Biomedical subjects

M Egelhaaf

Publications and source records attributed to M Egelhaaf.

At least 19 recordsLinked to original sources

PubMed2003

Visually guided orientation in flies: case studies in computational neuroethology.

To understand the functioning of nervous systems and, in particular, how they control behaviour we must bridge many levels of complexity from molecules, cells and synapses to perception behaviour. Although experimental analysis is a precondition for understanding by nervous systems, it is in no way sufficient. The understanding is aided at all levels of complexity by modelling. Modelling proved to be an inevitable tool to test the experimentally established hypotheses. In this review it will by exemplified by three case studies that the appropriate level of modelling needs to be adjusted to the particular computational problems that are to be solved. (1) Specific features of the highly virtuosic pursuit behaviour of male flies can be understood on the basis of a phenomenological model that relates the visual input to the motor output. (2) The processing of retinal image motion as is experienced by freely moving animals can be understood on the basis of a model consisting of algorithmic components and components which represent a simple equivalent circuit of nerve cells. (3) Behaviourally relevant features of the reliability of encoding of visual motion information can be understood by modelling the transformation of postsynaptic potentials into sequences of spike trains.

Animals↗

PubMed2003

FliMax, a novel stimulus device for panoramic and highspeed presentation of behaviourally generated optic flow.

A high-speed panoramic visual stimulation device is introduced which is suitable to analyse visual interneurons during stimulation with rapid image displacements as experienced by fast moving animals. The responses of an identified motion sensitive neuron in the visual system of the blowfly to behaviourally generated image sequences are very complex and hard to predict from the established input circuitry of the neuron. This finding suggests that the computational significance of visual interneurons can only be assessed if they are characterised not only by conventional stimuli as are often used for systems analysis, but also by behaviourally relevant input.

Animals↗

PubMed2003

Synaptic transfer of dynamic motion information between identified neurons in the visual system of the blowfly.

Synaptic transmission is usually studied in vitro with electrical stimulation replacing the natural input of the system. In contrast, we analyzed in vivo transfer of visual motion information from graded-potential presynaptic to spiking postsynaptic neurons in the fly. Motion in the null direction leads to hyperpolarization of the presynaptic neuron but does not much influence the postsynaptic cell, because its firing rate is already low during rest, giving only little scope for further reductions. In contrast, preferred-direction motion leads to presynaptic depolarizations and increases the postsynaptic spike rate. Signal transfer to the postsynaptic cell is linear and reliable for presynaptic graded membrane potential fluctuations of up to approximately 10 Hz. This frequency range covers the dynamic range of velocities that is encoded with a high gain by visual motion-sensitive neurons. Hence, information about preferred-direction motion is transmitted largely undistorted ensuring a consistent dependency of neuronal signals on stimulus parameters, such as motion velocity. Postsynaptic spikes are often elicited by rapid presynaptic spike-like depolarizations which superimpose the graded membrane potential. Although the timing of most of these spike-like depolarizations is set by noise and not by the motion stimulus, it is preserved at the synapse with millisecond precision.

Action Potentials↗

PubMed2001

Transfer of visual motion information via graded synapses operates linearly in the natural activity range.

Synaptic transmission between a graded potential neuron and a spiking neuron was investigated in vivo using sensory stimulation instead of artificial excitation of the presynaptic neuron. During visual motion stimulation, individual presynaptic and postsynaptic neurons in the brain of the fly were electrophysiologically recorded together with concentration changes of presynaptic calcium (Delta[Ca(2+)](pre)). Preferred-direction motion leads to depolarization of the presynaptic neuron. It also produces pronounced increases in [Ca(2+)](pre) and the postsynaptic spike rate. Motion in the opposite direction was associated with hyperpolarization of the presynaptic cell but only a weak reduction in [Ca(2+)](pre) and the postsynaptic spike rate. Apart from this rectification, the relationships between presynaptic depolarizations, Delta[Ca(2+)](pre), and postsynaptic spike rates are, on average, linear over the entire range of activity levels that can be elicited by sensory stimulation. Thus, the inevitably limited range in which the gain of overall synaptic signal transfer is constant appears to be adjusted to sensory input strengths.

Action Potentials↗

PubMed2001

Neural processing of naturalistic optic flow.

Stimuli traditionally used for analyzing visual information processing are much simpler than what an animal sees in normal life. When characterized with traditional stimuli, neuronal responses were found to depend on various parameters such as contrast, texture, or velocity of motion, and thus were highly ambiguous. In behavioral situations, all of these parameters change simultaneously and differently in different parts of the visual field. Thus it is hardly possible to predict from traditional analyses what information is encoded by neurons in behavioral situations. Therefore, we characterized an identified neuron in the optomotor system of the blowfly with image sequences as they were seen by animals walking in a structured environment. We conclude that during walking, the response of the neuron reflects the animal's turning direction nearly independently of the texture and spatial layout of the environment. Our findings stress the significance of analyzing the performance of neuronal circuits under their natural operating conditions.

