Association of visual objects and olfactory cues in Drosophila.

Context-dependent preferences in a choice between an upper and a lower visual object of otherwise identical appearance were recorded during stationary flight of the fruitfly, Drosophila melanogaster, in a flight simulator. The test animal was held in a fixed orientation at the center of a wing-beat processor that converts attempted turns into counter-rotations of a surrounding cylindrical panorama. This allowed the fly to maneuver the preferred object into the actual direction of flight. Single flies were trained to avoid a course toward the visual object that had been associated with the aversive odor benzaldehyde (BAL). Conditioned object avoidance was investigated in different treatment groups by collective evaluation of the scores from 80 long-lasting flights (> 1 hr). In addition to a significant cross-modal association, we found a striking long-term effect of transient exposure to BAL both in the embryonic and larval states. The preimaginal experience significantly increased the indifference to BAL in the adult flies. Disturbed vision does not account for this effect: Neither the perception nor the discrimination of the visual objects was significantly impaired in the investigated flies. Disturbed olfaction could explain the present results. Recently, however, preimaginal BAL uptake has been found to interfere directly with the retention of heat-shock-conditioned object avoidance.

Processing of artificial visual feedback in the walking fruit fly Drosophila melanogaster.

A computerized 360 degrees panorama allowed us to suppress most of the locomotion-induced visual feedback of a freely walking fly without neutralizing its mechanosensory system ('virtual open-loop' conditions). This novel paradigm achieves control over the fly's visual input by continuously evaluating its actual position and orientation. In experiments with natural visual feedback (closed-loop conditions), the optomotor turning induced by horizontal pattern motion in freely walking Drosophila melanogaster increased with the contrast and brightness of the stimulus. Conspicuously striped patterns were followed with variable speed but often without significant overall slippage. Using standard open-loop conditions in stationary walking flies and virtual open-loop or closed-loop conditions in freely walking flies, we compared horizontal turning induced by either horizontal or vertical motion of appropriately oriented rhombic figures. We found (i) that horizontal displacements and the horizontal-motion illusion induced by vertical displacements of the oblique edges of the rhombic figures elicited equivalent open-loop turning responses; (ii) that locomotion-induced visual feedback from the vertical edges of the rhombic figures in a stationary horizontal position diminished the closed-loop turning elicited by vertical displacements to only one-fifth of the response to horizontal displacements; and (iii) that virtual open-loop responses of mobile flies and open-loop responses of immobilized flies were equivalent in spite of delays of up to 0.1 s in the generation of the virtual stimulus. Horizontal compensatory turning upon vertical displacements of oblique edges is quantitatively consistent with the direction-selective summation of signals from an array of elementary motion detectors for the horizontal stimulus components within their narrow receptive fields. A compensation of the aperture-induced ambiguity can be excluded under these conditions. However, locomotion-induced visual feedback greatly diminished the horizontal-motion illusion in a freely walking fly. The illusion was used to assay the quality of open-loop simulation in the new paradigm.

The wake dynamics and flight forces of the fruit fly Drosophila melanogaster.

We have used flow visualizations and instantaneous force measurements of tethered fruit flies (Drosophila melanogaster) to study the dynamics of force generation during flight. During each complete stroke cycle, the flies generate one single vortex loop consisting of vorticity shed during the downstroke and ventral flip. This gross pattern of wake structure in Drosophila is similar to those described for hovering birds and some other insects. The wake structure differed from those previously described, however, in that the vortex filaments shed during ventral stroke reversal did not fuse to complete a circular ring, but rather attached temporarily to the body to complete an inverted heart-shaped vortex loop. The attached ventral filaments of the loop subsequently slide along the length of the body and eventually fuse at the tip of the abdomen. We found no evidence for the shedding of wing-tip vorticity during the upstroke, and argue that this is due to an extreme form of the Wagner effect acting at that time. The flow visualizations predicted that maximum flight forces would be generated during the downstroke and ventral reversal, with little or no force generated during the upstroke. The instantaneous force measurements using laser-interferometry verified the periodic nature of force generation. Within each stroke cycle, there was one plateau of high force generation followed by a period of low force, which roughly correlated with the upstroke and downstroke periods. However, the fluctuations in force lagged behind their expected occurrence within the wing-stroke cycle by approximately 1 ms or one-fifth of the complete stroke cycle. This temporal discrepancy exceeds the range of expected inaccuracies and artifacts in the measurements, and we tentatively discuss the potential retarding effects within the underlying fluid mechanics.

