1932 3 S Schuster R Strauss KG Götz 2002-00-00 12 1591 1594 Current Biology Insects can estimate distance or time-to-contact of surrounding objects from locomotion-induced changes in their retinal position and/or size. Freely walking fruit flies (Drosophila melanogaster) use the received mixture of different distance cues to select the nearest objects for subsequent visits. Conventional methods of behavioral analysis fail to elucidate the underlying data extraction. Here we demonstrate first comprehensive solutions of this problem by substituting virtual for real objects; a tracker-controlled 360° panorama converts a fruit fly's changing coordinates into object illusions that require the perception of specific cues to appear at preselected distances up to infinity. An application reveals the following: (1) en-route sampling of retinal-image changes accounts for distance discrimination within a surprising range of at least 8-80 body lengths (20-200 mm). Stereopsis and peering are not involved. (2) Distance from image translation in the expected direction (motion parallax) outweighs distance from image expansion, which accounts for impact-avoiding flight reactions to looming objects. (3) The ability to discriminate distances is robust to artificially delayed updating of image translation. Fruit flies appear to interrelate self-motion and its visual feedback within a surprisingly long time window of about 2 s. The comparative distance inspection practiced in the small fruit fly deserves utilization in self-moving robots. no notspecified http://www.kyb.tuebingen.mpg.de/ published 3 15017 15423 869 3 R Strauss KG Götz M Renner 2001-10-00 187 617 632 Journal of Comparative Physiology A Visual motion processing enables moving fruit flies to stabilize their course and altitude and to approach selected objects. Earlier attempts to identify task-specific pathways between two photoreceptor systems (R1-6, R7+8) and three steering parameters (wingstroke asymmetry, abdomen deflection, hindleg deflection) attributed course control and object fixation to R1-6 mediated simultaneous reactions of these parameters. The present investigation includes first results from fixed flying or freely walking ninaE17 mutants which cannot synthesize the R1-6 photoreceptor-specific opsin. Retention of about 12 percent of the normal course control and about 58 percent of the object fixation in these flies suggests partial input sharing for both responses and, possibly, a specialization for large-field (R1-6) and small-field (R7+8) motion. Such signals must be combined to perceive relative motion between an object and its background. The combining links found in larger species might explain a previously neglected interdependence of course control and object fixation in Drosophila. Output decomposition revealed an unexpected orchestration of steering. Wingstroke asymmetry and abdomen deflection do not contribute in fixed proportions to the yaw torque of the flight system. Different steering modes seem to be selected according to their actual efficiency under closed-loop conditions and to the degree of intended turning. An easy experimental access to abdominal steering is introduced. no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 43 3 HG Krapp R Hengstenberg M Egelhaaf 2001-00-00 85 724 734 Journal of Neurophysiology 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. no notspecified http://www.kyb.tuebingen.mpg.de/ published 10 15017 15423 33 3 P Callaerts S Leng J Clements J Clements D Cribbs YY Kang U Walldorf KF Fischbach R Strauss 2001-00-00 46 73 88 Journal of Neurobiology A role for the Pax-6 homologue eyeless in adult Drosophila brain development and function is described. eyeless expression is detected in neurons, but not glial cells, of the mushroom bodies, the medullar cortex, the lateral horn, and the pars intercerebralis. Furthermore, severe defects in adult brain structures essential for vision, olfaction, and for the coordination of locomotion are provoked by two newly isolated mutations of Pax-6/eyeless that result in truncated proteins. Consistent with the morphological lesions, we observe defective walking behavior for these eyeless mutants. The implications of these data for understanding postembryonic brain development and function in Drosophila are discussed. no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 1163 7 I Berndt E Wascher VH Franz KG Götz HH Bülthoff Sarasota, FL, USA2001-12-00 318 First Annual Meeting of the Vision Sciences Society (VSS 2001) Purpose: Looking through laterally mirroring prisms produces at least two changes in the phenomenal appearance of the world: When stretching your right arm, for example, visual feedback will indicate that it is your left arm that is moving. But not only will the 'wrong' limb seem to be moving, it will also move in the diametrically opposite direction. Usually output and feedback of an action 'fit' (i.e., go to and come from the same limb). But when looking through mirroring prisms, visual feedback comes from the opposite arm and opposite direction. In order to behave properly under these circumstances, some kind of recalibration has to occur. The contralateral hemisphere is more strongly involved in controlling these arm movements. It is possible that this recalibration alters the lateralization of the neural activity that controls these movements. To test for this, we recorded event-related potentials (ERPs) and event-related lateralizations (ERLs) of the EEG during pointing movements with and without laterally mirrored vision. Targets were presented either centrally or laterally. Results: We found effects of mirrored vision on the lateralization of neural activity. The relative involvement of the hemisphere ipsilateral to the SEEN target position (objective position is reversed with mirrored feedback) increased, especially around 300-400ms after stimulus onset. Additionally, differences in the ERPs around the same time after target onset were evident. Both effects were maximal around the parietal and parieto-occipital sites, suggesting modified stimulus processing. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1163.pdf published -318 15017 1542215017 15423 57 7 I Berndt E Wascher VH Franz KG Götz HH Bülthoff Tübingen, Germany2001-03-00 147 4. Tübinger Wahrnehmungskonferenz (TWK 2001) Schaut man durch eine rechts-links spiegelnde Brille, so beobachtet man zwei Phänomene: Zeigt man z.B. mit dem rechten Arm, dann sieht es so aus, als führe der linke Arm diese Bewegung aus. Zudem scheint die Bewegung in die entgegengesetzte Richtung zu verlaufen. Befehl und Rückmeldung stimmen also nicht mehr überein, sind gegenläufig. Ein effizientes Verhalten mit gespiegeltem Feedback erfordert eine Umkodierung der visuomotorischen Koordination. Diese sollte sich in einer Veränderung der neuronalen Aktivität im EEG niederschlagen. Wir fanden in einer vorangegangenen Studie, dass sich die verschiedenen Anteile einer Zeigebewegung in Lateralisierungen hirnelektrischer Potentiale im EEG (event-related lateralizations = ERLs) abbilden: Auswahl des Effektors, Lokalisation des Zielreizes, Bewegungsrichtung und Kontrolle der räumlich gerichteten Bewegung. Diese Lateralisierungen des EEG während der Zeigebewegung sollten sich auch durch die Spiegelung der visuellen Rückmeldung spezifisch verändern. Um dies zu untersuchen wurden EEG-Messungen während Zeigebewegungen mit und ohne Spiegelung des Gesichtsfeldes durchgeführt. Der Zielreiz wurde dabei entweder zentral oder lateralisiert (+/- 1,7 Grad) dargeboten. Es zeigte sich ein Effekt der Spiegelung auf die Lateralisierung des EEGs. Dieser bestand aus einer höheren Aktivierung der zum gesehenen Zielreiz ipsilateralen Hemisphäre im Vergleich zur ungespiegelten Bedingung. (Zu beachten ist, dass sich die objektive Position bei Spiegelung umkehrt.) Dieser Effekt trat ca. 300-400 ms nach Stimulus Onset auf und war maximal in parietalen und parieto-occipitalen Regionen. Die Spiegelung verursachte eine räumlich und zeitlich eingrenzbare Veränderung der Lateralisierung neuronaler Aktivität. Es liegt nahe, dass dies eine Modifikation der Zielreiz-Verarbeitung darstellt und durch die Umkodierung der visuomotorischen Koordination verursacht wird. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf57.pdf published -147 15017 1542215017 15423 330 2 R Hengstenberg Macmillan Magazines Ltd. Brunel Road, Basingstoke, Hampshire RG 21 2XS, UK 1999-00-00 18 19 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf330.pdf published 1 15017 15423 462 15 S Leng 1999-00-00 no notspecified published 15017 15423 235 3 R Hengstenberg 1998-00-00 392 757 758 Nature no notspecified http://www.kyb.tuebingen.mpg.de/ published 1 15017 15423 241 3 HG Krapp B Hengstenberg R Hengstenberg 1998-00-00 79 1902 1917 Journal of Neurophysiology The third visual neuropil (lobula plate) of the blowfly Calliphora erythrocephala is a center for processing motion information. It contains, among others, 10 individually identifiable "vertical system" (VS) neurons responding to visual wide-field motions of arbitrary patterns. We demonstrate that each VS neuron is tuned to sense a particular aspect of optic flow that is generated during self-motion. Thus the VS neurons in the fly supply visual information for the control of head orientation, body posture, and flight steering. To reveal the functional organization of the receptive fields of the 10 VS neurons, we determined with a new method the distributions of local motion sensitivities and local preferred directions at 52 positions in the fly's visual field. Each neuron was identified by intracellular staining with Lucifer yellow and three-dimensional reconstructions from 10-µm serial sections. Thereby the receptive-field organization of each recorded neuron could be correlated with the location and extent of its dendritic arborization in the retinotopically organized neuropil of the lobula plate. The response fields of the VS neurons, i.e., the distributions of local preferred directions and local motion sensitivities, are not uniform but resemble rotatory optic flow fields that would be induced by the fly during rotations around various horizontal axes. Theoretical considerations and quantitative analyses of the data, which will be presented in a subsequent paper, show that VS neurons are highly specialized neural filters for optic flow processing and thus for the visual sensation of self-motions in the fly. no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 147 3 JD Armstrong JS de Belle Z Wang K Kaiser 1998-00-00 5 102 114 Learning & Memory Paired brain centers known as mushroom bodies are key features of the circuitry for insect associative learning, especially when evoked by olfactory cues. Mushroom bodies have an embryonic origin, and unlike most other brain structures they exhibit developmental continuity, being prominent components of both the larval and the adult CNS, Here, we use cell-type- specific markers, provided by the P{GAL4} enhancer trap system, to follow specific subsets of mushroom body intrinsic and extrinsic neurons from the larval to the adult stage. We find marked structural differences between the larval and adult mushroom bodies, arising as the consequence of large-scale reorganization during metamorphosis. Extensive, though incomplete, degradation of the larval structure is followed by establishment of adult specific alpha and beta lobes, Kenyon cells of embryonic origin, by contrast, were found to project selectively to the adult gamma lobe, We propose that the gamma lobe stores information of relevance to both developmental stages, whereas the alpha and beta lobes have uniquely adult roles. no notspecified http://www.kyb.tuebingen.mpg.de/ published 12 15017 15423 258 3 R Strauss J Pichler 1998-00-00 182 411 423 Journal of Comparative Physiology A In arena experiments with the walking fruit fly, we found a remarkable persistence of orientation toward a landmark that disappeared during the fly's approach. The directional stability achieved by 'after-fixation' allows a fly to continue pursuit under natural conditions, where a selected target is frequently concealed by surrounding structures. The persistence of after-fixation was investigated in Buridan's paradigm, where a fly walks persistently back and forth between two inaccessible landmarks. Upon disappearance of a selected target, the flies maintained their intended course for more than 15 body lengths of approximately 2.5 mm in about 50% of the trials. About 13% even exceeded 75 body lengths. About 88% of the approaches clustered in equal portions around peaks at 2.4 s and 8.6 s. About 12% of the approaches persisted even longer. In contrast, a single peak at about 2.2 s is sufficient to describe the persistence of orientation in a random walk. The ability to pursue an invisible landmark is disturbed neither by a transient angular deviation from the course toward this landmark, when this target disappeared, nor by a distracting second landmark. Accordingly, after-fixation seems to require an internal representation of the direction toward the concealed target, and idiothetical course control to maintain this direction. no notspecified http://www.kyb.tuebingen.mpg.de/ published 12 15017 15423 236 3 R Hengstenberg 1998-00-00 392 231 232 Nature no notspecified http://www.kyb.tuebingen.mpg.de/ published 1 15017 15423 1013 7 R Strauss M Renner KG Götz 1998-00-00 New Neuroethology on the Move. Proceedings of the 26th Goettingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published 0 15017 15423 339 2 R Hengstenberg World Scientific Publishing 5 Tuh Tuck Link, 596224 Singapore 1998-00-00 20 22 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf339.pdf published 2 15017 15423 345 2 R Strauss Gesellschaft für wissenschaftliche Datenverarbeitung (GWDG) Am Fassberg, Turm 6, 37077 Göttingen 1998-00-00 53 78 no notspecified http://www.kyb.tuebingen.mpg.de/ published 25 15017 15423 1171 2 KG Götz World Scientific Publishing 5 Tuh Tuck Link, 596224 Singapore 1998-00-00 431 446 no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 338 2 KG Götz World Scientific Publishing, Singapore 1998 5 Toh Tuck Link, 596224 Singapore 1998-00-00 447 456 no notspecified http://www.kyb.tuebingen.mpg.de/ published 9 15017 15423 340 2 R Hengstenberg World Scientific Publishing 5 Tuh Tuck Link, 596224 Singapore 1998-00-00 41 52 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf340.pdf published 11 15017 15423 342 2 R Hengstenberg HG Krapp B Hengstenberg World Scientific Publishers 5 Tuh Tuck Link, 596224 Singapore 1998-00-00 53 70 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf342.pdf published 