Animals↗

PubMed2001

Two classes of visual motion sensitive interneurons differ in direction and velocity dependency of in vivo calcium dynamics.

Neurons exploit both membrane biophysics and biochemical pathways of the cytoplasm for dendritic integration of synaptic input. Here we quantify the tuning discrepancy of electrical and chemical response properties in two kinds of neurons using in vivo visual stimulation. Dendritic calcium concentration changes and membrane potential of visual interneurons of the fly were measured in response to visual motion stimuli. Two classes of tangential cells of the lobula plate were compared, HS-cells and CH-cells. Both neuronal classes are known to receive retinotopic input with similar properties, yet they differ in morphology, physiology, and computational context. Velocity tuning and directional selectivity of the electrical and calcium responses were investigated. In both cell classes, motion-induced calcium accumulation did not follow the early transient of the membrane potential. Rather, the amplitude of the calcium signal seemed to be related to the late component of the depolarization, where it was close to a steady state. Electrical and calcium responses differed with respect to their velocity tuning in CH-cells, but not in HS-cells. Furthermore, velocity tuning of the calcium response, but not of the electrical response differed between neuronal classes. While null-direction motion caused hyperpolarization in both classes, this led to a calcium decrement in CH-cells, but had no effect on the calcium signal in HS-cells, not even when calcium levels had been raised by a preceding excitatory motion stimulus. Finally, the voltage-[Ca2+]i-relationship for motion-induced, transient potential changes was steeper and less rectifying in CH-cells than in HS-cells. These results represent an example of dendritic information processing in vivo, where two neuronal classes respond to identical stimuli with a similar electrical response, but differing calcium response. This highlights the capacity of neurons to segregate two response components.

Animals↗

PubMed2001

Outdoor performance of a motion-sensitive neuron in the blowfly.

We studied an identified motion-sensitive neuron of the blowfly under outdoor conditions. The neuron was stimulated by oscillating the fly in a rural environment. We analysed whether the motion-induced neuronal activity is affected by brightness changes ranging between bright sunlight and dusk. In addition, the relationship between spike rate and ambient temperature was determined. The main results are: (1) The mean spike rate elicited by visual motion is largely independent of brightness changes over several orders of magnitude as they occur as a consequence of positional changes of the sun. Even during dusk the neuron responds strongly and directionally selective to motion. (2) The neuronal spike rate is not significantly affected by short-term brightness changes caused by clouds temporarily occluding the sun. (3) In contrast, the neuronal activity is much affected by changes in ambient temperature.

Adaptation, Ocular↗

PubMed2001

Binocular contributions to optic flow processing in the fly visual system.

Integrating binocular motion information tunes wide-field direction-selective neurons in the fly optic lobe to respond preferentially to specific optic flow fields. This is shown by measuring the local preferred directions (LPDs) and local motion sensitivities (LMSs) at many positions within the receptive fields of three types of anatomically identifiable lobula plate tangential neurons: the three horizontal system (HS) neurons, the two centrifugal horizontal (CH) neurons, and three heterolateral connecting elements. The latter impart to two of the HS and to both CH neurons a sensitivity to motion from the contralateral visual field. Thus in two HS neurons and both CH neurons, the response field comprises part of the ipsi- and contralateral visual hemispheres. The distributions of LPDs within the binocular response fields of each neuron show marked similarities to the optic flow fields created by particular types of self-movements of the fly. Based on the characteristic distributions of local preferred directions and motion sensitivities within the response fields, the functional role of the respective neurons in the context of behaviorally relevant processing of visual wide-field motion is discussed.

Animals↗

PubMed2001

Neuronal processing of behaviourally generated optic flow: experiments and model simulations.

The stimuli traditionally used for analysing visual information processing are much simpler than what an animal sees when moving in its natural environment. Therefore, we analysed in a previous study the performance of an identified neuron in the optomotor system of the fly by using as visual stimuli image sequences that were experienced by the animal while walking in a structured environment. These electrophysiological experiments revealed that the fly visual system computes from behaviourally generated optic flow a rather unambiguous representation of the animal's self-motion. In contrast to conclusions based on simple stimuli, the directions of turns are represented by an interneuron, the HSE cell, quite independent of the spatial layout of the environment and its textural properties when the cell is stimulated with behaviourally generated optic flow. This conclusion is substantiated here by further experimental evidence. Moreover, it is shown that the largely unambiguous responses of the HSE cell to behaviourally generated optic flow can be replicated to a large extent by a network model of the fly's visual motion pathway. These results stress the significance of naturalistic stimuli for analysing what is encoded by neuronal circuits under natural operating conditions.

