Biomedical subjects

M Egelhaaf

Publications and source records attributed to M Egelhaaf.

At least 37 records · Page 2Linked to original sources

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

Speed tuning in elementary motion detectors of the correlation type.

A prominent model of visual motion detection is the so-called correlation or Reichardt detector. Whereas this model can account for many properties of motion vision, from humans to insects (review, Borst and Egelhaaf 1989), it has been commonly assumed that this scheme of motion detection is not well suited to the measurement of image velocity. This is because the commonly used version of the model, which incorporates two unidirectional motion detectors with opposite preferred directions, produces a response which varies not only with the velocity of the image, but also with its spatial structure and contrast. On the other hand, information on image velocity can be crucial in various contexts, and a number of recent behavioural experiments suggest that insects do extract velocity for navigational purposes (review, Srinivasan et al. 1996). Here we show that other versions of the correlation model, which consists of a single unidirectional motion detector or incorporates two oppositely directed detectors with unequal sensitivities, produce responses which vary with image speed and display tuning curves that are substantially independent of the spatial structure of the image. This surprising feature suggests simple strategies of reducing ambiguities in the estimation of speed by using components of neural hardware that are already known to exist in the visual system.

Models, Theoretical↗

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↗

PubMed1997

Edge detection by landing honeybees: behavioural analysis and model simulations of the underlying mechanism.

The mechanism of edge detection in the honeybee was investigated by examining the effects of combining different kinds of visual cues that define an edge. Free-flying bees were trained to land at three different types of edges which were defined by texture and relative motion cues either in isolation or in combination with each other. Bees are able to detect and land at the three types of edges, but do so with different frequencies. In contrast to the naive expectation that edges jointly defined by two cues can be detected better than those defined by a single cue in isolation, the combination of the cues does not increase and may even decrease the detectability of an edge. When bees land at an edge the orientation of their body axis is strongly affected by the visual cues defining this edge. Model simulations were performed to test whether the experimental findings can be explained on the basis of a single edge detection mechanism sensitive to both types of visual cues. In the model, the information from both types of cues is sensed by two fields of movement detectors that receive their input signals from two adjacent patches in the visual field. The output of all detectors subserving either patch is pooled by integrating cells. The signals of the two integrating cells subserving the two adjacent patches are compared at a subtraction stage. The resulting signal is then rectified and forms the output signal of the model. The model simulations closely resemble the experimental results, thus providing evidence that edge detection by the bee could be mediated by a single mechanism.

Animals↗

PubMed1997

How reliably does a neuron in the visual motion pathway of the fly encode behaviourally relevant information?

How reliably neurons convey information depends on the extent to which their activity is affected by stochastic processes which are omnipresent in the nervous system. The functional consequences of neuronal noise can only be assessed if the latter is related to the response components that are induced in a normal behavioural situation. In the present study the reliability of neural coding was investigated for an identified neuron in the pathway processing visual motion information of the fly (Lucilia cuprina). The stimuli used to investigate the neuronal performance were not exclusively defined by the experimenter. Instead, they were generated by the fly itself, i.e. by its own actions and reactions in a behavioural closed-loop experiment, and subsequently replayed to the animal while the activity of an identified motion-sensitive neuron was recorded. Although the time course of the neuronal responses is time-locked to the stimulus, individual response traces differ slightly from each other due to stochastic fluctuations in the timing and number of action potentials. Individual responses thus consist of a stimulus-induced and a stochastic response component. The stimulus-induced response component can be recovered most reliably from noisy neuronal signals if these are smoothed by intermediate-sized time windows (40-100 ms). At this time scale the best compromise is achieved between smoothing out the noise and maintaining the temporal resolution of the stimulus-induced response component. Consequently, in the visual motion pathway of the fly, behaviourally relevant motion stimuli can be resolved best at a time scale where the timing of individual spikes does not matter.

Animals↗

PubMed1995

Mechanisms of dendritic integration underlying gain control in fly motion-sensitive interneurons.