17 15017 15423 281 7 S Leng R Strauss Göttingen, Germany1998-05-00 259 26th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -259 15017 15423 335 7 U Wannek R Strauss Göttingen, Germany1998-05-00 258 26th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -258 15017 15423 1195 7 R Strauss M Renner KG Götz Göttingen, Germany1998-05-00 421 26th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -421 15017 15423 276 7 MO Franz R Hengstenberg HG Krapp Göttingen, Germany1998-05-00 419 26th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf276.pdf published -419 15017 15423 367 3 HG Krapp R Hengstenberg 1997-00-00 37 225 234 Vision Research We present a method to determine, within a few seconds, the local preferred direction (LPD) and local motion sensitivity (LMS) in small patches of the receptive fields of wide-held motion-sensitive neurons. This allows us to map, even during intracellular recordings, the distribution of LPD and LMS over the huge receptive fields of neurons sensing self-motions of the animal. Comparisons of the response field of a given neuron with the optic flow fields caused by different movements in space, allows us to specify the particular motion of the animal sensed by that neuron. Copyright (C) 1996 Elsevier Science Ltd. no notspecified http://www.kyb.tuebingen.mpg.de/ published 9 15017 15423 362 3 A Guo KG Götz 1997-00-00 4 192 204 Learning & Memory 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 ny 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 BAI, 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. no notspecified http://www.kyb.tuebingen.mpg.de/ published 12 15017 15423 382 3 C Varnam R Strauss JS de Belle M Sokolowski 1997-00-00 11 99 115 Journal of Neurogenetics no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 381 3 R Strauss S Schuster KG Götz 1997-00-00 200 1281 1296 Journal of Experimental Biology no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 1304 7 R Hengstenberg 1997-00-00 15 19 International School of Biophysics 1994 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1304.pdf published 4 15017 15423 410 7 S Leng R Strauss Göttingen, Germany1997-05-00 294 25th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -294 15017 15423 1194 7 M Renner KG Götz Göttingen, Germany1997-05-00 295 25th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -295 15017 15423 426 7 R Strauss T Trinath S Leng Göttingen, Germany1997-05-00 293 25th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -293 15017 15423 436 7 U Wannek R Strauss Göttingen, Germany1997-05-00 292 25th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -292 15017 15423 422 7 S Schuster Göttingen, Germany1997-05-00 291 25th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -291 15017 15423 539 3 G Stange R Hengstenberg 1996-02-00 2 64 209 218 Journal of Neuroscience Methods We present a method to record and simultaneously display the three rotatory components of arbitrary head turns of an insect flying stationarily in a wind tunnel or walking on a treadmill. An elongated marker, placed on the fly's forehead, is video- recorded from ahead under deep red stroboscopic illumination, invisible to the insect. A fast on-board image processor of a PC video-adapter (True Vision, AT-Vista), programmed in its native code, extracts position and orientation of the marker in the video-image. The host PC transforms these data into calibrated head angles and displays stimulus and response components after 40 ms processing time at a rate of 50 frames per second. Head turns are measured relative to the fly's trunk even when the fly is rotated around its body axis provided that it is aligned with the video-axis. Technical tests, as well as recordings from live flies responding to various stimuli, illustrate the performance and accuracy of the procedure. This minimally invasive method of motion recording should be easily adaptable to other insects and to similar movements of small parts. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf539.pdf published 9 15017 15423 526 3 F-O Lehmann KG Götz 1996-00-00 179 311 322 Journal of Comparative Physiology A-Sensory Neural And Behavioral Physiology 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. no notspecified http://www.kyb.tuebingen.mpg.de/ published 11 15017 15423 525 3 HG Krapp R Hengstenberg 1996-00-00 384 463 466 Nature Humans, animals and some mobile robots use visual motion cues for object detection and navigation in structured surroundings (1-4). Motion is commonly sensed by large arrays of small field movement detectors, each preferring motion in a particular direction (5, 6). Self-motion generates distinct 'optic flow fields' in the eyes that depend on the type and direction of the momentary locomotion (rotation, translation) (7). To investigate how the optic flow is processed at the neuronal level, we recorded intracellularly from identified interneurons in the third visual neuropile of the blowfly (8). The distribution of local motion tuning over their huge receptive fields was mapped in detail. The global structure of the resulting 'motion response fields' is remarkably similar to optic flow fields. Thus, the organization of the receptive fields of the so-called VS neurons (9,10) strongly suggests that each of these neurons specifically extracts the rotatory component of the optic flow around a particular horizontal axis. Other neurons are probably adapted to extract translatory flow components. This study shows how complex visual discrimination can be achieved by task-oriented preprocessing in single neurons. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf525.pdf published 3 15017 15423 446 3 JS de Belle M Heisenberg 1996-00-00 93 9875 9880 Proceedings of the National Academy of Sciences of the United States of America Mutations in 12 genes regulating Drosophila melanogaster mushroom body (MB) development were each studied in two genetic backgrounds. In all cases, brain structure was qualitatively or quantitatively different after replacement of the "original" genetic background with that of the Canton Special wild-type strain. The mushroom body miniature gene (mbm) was investigated in detail. mbm supports the maintenance of MB Kenyon cell fibers in third instar larvae and their regrowth during metamorphosis. Adult mbm(1) mutant females are lacking many or most Kenyon cell fibers and are impaired in MB-mediated associative odor learning. We show here that structural defects in mbm(1) are apparent only in combination with an X-linked, dosage-dependent modifier (or modifiers). In the Canton Special genetic background, the mbm(1) anatomical phenotype is suppressed, and MBs develop to a normal size. However, the olfactory learning phenotype is not fully restored, suggesting that submicroscopic defects persist in the MBs. Mutant mbm(1) flies with full-sized MBs have normal retention but show a specific acquisition deficit that cannot be attributed to reductions in odor avoidance, shock reactivity, or locomotor behavior. We propose that polymorphic gene interactions (in addition to ontogenetic factors) determine MB size and, concomitantly, the ability to recognize and learn odors. no notspecified http://www.kyb.tuebingen.mpg.de/ published 5 15017 15423 543 3 CJ Varnam R Strauss JS de Belle MB Sokolowski 1996-00-00 11 99 115 Journal of Neurogenetics