Activation phase ensures kinematic efficacy in flight-steering muscles of Drosophila melanogaster.

During tethered flight in Drosophila melanogaster, spike activity of the second basalar flight-control muscle (M.b2) is correlated with an increase in both the ipsilateral wing beat amplitude and the ipsilateral flight force. The frequency of muscle spikes within a burst is about 100 Hz, or 1 spike for every two wing beat cycles. When M.b2 is active, its spikes tend to occur within a comparatively narrow phase band of the wing beat cycle. To understand the functional role of this phase-lock of firing in the control of flight forces, we stimulated M.b2 in selected phases of the wing beat cycle and recorded the effect on the ipsilateral wing beat amplitude. Varying the phase timing of the stimulus had a significant effect on the wing beat amplitude. A maximum increase of wing beat amplitude was obtained by stimulating M.b2 at the beginning of the upstroke or about 1 ms prior to the narrow phase band in which the muscle spikes typically occur during flight. Assuming a delay of 1 ms between the stimulation of the motor nerve and muscle activation, these results indicate that M.b2 is activated at an instant of the stroke cycle that produces the greatest effect on wing beat amplitude.

Optomotor control of course and altitude in Drosophila melanogaster is correlated with distinct activities of at least three pairs of flight steering muscles.

Flight control in the fruitfly Drosophila melanogaster is achieved by minute sets of muscles on either side of the thorax. Control responses of wings and muscles were elicited during fixed flight by moving a striped pattern in front of the eyes. For example, pattern motion from the lower right to the upper left signals to the test fly a rotatory course deviation to the right and simultaneously a translatory altitude displacement downwards. The counteracting response to the displacement of the retinal image is an increase in thrust and lift on the right, accomplished mainly by increasing the wingbeat amplitude (WBA) on that side. A comparison of such responses with the simultaneously recorded action potentials in the prominent basalar muscles M.b1 and M.b2 and axillary muscles M.I1 and M.III1 on either side suggests that three of these muscles act on the WBA more or less independently and contribute to the optomotor control of course and altitude. During flight, M.b1 is almost continuously active with a frequency equal to or slightly below 1 spike per wingbeat cycle. The spikes occur within a narrow phase interval of this cycle, normally at the beginning of the transition from upstroke to downstroke. However, the visual stimulus described above causes a substantial phase lead in M.b1 on the right; the spikes occur shortly before the end of the upstroke. Such phase shifts are accompanied by comparatively smooth 'tonic' responses of the WBA. The activities of M.b2 and M.I1 are normally very low. However, the stimulus described above activates M.b2 on the right in a phase interval approximately two-thirds into the upstroke and M.I1 on the left in a phase interval at the beginning of the downstroke. The spikes tend to occur in bursts. These bursts are correlated with WBA-increasing 'hitches' (rapid changes in amplitude) on the right and WBA-decreasing hitches on the left. As fast 'phasic' responses, the burst-induced hitches are likely to account for the course-controlling 'body saccades' observed during free flight. For unknown reasons, M.I1 is activated by pattern motion but cannot conceivably assist the other muscles in altitude control. Unlike its homologues in larger flies (Musca domestica, Calliphora erythrocephala), M.III1 does not participate in optomotor flight control. Its activation seems to support the termination of flight and wing retraction at rest. The essential properties of the three pairs of muscles M.b1, M.b2 and M.I1 resemble those found in larger flies; the muscles are controlled by motion detectors with muscle-specific 'preferred directions' in the hexagonal array of retinal elements. Optomotor control of the three pairs of muscles in Drosophila melanogaster could explain most, but not all, of the WBA responses recorded so far.

The active control of wing rotation by Drosophila.