Algorithms↗

PubMed2000

Reliability of a fly motion-sensitive neuron depends on stimulus parameters.

The variability of responses of sensory neurons constrains how reliably animals can respond to stimuli in the outside world. We show for a motion-sensitive visual interneuron of the fly that the variability of spike trains depends on the properties of the motion stimulus, although differently for different stimulus parameters. (1) The spike count variances of responses to constant and to dynamic stimuli lie in the same range. (2) With increasing stimulus size, the variance may slightly decrease. (3) Increasing pattern contrast reduces the variance considerably. For all stimulus conditions, the spike count variance is much smaller than the mean spike count and does not depend much on the mean activity apart from very low activities. Using a model of spike generation, we analyzed how the spike count variance depends on the membrane potential noise and the deterministic membrane potential fluctuations at the spike initiation zone of the neuron. In a physiologically plausible range, the variance is affected only weakly by changes in the dynamics or the amplitude of the deterministic membrane potential fluctuations. In contrast, the amplitude and dynamics of the membrane potential noise strongly influence the spike count variance. The membrane potential noise underlying the variability of the spike responses in the motion-sensitive neuron is concluded to be affected considerably by the contrast of the stimulus but by neither its dynamics nor its size.

Action Potentials↗

PubMed2000

Performance of fly visual interneurons during object fixation.

Neurons involved in the processing of optic flow are usually analyzed using stimuli designed by the experimenter. However, in real life optic flow depends on locomotive behavior. We characterized the performance of motion-sensitive neurons in the visual system of the fly using optic flow as occurring in behavioral situations during object fixation. Optic flow generated by tethered flying flies in a flight simulator was subsequently replayed while recording the responses of two cell types in the fly's motion pathway presumably involved in the detection of objects and of deviations from a straight flight course, respectively. FD1b cells, which are representatives of the so-called figure-detection cells, responded very specifically to object motion. Although object selectivity of these cells is attributable to inhibition during large-field motion, the influence of background motion during object fixation was almost negligible. In contrast, the cells of the so-called horizontal system (HS cells) are most sensitive to background motion, as elicited during deviations of the animal from its course. During object fixation, the responses of HS cells depended on both object and background motion. The simulated distance of the background to the fly did not have a strong influence on the responses of either cell type. The specificity for detecting deviations from a straight course is enhanced by subtraction of the signals of HS cells in both halves of the brain. In contrast, the FD1b cells in the two halves of the brain need to interact in a nonlinear way to ensure efficient detection of objects.

Action Potentials↗

PubMed2000

Detection of object motion by a fly neuron during simulated flight.

Object detection on the basis of relative motion was investigated in the fly at the neuronal level. A representative of the figure detection cells (FD-cells), the FD1b-cell, was characterized with respect to its responses to optic flow which simulated the presence of an object during translatory flight. The figure detection cells reside in the fly's third visual neuropil and are believed to play a central role in mediating object-directed turning behaviour. The dynamical response properties as well as the mean response amplitudes of the FD1b-cell depend on the temporal frequency of object motion and on the presence or absence of background motion. The responses of the FD1b-cell to object motion during simulated translatory flight were compared to behavioural responses of the fly as obtained with identical stimuli in a previous study. The behavioural responses could only partly be explained on the basis of the FD1b-cell's responses. Further processing between the third visual neuropil and the final motor output has to be assumed which involves (1) facilitation of the object-induced responses during translatory background motion at moderate temporal frequencies, and (2) inhibition of the object-induced turning responses during translatory background motion at high temporal frequencies.

Animals↗

PubMed2000

Optomotor course control in flies with largely asymmetric visual input.

We have studied freely flying and walking flies as well as flies flying in a flight simulator in order to discover how functionally blinding one of the eyes affects the fly's ability to move straight. It is hard to tell just by observing the animals' movements whether they have been deprived of vision in one eye. Statistical analysis is need to show that there are differences in the locomotory paths of monocular and binocular flies: monocular flies tend to turn slightly towards the side of the seeing eye. It is possible that the superimposed translational and rotational optic flow fields, generated on the trajectory of monocular flies, sum to zero net flow. This overall flow over the retina of the open eye might lead to a state of optomotor equilibrium.

Animals↗

PubMed2000

Neuronal representation of optic flow experienced by unilaterally blinded flies on their mean walking trajectories.

