show strong (~150±/F) and stable responses to motion in their preferred direction. These calcium responses correlate with the fly's own walking optomotor response in that direction. Do responses of HS neurons change depending on the behavioral state of the animal? We will present results that compare responses in walking flies to those in flying flies. In our poster, we will describe the details of setups that allow us to perform stable recordings from identified motion-sensitive interneurons in the visual system of walking and flying Drosophila. These recordings represent the first physiological recordings in behaving Drosophila and provide a platform for future explorations of decision-making and sensory-motor transformations in this powerful genetic model organism.

Behavioural state affects motion-sensitive neurones in the fly visual system

The strength of stimulus-induced responses at the neuronal and the behavioural level often depends on the internal state of an animal. Within pathways processing sensory information and eventually controlling behavioural responses, such gain changes can originate at several sites. Using motion-sensitive lobula plate tangential cells (LPTCs) of blowflies, we address whether and in which way information processing changes for two different states of motor activity. We distinguish between the two states on the basis of haltere movements. Halteres are the evolutionarily transformed hindwings of flies. They oscillate when the animals walk or fly. LPTCs mediate, amongst other behaviours, head optomotor responses. These are either of large or small amplitude depending on the state of motor activity. Here we find that LPTC responses also depend on the motor activity of flies. In particular, LPTC responses are enhanced when halteres oscillate. Nevertheless, the response changes of LPTCs do not account for the corresponding large gain changes of head movements. Moreover, haltere activity itself does not change the activity of LPTCs. Instead, we propose that a central signal associated with motor activity changes the gain of head optomotor responses and the response properties of LPTCs.