This paper investigates the temporal control of a fast wing rotation in flies, the ventral flip, which occurs during the transition from downstroke to upstroke. Tethered flying Drosophila actively modulate the timing of these rapid supinations during yaw responses evoked by an oscillating visual stimulus. The time difference between the two wings is controlled such that the wing on the outside of a fictive turn rotates in advance of its contralateral partner. This modulation of ventral-flip timing between the two wings is strongly coupled with changes in wing-stroke amplitude. Typically, an increase in the stroke amplitude of one wing is correlated with an advance in the timing of the ventral flip of the same wing. However, flies do display a limited ability to control these two behaviors independently, as shown by flight records in which the correlation between ventral-flip timing and stroke amplitude transiently reverses. The control of ventral-flip timing may be part of an unsteady aerodynamic mechanism that enables the fly to alter the magnitude and direction of flight forces during turning maneuvers.

Cortical templates for the self-organization of orientation-specific d- and l-hypercolumns in monkeys and cats.

Blasdel and Salama's sensory maps of orientation-selective edge detectors in the monkey striate cortex can be reduced to an idealized scheme in which orientation hypercolumns of the d- and l-type occur in alternating sequence (Fig. 1). This scheme resolves the apparent contradiction between linear and circular arrangements of successive edge directions in earlier accounts. The actual configuration of hypercolumns is in register with two possible templates for the self-organization of orientation selectivity: the isometric cytochrome oxidase blobs of the colour system, and the anisometric slabs of the ocular dominance system. The centers of the hypercolumns coincide with the blobs. Simulation of cortical self-organization shows this co-incidence even in the absence of template-specific interactions. However, blobs and slabs are symmetrical to these centers, and therefore no templates for the asymmetrical distribution of preferred orientation in the hypercolumns. The present simulation derives the pre-natal formation of an initial scheme from a hypothetical gradient of nervous activity. Post-natal formation, or maturation, of this scheme is achieved by visual experience. Simulation of corresponding interactions between simultaneously activated neurons illustrates both the gain in orientation selectivity (Figs. 2 and 3), and the optimization of farfield diversity and nearfield conformity (Figs. 4 and 5). The results are compatible with the actual distribution of blob-centered d- and l-hypercolumns, iso-orientation modules and orientation fractures in the monkey. A surprisingly similar distribution of blobless d- and l-hypercolumns is expected in the absence of the colour system. Applied to the apparently blobless cortex of the cat, the scheme explains the modulation of deoxyglucose uptake along the iso-orientation bands in a report of Löwel, Freeman, and Singer.

Do "d-blob" and "l-blob" hypercolumns tessellate the monkey visual cortex?

The orientation selective neurons in the monkey striate cortex seem to be organized in pairs of mirror symmetrical hypercolumnar patches, each of which centered by a "cytochrome oxidase blob." A simple scheme derived from recent data of Blasdel and Salama reconciles earlier models assuming either linear or circular representation of the preferred direction of edges in the visual field.

[Normal development of the fruitfly Drosophila in VLF magnetic fields (author's transl)]. / Normale Entwicklung der Fliege Drosophila in Niederfrequenten Magnetfeldern

Attempts to substantiate irreversible actions of a variety of magnetic fields on the fruitfly, Drosophila melanogaster, have been successful and unsuccessful in about equal numbers. The most conspicuous mutagenic effects apparently induced by pulsed HF-fields failed to appear under continuous electromagnetic irradiation. This seems to correlate the observed damage with the VLF-components of the pulsed fields. The present investigation is motivated by the occurrence of these components both in the atmosphere and in the vicinity of electrical appliances. A strain of normally viable wild type males and subnormally viable Attached X y w females was used in which the yield, and the sex ratio, of the progeny indicate, respectively, the extent of developmental damage and of sex-linked recessive lethal mutation induced by the exposure of detrimental conditions. Evaluation of 73,800 flies from subsequent generations of a control group and two test groups raised in steady, or rotating, homogeneous 9.6 kHz magnetic fields of about 2.5 G did not reveal any developmental or hereditarc load in the test groups. (Pressman 1970; Mittler 1973).