Asymmetries in the optic flow on both eyes may indicate an unintended turn of an animal and evoke compensatory optomotor responses. On a straight path in an evenly structured environment, the optic flow on both eyes is balanced corresponding to a state of optomotor equilibrium. When one eye is occluded an optomotor equilibrium is expected to be reached on a curved path provided that the translatory optic flow component is cancelled by a superimposed rotation. This hypothesis is tested by analysing how the HSE cell, a constituent element of the fly's optomotor system, represents optic flow in behavioural situations. The optic flow as seen on the average trajectory of freely walking monocular flies is reconstructed. This optic flow is used as stimulus of the HSE cell in electrophysiological experiments and as input of a model of the fly's optomotor system. The responses of the HSE cell and of the model fluctuate around the resting potential. On average, they are much smaller than the responses evoked by optic flow experienced on a straight path. These results corroborate the hypothesis that the mean trajectory of monocular flies corresponds to a path of optomotor equilibrium.

Animals↗

PubMed2000

Response latency of a motion-sensitive neuron in the fly visual system: dependence on stimulus parameters and physiological conditions.

The response latency of an identified motion-sensitive neuron in the blowfly visual system strongly depends on stimulus parameters. The latency decreases with increasing contrast and temporal frequency of a moving pattern, but changes only little when the pattern size and thus the number of activated inputs is increased. The latency does not only depend on visual stimuli, but is also affected by temperature changes and the age of the fly. Since response latencies cover a range of one order of magnitude, the latency changes are expected to be of relevance in visually guided orientation behaviour.

Animals↗

PubMed2000

Synaptic interactions increase optic flow specificity.

Representations of optic flow are encoded in fly tangential neurons by pooling the signals of many retinotopically organized local motion-sensitive inputs as well as of other tangential cells originating in the ipsi- and contralateral half of the brain. In the so called HSE cell, a neuron involved in optomotor course control, two contralateral input elements, the H1 and H2 cells, mediate distinct EPSPs. These EPSPs frequently elicit spike-like depolarizations in the HSE cell. The synaptic transmission between the H2 and the HSE cell is analysed in detail and shown to be very reliable with respect to the amplitude and time-course of the postsynaptic potential. As a consequence of its synaptic input, the HSE cell responds best to wide-field motion, such as that generated on the eyes when the animal turns about its vertical body axis. It is shown that the specificity of the HSE cell for this type of optic flow is much enhanced if rapid membrane depolarizations, such as large-amplitude EPSPs or spike-like depolarizations, are taken into account rather than the average membrane potential.

Action Potentials↗

PubMed2000

Dendritic calcium accumulation associated with direction-selective adaptation in visual motion-sensitive neurons in vivo.

Motion adaptation in directionally selective tangential cells (TC) of the fly visual system has previously been explained as a presynaptic mechanism. Based on the observation that adaptation is in part direction selective, which is not accounted for by the former models of motion adaptation, we investigated whether physiological changes located in the TC dendrite can contribute to motion adaptation. Visual motion in the neuron's preferred direction (PD) induced stronger adaptation than motion in the opposite direction and was followed by an afterhyperpolarization (AHP). The AHP subsides in the same time as adaptation recovers. By combining in vivo calcium fluorescence imaging with intracellular recording, we show that dendritic calcium accumulation following motion in the PD is correlated with the AHP. These results are consistent with a calcium-dependent physiological change in TCs underlying adaptation during continuous stimulation with PD motion, expressing itself as an AHP after the stimulus stops. However, direction selectivity of adaptation is probably not solely related to a calcium-dependent mechanism because direction-selective effects can also be observed for fast moving stimuli, which do not induce sizeable calcium accumulation. In addition, a comparison of two classes of TCs revealed differences in the relationship of calcium accumulation and AHP when the stimulus velocity was varied. Thus the potential role of calcium in motion adaptation depends on stimulation parameters and cell class.

Adaptation, Physiological↗

PubMed2000

Object fixation by the blowfly during tethered flight in a simulated three-dimensional environment.

The ability of flies to detect and fixate objects moving relative to their background was investigated in a flight simulator during translational tethered flight. The fly experienced optic flow that depended on its own actions and reactions in a similar way as in free-flight (closed-loop) conditions. Fixation of an object required turning responses towards it. The simulated distances between the fly, object and background were varied systematically by changing the velocities with which the object and the background pattern moved from the frontal to the back part of the fly's visual field. Fixation responses were only elicited when the object was simulated to be closer than the background. The fly's fixation performance was better with close than with more distant objects. Since, under many stimulus conditions, fixation responses were either elicited or entirely failed to be elicited, it is concluded that object fixation behaviour is gated in the visuo-motor pathway.

Animals↗