The central complex (CC) is a prominent component of the adult insect brain. In Drosophila melanogaster, mutations which alter CC structure also impair adult locomotion. This has led to the suggestion that the CC functions as a higher organizer of adult locomotor patterns (Strauss and Heisenberg, 1993). In the present study, we describe altered larval behavior resulting from mutations in six CC structural genes. Differences from the control strain were found for larvae from each CC mutant strain in at least one of three assays. central body defect(1) (cbd(1)), central complex deranged(1) (ccd(1)), central brain deranged(1) (ceb(1)) and central complex(1) (cex(1)) larvae all had general defects in locomotion (on a non-nutritive agar surface). Both ellipsoid body open(2) (ebo(2)) and no bridge(1) (nob(1)) had larval foraging behavior defects (on a nutritive yeast surface). Only cex(1) larvae required significantly longer time in a roil over assay of muscle tone. Genetic analysis suggested that nob(1) interacts additively with two other genes influencing larval foraging behavior, foraging (for) and Chaser (Car). Sor also had an influence on adult foraging, whereas here we found that Csr did not. We did not include adult foraging behavior tests of the CC mutants due to general locomotion defects in these flies (Strauss and Heisenberg, 1993). no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 517 3 G Heide KG Götz 1996-00-00 199 1711 1726 Journal Of Experimental Biology no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 447 3 MH Dickinson KG Götz 1996-00-00 199 2085 2104 Journal of Experimental Biology no notspecified http://www.kyb.tuebingen.mpg.de/ published 19 15017 15423 1193 7 S Schuster KG Götz Göttingen, Germany1996-05-00 353 24th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -353 15017 15423 598 7 U Wannek R Strauss 1996-00-00 133 In: Brain and Evolution, Vol. II,(Eds.) N. Elsner and H.-U. Schnitzler. Thieme, Stuttgart no notspecified http://www.kyb.tuebingen.mpg.de/ published -133 15017 15423 580 7 S Leng R Strauss 1996-00-00 134 Brain and Evolution, Vol. II, (Eds.) N. Elsner, H.-U. Schnitzler. Thieme, Stuttgart no notspecified http://www.kyb.tuebingen.mpg.de/ published -134 15017 15423 574 7 R Hengstenberg HG Krapp 1996-00-00 349 Brain and Evolution, Vol. II, (Eds.) N. Elsner, H.U. Schnitzler. Thieme, Stuttgart 1996 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf574.pdf published -349 15017 15423 599 7 J Wulf JS de Belle C Helfrich-Förster 1996-00-00 29 In: Brain and Evolution, Vol. I, (Eds.) N. Elsner and H.-U. Schnitzler. Thieme, Stuttgart no notspecified http://www.kyb.tuebingen.mpg.de/ published -29 15017 15423 567 7 JS de Belle M Heisenberg 1996-00-00 24 J. Neurogenet. no notspecified http://www.kyb.tuebingen.mpg.de/ published -24 15017 15423 591 7 S Schuster KG Götz 1996-00-00 353 Brain and Evolution, Vol. II, (Eds.) N. Elsner, H.U. Schnitzler. Thieme, Stuttgart no notspecified http://www.kyb.tuebingen.mpg.de/ published -353 15017 15423 592 7 R Strauss T Trinath 1996-00-00 135 In: Brain and Evolution, Vol. II, (Eds.) N. Elsner and H.-U. Schnitzler. Thieme, Stuttgart no notspecified http://www.kyb.tuebingen.mpg.de/ published -135 15017 15423 613 15 S Schuster 1996-00-00 no notspecified published 15017 15423 1181 3 KG Götz 1995-06-09 Triest Triest International Centre for Theoretical Physics no notspecified http://www.kyb.tuebingen.mpg.de/ published 0 15017 15423 1182 3 KG Götz 1995-06-09 Triest Triest International Centre for Theoretical Physics no notspecified http://www.kyb.tuebingen.mpg.de/ published 0 15017 15423 1183 3 KG Götz 1995-06-09 Triest Triest International Centre for Theoretical Physics no notspecified http://www.kyb.tuebingen.mpg.de/ published 0 15017 15423 621 3 JS de Belle 1995-00-00 15 245 247 Neuron no notspecified http://www.kyb.tuebingen.mpg.de/ published 2 15017 15423 1303 7 HG Krapp R Hengstenberg Cambridge, UK1995-09-00 255 4th International Congress of Neuroethology The control of locomotion in a given environment requires information about instantaneous self-motion. Visually oriented animals, including man, may gain such information by analyzing the momentary optic flow pattern generated over both eyes during relative movement between animal and environment. Optic flow patterns can be described by vector fields where each single vector indicates the direction and velocity of the local relative movement at a certain position within the visual field. An optic flow pattern depends upon a set of motion parameters, namely (i) the direction of gaze and (ii) the rotatory and (iii) translatory components of self-motion. The translatory flow vectors also depend an the distance between visual objects and the eyes. Therefore, optic flow fields contain valuable information about the 3D-layout of the surroundings and instantaneous self-motion (Koenderink and van Doorn, 1987). About 50 motion-sensitive, wide-field interneurons which are assumed to be' involved in locomotor control are located in the third visual neuropil (lobula plate) of the blowfly's (Calliphora erythrocephala) visual system (Hausen, 1993). The output of many direction-specific movement detectors (EMDS) with small receptive fields are spatially integrated in a retinotopic manner an the dendrites of these interneurons. Are such interneurons adapted to sense specific aspects of the momentary optic flow field? To address this question, we investigated the receptive field organization of 10 identifiable interneurons of the so called vertical-system (VS; Hengstenberg, 1982) in great detail. We recorded intracellularly from the VS-neurons to determine the spatial distribution of local preferred directions and motion sensitivities at 52 positions spaced equally over the ipsilateral visual hemisphere (for method see: Menzel and Hengstenberg, 1991; Krapp and Hengstenberg 1992). The resulting response fields of the VS-neurons (about 90 recordings) show striking similarities to optic flow fields generated by specific motions in space (Krapp and Hengstenberg, 1994). By applying an iterative least square formalism (Koenderink and van Doorn, 1987) to the response fields we calculated the optimal self-motion parameters (translatory and rotatory component) for each VS-neuron. These parameters describe an optic flow field that best fits the respective measured response field. To find out whether the VS-neurons are functionally tuned more to the translatory or to the rotatory component of self-motion we systematically varied the optimal motion parameters. The error between the measured response field and the calculated optic flow field increases if both the translatory and the rotatory component deviate from the optimal motion parameters. The increase in the error is almost the same if only the rotatory component is varied. In contrast, if the translatory component is varied and the rotatory component is kept optimal the increase in the error is considerably smaller. The analysis of the response fields of the VS-neurons leads to the following conclusion: the VS-neurons are functionally tuned to sense rotations around different horizontally aligned body axes. The neurons VS1-VS3 are optimized to sense optic flow fields generated during nose-up pitch. VS4-VS7 are filter neurons for counterclockwise roll and VS8-VS10 are adapted to rotations around an axis that lies between the pitch and roll axes. Thus, the signals of the VS-neurons could contribute directly to visual flight control and gaze stabilization. