Biomedical subjects

M Egelhaaf

Publications and source records attributed to M Egelhaaf.

At least 19 recordsLinked to original sources

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

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↗

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↗

PubMed1999

Variability in spike trains during constant and dynamic stimulation.

In a recent study, it was concluded that natural time-varying stimuli are represented more reliably in the brain than constant stimuli are. The results presented here disagree with this conclusion, although they were obtained from the same identified neuron (H1) in the fly's visual system. For large parts of the neuron's activity range, the variability of the responses was very similar for constant and time-varying stimuli and was considerably smaller than that in many visual interneurons of vertebrates.

Action Potentials↗

PubMed1999

Encoding of motion in real time by the fly visual system.

Direction-selective cells in the fly visual system that have large receptive fields play a decisive role in encoding the time-dependent optic flow the animal encounters during locomotion. Recent experiments on the computations performed by these cells have highlighted the significance of dendritic integration and have addressed the role of spikes versus graded membrane potential changes in encoding optic flow information. It is becoming increasingly clear that the way optic flow is encoded in real time is constrained both by the computational needs of the animal in visually guided behaviour as well as by the specific properties of the underlying neuronal hardware.

Animals↗

PubMed1999

In vivo calcium accumulation in presynaptic and postsynaptic dendrites of visual interneurons.

In this comparative in vivo study of dendritic calcium accumulation, we describe the time course and spatial integration properties of two classes of visual interneurons in the lobula plate of the blowfly. Calcium accumulation was measured during visual motion stimulation, ensuring synaptic activation of the neurons within their natural spatial and temporal operating range. The compared cell classes, centrifugal horizontal (CH) and horizontal system (HS) cells, are known to receive retinotopic input of similar direction selectivity, but to differ in morphology, biophysics, presence of dendrodendritic synapses, and computational task. 1) The time course of motion-induced calcium accumulation was highly invariant with respect to stimulus parameters such as pattern contrast and size. In HS cells, the rise of [Ca(2+)](i) can be described by a single exponential with a time constant of 5-6 s. The initial rise of [Ca(2+)](i) in CH cells was much faster (tau approximately 1 s). The decay time constant in both cell classes was estimated to be at least 3.5 times longer than the corresponding rise time constant. 2) The voltage-[Ca(2+)](i) relationship was best described by an expansive nonlinearity in HS cells and an approximately linear relationship in CH cells. 3) Both cell classes displayed a size-dependent saturation nonlinearity of the calcium accumulation. Although in CH cells calcium saturation was indistinguishable from saturation of the membrane potential, saturation of the two response parameters differed in HS cells. 4) There was spatial overlap of the calcium signal in response to nonoverlapping visual stimuli. Both the area and the amplitude of the overlap profile was larger in CH cells than in HS cells. Thus calcium accumulation in CH cells is spatially blurred to a greater extent than in HS cells. 5) The described differences between the two cell classes may reflect the following computational tasks of these neurons: CH cells relay retinotopic information within the lobula plate via dendritic synapses with pronounced spatial low-pass filtering. HS cells are output neurons of the lobula plate, in which the slow, local calcium accumulation may be suitable for local modulatory functions.

Animals↗

PubMed1999

Temperature-dependence of neuronal performance in the motion pathway of the blowfly calliphora erythrocephala

Raising the head temperature within a behaviourally relevant range has strong effects on the performance of an identified neuron, the H1 neuron, in the visual motion pathway of blowflies. The effect is seen as an increase in the mean amplitude of the responses to motion under both transient and steady-state conditions, a considerable decrease in the response latency and an improvement in the reliability of the responses to motion. These temperature-dependent effects are independent of whether the animal is exposed to transient temperature changes or is maintained continuously at the same temperature for its entire life. The changes in the neuronal response properties with temperature may be of immediate functional significance for the animal under its normal operating conditions. In particular, the decrease in latency and the improvement in the reliability with increasing temperature may be relevant for the fly when executing its extremely virtuosic flight manoeuvres.

Journal Article↗

PubMed1998

Temporal precision of the encoding of motion information by visual interneurons.

BACKGROUND: There is much controversy about the timescale on which neurons process and transmit information. On the one hand, a vast amount of information can be processed by the nervous system if the precise timing of individual spikes on a millisecond timescale is important. On the other hand, neuronal responses to identical stimuli often vary considerably and stochastic response fluctuations can exceed the mean response amplitude. Here, we examined the timescale on which neural responses could be locked to visual motion stimuli. RESULTS: Spikes of motion-sensitive neurons in the visual system of the blowfly are time-locked to visual motion with a precision in the range of several tens of milliseconds. Nevertheless, different motion-sensitive neurons with largely overlapping receptive fields generate a large proportion of spikes almost synchronously. This precision is brought about by stochastic rather than by motion-induced membrane-potential fluctuations elicited by the common peripheral input. The stochastic membrane-potential fluctuations contain more power at frequencies above 30-40 Hz than the motion-induced potential changes. A model of spike generation indicates that such fast membrane-potential changes are a major determinant of the precise timing of spikes. CONCLUSIONS: The timing of spikes in neurons of the motion pathway of the blowfly is controlled on a millisecond timescale by fast membrane-potential fluctuations. Despite this precision, spikes do not lock to motion stimuli on this timescale because visual motion does not induce sufficiently rapid changes in the membrane potential.

Action Potentials↗

PubMed1997

Synapse distribution on VCH, an inhibitory, motion-sensitive interneuron in the fly visual system.

In this study, the distribution of synapses in the ventral centrifugal horizontal (VCH)-a nonspiking, inhibitory, motion-sensitive interneuron in the third visual ganglion (lobula plate) of the blowfly Calliphora erythrocephala-was examined by electron microscopy and electrophysiology. The frequency histograms of excitatory and inhibitory postsynaptic potential amplitudes recorded from the VCH during contralateral stimulus presentation suggest the existence of three neurons, two excitatory and one inhibitory, mediating contralateral input. To localize input and output regions of the VCH, we investigated the ultrastructure of its two arborisation areas after intracellular iontophoretic injection of horseradish peroxidase. The VCH has input synapses in its arborisation in the protocerebrum and in its arborisation in the lobula plate. Output synapses were found exclusively in the lobula plate. Thus, the large dendritic arbor of the VCH in the lobula plate serves simultaneously as an input and an output region. There, we found input and output synapses in close vicinity (0.5-1.5 microm) to each other. Taking into account that the VCH receives retinotopicly arranged input from the ipsilateral eye in the lobula plate, the close location of input and output synapses in the VCH suggests that the spatial organization of its retinotopic synaptic input is more or less conserved in its inhibitory output pattern. The VCH has been proposed to inhibit the figure detection 1 (FD1), another neuron of the lobula plate, that responds preferentially to small moving objects. These results suggest that the FD1 may receive inhibitory inputs from the VCH in the lobula plate, where the dendritic arbors of both neurons overlap.

Animals↗