In the compensatory optomotor response of the fly the interesting phenomenon of gain control has been observed by Reichardt and colleagues (Reichardt et al., 1983): The amplitude of the response tends to saturate with increasing stimulus size, but different saturation plateaus are assumed with different velocities at which the stimulus is moving. This characteristic can already be found in the motion-sensitive large field neurons of the fly optic lobes that play a role in mediating this behavioral response (Hausen, 1982; Reichardt et al., 1983; Egelhaaf, 1985; Haag et al., 1992). To account for gain control a model was proposed involving shunting inhibition of these cells by another cell, the so-called pool cell (Reichardt et al., 1983), both cells sharing common input from an array of local motion detectors. This article describes an alternative model which only requires dendritic integration of the output signals of two types of local motion detectors with opposite polarity. The explanation of gain control relies on recent findings that these input elements are not perfectly directionally selective and that their direction selectivity is a function of pattern velocity. As a consequence, the resulting postsynaptic potential in the dendrite of the integrating cell saturates with increasing pattern size at a level between the excitatory and inhibitory reversal potentials. The exact value of saturation is then set by the activation ratio of excitatory and inhibitory input elements which in turn is a function of other stimulus parameters such as pattern velocity. Thus, the apparently complex phenomenon of gain control can be simply explained by the biophysics of dendritic integration in conjunction with the properties of the motion-sensitive input elements.

Animals↗

PubMed1995

A preparation of the blowfly (Calliphora erythrocephala) brain for in vitro electrophysiological and pharmacological studies.

We describe a method for the preparation and maintenance of the blowfly (Calliphora erythrocephala) brain in a recording chamber under in vitro conditions in a semi-slice configuration. Large identification neurones in the posterior part of the 3rd optic lobe (lobula plate) can be penetrated easily with microelectrodes. The so-called vertical system (VS) cells which respond to vertical image motion in vivo could be encountered best because their axons are escorted individually by specific tracheae. Fluorescent stained cells show their natural shape as being in vivo. Electrophysiological properties of the cells investigated so far, i.e., resting potential (about -40 mV) and firing properties (single rebound spikes), are comparable to recordings in intact flies. Initial pharmacological experiments on VS cells in this preparation reveal that iontophoretical application of acetylcholine and carbamylcholine results in depolarization. VS cells also respond to bath-applied nicotine (1 microM) with a slow depolarization of their membrane potential in normal fly saline as well as in a Ca(2+)-free saline, suggesting direct cholinergic input via nicotinic receptors. The suitability of the preparation for a wide range of electrophysiological and pharmacological studies is discussed.

Acetylcholine↗

PubMed1995

Calcium accumulation in visual interneurons of the fly: stimulus dependence and relationship to membrane potential.

1. The large motion-sensitive tangential neurons in the fly third visual neuropil spatially pool the postsynaptic signals of many local elements. The changes in membrane potential and calcium concentration induced in these cells by visual motion are analyzed in vivo by simultaneous optical and intracellular voltage recording techniques. 2. Visual motion in the preferred direction leads to depolarization of the cell and to calcium accumulation mainly in the axon terminal, the soma, and the dendritic tree. During motion in the null direction, the cell hyperpolarizes and virtually no changes in calcium concentration can be observed. 3. Dendritic calcium accumulation is first restricted to those dendritic branches that are close to the sites of direct synaptic input. In other parts of the dendrite the calcium concentration increases more slowly and usually reaches only lower levels. 4. Calcium starts accumulating at the onset of motion. However, the calcium concentration reaches its final steady-state level much later than the corresponding membrane potential changes. Even if these are completely transient at high temporal frequencies of pattern motion, the calcium signal stays high until the stimulus pattern stops moving. 5. The amplitude of the calcium signal depends on the temporal frequency of pattern motion in a similar way as do the corresponding membrane potential changes. However, there exist differences that can be attributed to the different time courses of both signals. 6. Depolarization of the dendritic tree by current injection through a microelectrode leads to similar changes in calcium accumulation as does activation by synaptic input, suggesting that calcium enters the cell via voltage-dependent channels. The possible function of calcium channels for dendritic integration of synaptic input is discussed.

Animals↗