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1303.pdf published -255 15017 15423 1192 7 S Schuster KG Götz R Strauss Göttingen, Germany1995-03-00 403 23rd Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -403 15017 15423 1301 7 HG Krapp R Hengstenberg 1995-00-00 404 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1301.pdf published -404 15017 15423 1302 7 R Hengstenberg 1995-00-00 255 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1302.pdf published -255 15017 15423 1299 3 G Nalbach R Hengstenberg 1994-00-00 175 695 708 J. Comp. Physiol. A no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1299.pdf published 13 15017 15423 1300 2 R Hengstenberg Vandenhoek & Ruprecht Göttingen, Germany 1994-00-00 199 207 Jahruch der Max-Planck-Gesellschaft 1994 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1300.pdf published 8 15017 15423 1170 2 KG Götz edited by Schildberger, K. & Elsner, N. Progress in Zoology 39. G. Fischer, Stuttgart 1994-00-00 47 59 no notspecified http://www.kyb.tuebingen.mpg.de/ published 12 15017 15423 1191 7 S Schuster KG Götz Göttingen, Germany1994-05-00 304 22nd Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -304 15017 15423 515 7 HG Krapp B Hengstenberg R Hengstenberg 1994-00-00 453 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf515.pdf published -453 15017 15423 1298 3 H Schuppe R Hengstenberg 1993-00-00 173 143 149 J. Comp. Physiol. A The 3 Ocelli of the blowfly Calliphora erythrocephala, grouped close together on the top of the head (Fig.1), have large, extensively overlapping visual fields. Together they view the entire upper hemisphere of the surroundings plus part of the lower hemisphere (Fig. 5, 7). It is shown for the lateral ocelli that despite the underfocussing of the ocellar lens large patterns are imaged on the receptor mosaic. Because of the astigmatism of the lens, patterns in longitudinal orientations are more accurately represented than in others (Fig. 3). Nevertheless, an artifical horizon rotated around the long axis of the animal does not elicit head roll. Likewise, changes of overall brightness in the visual field of the median and one lateral ocellus elicit only weak phasic-tonic &quot;dorsal light responses&quot; of the animal which supplement the tonic dorsal light responses mediated by the compound eyes (Figs. 9, 10). Our results show that, in Calliphora, the ocelli have little influence on head orientation during flight, and must be assumed to serve other functions. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1298.pdf published 6 15017 15423 563 3 MH Dickinson KG Götz 1993-00-00 182 173 189 J.exp.Biol. no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 562 3 MH Dickinson KG Götz 1993-00-00 174 45 64 J.exp.Biol. no notspecified http://www.kyb.tuebingen.mpg.de/ published 19 15017 15423 1296 2 R Hengstenberg Elsevier Science Publ., Amsterdam 1993-00-00 285 298 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1296.pdf published 13 15017 15423 1167 2 KG Götz Steiner Stuttgart, Germany 1993-00-00 110 114 Jahrbuch 1992 der Akademie der Wissenschaften und der Literatur no notspecified http://www.kyb.tuebingen.mpg.de/ published 4 15017 15423 1169 2 KG Götz edited by Wang, B.Y. & Cai, N.S. The School of Life Sciences, Fudan University, Shanghai 1993-00-00 1 14 no notspecified http://www.kyb.tuebingen.mpg.de/ published 13 15017 15423 1297 7 HG Krapp R Hengstenberg 1993-00-00 357 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1297.pdf published -357 15017 15423 1168 41 KG Götz 1295 3 T Preuss R Hengstenberg 1992-00-00 171 483 493 J. Comp. Physiol. A The blowfly Calliphora has a mobile head and various, presumably proprioceptive, sense organs in the neck region. The &quot;prosternal organs&quot; are a pair of mechanosensory hair fields, each comprising ca. 110 sensilla. We studied their structure (Figs. 2-4), kinematics (Figs.5, 6) and,&nbsp; after surgery, their influence on head posture (Figs. 7-11) in order to reveal their specific function. The hair sensilla are stucturally polarized, all in roughly the same direction, and are stimulated by dorsoventral bending of the hairs (Figs. 3, 4). This occurs indirectly by flap- movements of two contact sclerites (Figs. 3, 6); they move in the same direction during pitch turns of the head, in opposite directions during roll turns, and barely at all during yaw turns of the head (Fig. 5). Bending and arresting all hairs of one field elicits a head roll bias to the non-operated side (Fig. 7) during tethered flight in visually featureless surroundings. In contrast, shaving all hairs of one field elicits a head roll to the operated side (Figs. 8-10). The surgically induced bias of head posture is not compensated within&nbsp; three days (Fig. 10). Our results show that the prosternal organs of Calliphora sense pitch and roll turns of the fly's head, and control at least its roll position. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1295.pdf published 10 15017 15423 1291 2 R Hengstenberg Oxford Univ. Press, NY 1992-00-00 49 55 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1291.pdf published 6 15017 15423 1190 7 F-O Lehmann KG Götz Göttingen, Germany1992-06-00 173 20th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -173 15017 15423 1293 7 R Hengstenberg 1992-00-00 305 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1293.pdf published -305 15017 15423 1294 7 HG Krapp R Hengstenberg 1992-00-00 306 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1294.pdf published -306 15017 15423 561 3 KG Götz 1991-00-00 95 279 286 Zool.Jb.Physiol. no notspecified http://www.kyb.tuebingen.mpg.de/ published 7 15017 15423 1287 3 R Hengstenberg 1991-00-00 3 19 29 The Neuroscience Flies move their eyes by turning their heads either spontaneously or in response to unexpected disturbances of their preferred flight attitude. They use several visual and mechanosensory cues to keep flight balance and to stabilize their eyes relative to the surroundings by compensatory head/eye movements. The various sensory subsystems have different speed characteristics and cooperate autonomously to provide the fly with fast and accurate visual stabilization. Head and trunk are coordinated by neck sense organs affecting head posture as well as flight torque. The functional structure of the fly's gaze control system changes its input-and output configuration when the fly alternates between flight and walking no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1287.pdf published 10 15017 15423 1288 3 R Hengstenberg 1991-00-00 95 297 304 Zoologische Jahrbücher: Abteilung für allgemeine Zoologie und Physiologie der Tiere no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1288.pdf published 7 15017 15423 1285 7 J Menzel R Hengstenberg 1991-00-00 274 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1285.pdf published -274 15017 15423 1286 7 R Hengstenberg 1991-00-00 273 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1286.pdf published -273 15017 15423 1290 7 R Hengstenberg 1991-00-00 344 345 Verhdlg. d. Deutsch. Zool. Ges. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1290.pdf published 1 15017 15423 1289 7 R Hengstenberg 1991-00-00 100 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1289.pdf published -100 15017 15423 560 3 JM Zanker KG Götz 1990-00-00 327 19 44 Phil.Trans.R.Soc.Lond.B no notspecified http://www.kyb.tuebingen.mpg.de/ published 25 15017 15423 1189 7 F-O Lehmann KG Götz Göttingen, Germany1990-06-00 77 18th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -77 15017 15423 1284 7 T Preuss R Hengstenberg Göttingen, Germany1990-06-00 78 18th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1284.pdf published -78 15017 15423 1165 2 KG Götz edited by Nachtigall, W. Biona report 8. G. Fischer, Stuttgar7 1989-00-00 27 38 no notspecified http://www.kyb.tuebingen.mpg.de/ published 11 15017 15423 1166 2 KG Götz edited by Singh, R.N. & Strausfeld, N.J. Plenum Press, New York 1989-00-00 139 153 no notspecified http://www.kyb.tuebingen.mpg.de/ published 14 15017 15423 763 7 S Edelman HH Bülthoff D Weinshall Zichron Yaakov, Israel1989-09-00 517 Twelfth European Conference on Visual Perception (ECVP 1989) no notspecified http://www.kyb.tuebingen.mpg.de/ published -517 15017 15423 764 7 HH Bülthoff D Kersten Zichron Yaakov, Israel1989-09-00 504 Twelfth European Conference on Visual Perception (ECVP 1989) no notspecified http://www.kyb.tuebingen.mpg.de/ published -504 15017 15423 1283 7 R Hengstenberg 1989-00-00 257 258 no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1283.pdf published 1 15017 15423 559 3 KG Götz 1988-03-00 4 58 213 223 Biological Cybernetics 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. no notspecified http://www.kyb.tuebingen.mpg.de/ published 10 15017 15423 1277 3 PG Layer S Rommel HH Bülthoff R Hengstenberg 1988-03-00 3 251 587 595 Cell and Tissue Research AChE-positive cells suddenly amass in a superficial layer of the neuroepithelium; this layer finally covers, in a sheat-like manner, the entire surface of the embryonic chicken brain. This feature is functionally not understood; however, it appears shortly after the neurons become post-mitotic, and the lateral extensions of this layer can easily be traced using histochemistry on serial brain sections. The layer can therefore be exploited to delineate spatially the waves of onset of biochemical tissue differentiation. We have studied whole brains between stages 11 and 30 and provide the first complete spatial schemes of brain differentiation based on computer-reconstructed, two- and three-dimensional maps. The brain does not differentiate in one smooth coherent wave, but instead five separate primary AChE-activation zones are detected: the first originating at stage 11 (ldquorhombencephalic waverdquo), the second at the same time (ldquomidbrain waverdquo), the third at stage 15 ("ldquotectal waverdquo). A fourth zone develops later, at stage 18, from the bottom part of the telencephalon to the top. Retinal development also starts at stage 18. In a given area, it appears that AChE-development shortly precedes that of the formation of major fiber tracts. AChE might therefore represent a prerequisite for fiber growth and pathfinding. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/independent_spatial_waves_of_biochemical_differentiation_1277[0].pdf published 8 15017 15423 1281 3 R Hengstenberg 1988-03-00 2 163 151 165 Journal of Comparative Physiology A In the blowflyCalliphora flying stationarily in a wind tunnel, compensatory head movements were elicited by rolling the fly about its longitudinal axis (Fig. 1). Responses were recorded on video tape, and evaluated by single frame analysis. Active head movements were observed in response to visual and mechanosensory stimuli (Fig. 2). They are not made or caused by the head's inertial momentum (Fig. 11). Gravity, used by walking flies to align their head with the vertical, does not seem to be perceived during flight (Figs. 3–6) but has a passive stabilizing effect upon the flight attitude (Fig. 7). A difference in aerodynamical load of the two wings elicits a transient head roll partly compensating a banked attitude (Figs. 4–6). The majority of campaniform sensilla at the wing base seems suitable to measure wing load. Steady roll motion elicits a steady compensatory head roll which does not vanish even after 8 min of rotation at constant angular velocity (Fig. 8). Roll motion is most efficient at high roll speeds (100‡/s<w<2000‡/s). Mechanical motion perception fails if both halteres are disabled by arresting their oscillation or by amputation of the haltere knobs (Fig. 11). Flies with only one haltere intact cannot distinguish pitch from roll, but with respect to the sense of rotation they still respond bidirectionally (Fig. 12). Haltere dynamics and the response characteristics of haltere sensilla are discussed on the basis of recent results. Head/body coordination is demonstrated in the absence of any roll stimulus (Fig. 3 a). The role of resilience of the neck skeleton, and that of different neck sense organs are discussed. Mechanosensory roll control inCalliphora depends upon the locomotor state: When walking, the fly aligns its head vertically by gravity perception (Horn 1982). When flying, it controls only fast rotations. Passive attitude stabilization and visual means of control are required to maintain an upright flight attitude and head orientation. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1281.pdf published 14 15017 15423 1282 7 R Hengstenberg N Bayer Bielefeld, Germany1988-05-00 203 81. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1282.pdf published -203 15017 15423 1278 7 R Hengstenberg K Hausen B Hengstenberg Göttingen, Germany1988-00-00 129 16. Göttinger Neurobiologentagung no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1278.pdf published -129 15017 15423 1279 7 K Hausen R Hengstenberg T Wiegand Göttingen, Germany1988-00-00 130 16. Göttinger Neurobiologentagung no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1279.pdf published -130 15017 15423 1280 7 H Schuppe R Hengstenberg Göttingen, Germany1988-00-00 215 16. Göttinger Neurobiologentagung no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1280.pdf published -215 15017 15423 557 3 KG Götz 1987-05-00 2-3 56 107 109 Biological Cybernetics 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. no notspecified http://www.kyb.tuebingen.mpg.de/ published 2 15017 15423 556 3 KG Götz 1987-03-00 1 128 35 46 Journal of Experimental Biology Tethered flight in a 3-day-old female Drosophila was sustained for 32.2 h with only short interruptions during uptake of sucrose solution. The course-control reactions derived from the difference of the wingbeat amplitudes on either side have been used to simulate the rotatory displacement of the surrounding landmarks during a comparable turn in free flight. Stabilization of a target in the preferred area of the visual field requires continuous visual attention. A rate of about 5 course-correcting manoeuvres per second was maintained throughout the experiment. Drosophila seems to be able to cover long distances in search of a favourable habitat. Flight-specific carbohydrate consumption is equivalent to a metabolic power input per body weight of about 18 W N−1. The tethered fly produces about 40 % of the lift required to sustain hovering flight. The resulting mechanochemical efficiency of about 0.04-0.07 is within the expected order of magnitude for flying insects. Expenditure of reserve substances may account for the difference between the comparatively low power input of about 7 WN−1 derived from carbohydrate uptake in the first hours of flight (Wigglesworth, 1949), and the actual metabolic turnover of about 21WN−1 derived from oxygen consumption during this period (Laurie-Ahlberg et al. 1985). Weis-Fogh's ‘clap and fling’, a widespread lift-generating process exploiting the aerodynamic wing interference at the dorsal end of the wingbeat, was in action throughout the flight. However, there were two significant modifications (as first conceived by Ellington, 1980): (1) during ‘clap’, there is a progress of wing contact from the leading to the trailing edge, which is likely to ‘squeeze’ a thrust-generating jet of air to the rear; (2) during ‘fling’, there is a progress of wing separation in the same direction, which is described as a ‘peel’ resembling the progressive separation of two plastic foils pulled apart against forces of mutual attraction. The wings of the test fly survived about 23 million such peels without damage. Increasing airspeed decreases the intensity of ‘clap and fling’ in Drosophila: results obtained in the wind tunnel show the transition to a ‘near clap and fling’, lacking mutual wing contact. no notspecified http://www.kyb.tuebingen.mpg.de/ published 11 15017 15423 1276 7 K Hausen R Hengstenberg New Orleans, LA, USA1987-11-00 17th Annual Meeting of the Society for Neuroscience (Neuroscience 1987) no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1276.pdf published 0 15017 15423 1275 7 M Gewecke K Kirschfeld R Feiler E Hartwieg R Hengstenberg G Lenz Göttingen, Germany1987-00-00 148 15th Göttingen Neurobiology Conference no notspecified http://www.kyb.tuebingen.mpg.de/ published -148 15017 15423 558 10 KG Götz 1272 3 R Hengstenberg DC Sandeman B Hengstenberg 1986-05-00 1249 227 455 482 Proceedings of the Royal Society of London B Video records were made of the blowfly Calliphora erythrocephala L. mainly during tethered flight in a wind-tunnel, to study its movements about the longitudinal body axis (roll). During undisturbed flight, flies hold their head on average aligned with the body but may turn it about all three body axes. Pitch and yaw turns of the head are comparatively small (20 degrees), whereas roll turns can be large (90 degrees), and fast (1200 degrees s$^{-1}$). When passively rolled, flies produce compensatory head movements during walking or flight; at rest this reflex is turned off. Flies perceive a static misalignment relative to the vertical, as well as roll motion up to 10 000 degrees s$^{-1}$. Within this range flies counteract an imposed roll with maximal gain at about 1000 degrees s$^{-1}$. Compensatory head movements are made with very low latency (down to $\Delta $t $\approx $ 5 ms), and with considerable speed (up to $\omega $ = 1000 degrees s$^{-1}$). Flies may `disregard an apparent deviation from their correct orientation, and may superimpose spontaneous head movements on those elicited by a stimulus. Compensatory head movements generally undercompensate the imposed misalignment. Simultaneously, however, flies modify their wing pitch and wingbeat amplitude to produce a compensatory roll torque. Since head and body roll act simultaneously and in the same direction, the overall speed and degree of head realignment, relative to external coordinates, increase considerably. This is certainly an advantage for flight in turbulent air. In still air, without need to correct an imposed misalignment, flies nevertheless produce spontaneous fluctuations of their flight torque, and head roll movements in the opposite direction. This is to be expected if flies intend to keep their eyes aligned with the coordinates of the environment while spontaneously performing banked turns. The limits of fly vision and the advantages of compensatory head movements for different visually guided behaviour are discussed. Compensatory head roll movements give flies greater manoeuvrability when cruising than the visual system would allow, without such a stabilizing reflex. no notspecified http://www.kyb.tuebingen.mpg.de/ published 27 15017 15423 1162 2 KG Götz Max-Planck-Gesellschaft München, Germany 1986-00-00 280 286 Jahrbuch der Max-Planck-Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de/ published 6 15017 15423 1273 7 G Nalbach R Hengstenberg München, Germany1986-05-00 229 79. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1273.pdf published -229 15017 15423 1274 7 G Nalbach R Hengstenberg Göttingen, Germany1986-00-00 14. Göttinger Neurobiologentagung no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1274.pdf published 0 15017 15423 553 3 KG Götz R Biesinger 1985-05-00 3 156 319 327 Journal of Comparative Physiology A The term ‘centrophobism’ is introduced to describe a newly discovered modification of search behavior in the walking fruitfly,Drosophila melanogaster: the avoidance of the center of an arena after diethylether narcosis. Evidence for the effect is obtained by comparison of the tracks of etherized and non-etherized flies under the influence of olfactory attractant around the center of the arena (Fig. 3). The tracks can be distinguished by their mean radial distance from the central district of the arena. ‘Centrophobia’ denotes the relative difference of the distances of etherized flies and non-etherized controls (Fig. 4). Etherized flies avoid the center of the arena in spite of the attraction of olfactory, thermal or visual cues. The avoidance is significant even in the absence of conspicuous sensory cues for the discrimination of center and surround. The centrophobia obtained in the arena can be used to estimate the efficacy of attractants in the non-etherized control flies (Figs. 6, 7). The lowest possible dose of ether sufficient to elicit narcosis is sufficient to induce centrophobia. None of the other prevalent insect anaesthetics, CO2, N2 and cold, substitutes ether in the present experiments (Figs. 8, 9). Centrophobia arises immediately after ether narcosis. Once induced the effect lasts apparently undiminished for the life time of the flies (Fig. 9). Centrophobia has been found in either sex of the 9 strains tested so far (Fig. 5). Four strains including mutants deficient in wing formation (vestigial) or learning (dunce) show either temporal decline or partial suppression of centrophobia. The anomalous properties are actually due to enhanced spontaneous centrophobism in the non-etherized control groups of these strains (Fig. 10). no notspecified http://www.kyb.tuebingen.mpg.de/ published 8 15017 15423 554 3 KG Götz R Biesinger 1985-05-00 3 156 329 337 Journal of Comparative Physiology A Ether-induced avoidance of the center of an arena by a walking fly does not seem to be the outcome of at least two post-narcotic effects of ether vapor: the inactivation of acetylcholinesterase (Gage et al. 1979), and the inactivation of locomotion (van Dijken et al. 1977). The latter is actually due to a change in the action pattern of search and search control. The centrophobism arising either irreversibly in response to ether treatment, or reversibly in the course of accommodation to a new territory, increases the probability of brief stops at the outer boundary of the arena (Figs. 2–4). Acquisition and maintenance of ‘orientedness’ by exploration of the available territory or evaluation of sensory aids to orientation appears indispensible if a fly wants to avoid the center of the arena. However, centrophobism can be explained without assumption of voluntary behavior. Persistence of direction during random walk in the arena is sufficient to divert locomotor activity from the center to the surround (Figs. 5, 6). The centrophobism found, so far, is equivalent to a ‘mean free path’ of about 4 cm in etherized flies, and about 2 cm in non-etherized flies (Table 1). Search control by variation of persistence in the track of a fly is compatible with results obtained in other insects. no notspecified http://www.kyb.tuebingen.mpg.de/ published 8 15017 15423 1161 7 G Heide M Spüler KG Götz K Kamper Hamburg, Germany1985-00-00 215 222 Symposium 4.5 from the XVII. International Congress of Entomology 1984 no notspecified http://www.kyb.tuebingen.mpg.de/ published 7 15017 15423 1271 7 R Hengstenberg Wien, Austria1985-06-00 228 78. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1271.pdf published -228 15017 15423 555 10 KG Götz 551 3 KG Götz C Wehrhahn 1984-11-00 2 51 129 134 Biological Cybernetics Specialized networks of movement detectors in the antero-inferior field of the eyes of the fruitfly, Drosophila melanogaster, and the housefly, Musca domestica, respond to upward (or downward) drift of the retinal images by excitation (or inhibition) of the lift-generating force of flight. The influence of the direction of pattern movement upon the altitude control response has been investigated under conditions of fixed flight in still air. Matched model analysis of the available response curves suggests the predominance of unidirectional movement detectors in these networks. Homologous wingbeat-inhibiting detectors in the specified fields of the eyes of the two species respond preferentially to pattern movement from antero-superior to postero-inferior. The arrangement of wingbeat-exciting detectors seems to follow different schemes: These detectors respond preferentially to movement from inferior to superior in Drosophila, and to movement from antero-inferior to postero-superior in Musca. The wingbeat-exciting network in Musca is restricted to a comparatively small antero-equatorial area of the specified fields of the eyes. The combination of the two types of detectors in this area establishes a powerful lift control system which is particularly sensitive to minute deviations from a given level of flight. no notspecified http://www.kyb.tuebingen.mpg.de/ published 5 15017 15423 552 3 KG Götz U Wandel 1984-11-00 2 51 135 139 Biological Cybernetics Drift of the retinal images of the surroundings elicits optomotor responses of flight control in the fruitfly, Drosophila melanogaster, and in the housefly, Musca domestica. The present investigation deals with the responses of tethered flies in still air. The responses were elicited by continuous movement of striped patterns in front of the eyes, and characterized by the magnitude and elevation of the resulting force of flight which is the average of the forces produced during a wingbeat cycle. The force of flight is resolved into the upward directed lift and the forward directed thrust. In either species, pattern movement acts upon the magnitude, but not upon the elevation of the force of flight. The elevation relative to the longitudinal body axis is almost invariably 24° in Drosophila, and 29° in Musca. The lift/thrust ratio in still air is fixed accordingly, and can be changed only by variation of the body angle. Keeping an angle of minimum body drag does not contribute significantly to the efficiency of insect flight at very low Reynolds numbers (Re). Control of the lift/thrust ratio by variation of the body angle is, therefore, less surprising in Drosophila where Re is in the order of 102, than in Musca, where Re is in the order of 103. Control of this ratio without variation of the body angle is actually established in insects flying at even higher Re. Covariance of lift and thrust in the investigated flies is achieved by control of wingbeat amplitude or wingbeat frequency, but not by control of wing pitch or stroke plane. A change in the latter parameters would have deflected the force of flight and is, therefore, inconsistent with the constant elevation found in the present experiments. The results obtained, so far, do not exclude active deflections of the force vector during occasional bouts of aerobatics, or passive deflections of this vector during flight at non-zero airspeed. no notspecified http://www.kyb.tuebingen.mpg.de/ published 4 15017 15423 1270 2 R Hengstenberg Springer Berlin, Germany 1984-00-00 121 134 Localization and Orientation in Biology and Engineering no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1270.pdf published 13 15017 15423 1160 7 KG Götz Bonn, Germany1983-05-00 83 99 76. Jahresversammlung der Deutschen Zoologischen Gesellschaft Researchers have studied the stimulus-response characteristics of insects for many years. They try to obtain insight into the mechanisms of visual processing and the interaction of sensory input and motor output. The fly is one of the best-studied insects for this purpose. Of particular interest is the selection of appropriate behaviours in response to visual input. This thesis focuses on the selection of two closely related behaviours in the fly: optomotor response and object fixation. no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 1085 7 KG Götz Saarbrücken, Germany1983-00-00 21 34 Symposium Physiology and Biophysics of Insect Flight 1982 no notspecified http://www.kyb.tuebingen.mpg.de/ published 13 15017 15423 848 2 R Hengstenberg HH Bülthoff B Hengstenberg Springer Berlin, Germany 1983-00-00 183 205 Functional neuroanatomy no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf848.pdf published 22 15017 15423 1188 7 KG Götz R Biesinger 1983-10-00 39 Drosophila Information Service no notspecified http://www.kyb.tuebingen.mpg.de/ published -39 15017 15423 1187 7 KG Götz Bonn, Germany1983-05-00 244 76. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de/ published -244 15017 15423 1269 7 R Hengstenberg Bonn, Germany1983-05-00 246 76. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1269.pdf published -246 15017 15423 854 3 HH Bülthoff KG Götz M Herre 1982-12-00 4 148 471 481 Journal of Comparative Physiology Movement-induced visual orientation in flies depends largely upon predictable responses which establish simple ldquooptomotor balancerdquo or complex ldquopseudo searchrdquo in the appropriate visual environment. Less conspicuous course diverting spontaneous actions of the flies become important in pattern-induced visual orientation. The apparently stochastic spontaneous actions of the houseflyMusca domestica still allow powerful probabilistic predictions of orientation during stationary flight (Reichardt and Poggio 1981). The predominance of non-stochastic spontaneous actions such as ldquobody saccadesrdquo, focussing and shift of ldquovisual attentionrdquo, plasticity of response components etc. in the fruitflyDrosophila melanogaster (Heisenberg and Wolf 1979–1980) accounts for complementary behavioural options which reduce the relevance of probabilistic predictions of orientation in this fly. The conjecture of ldquocomplementary optionsrdquo is based on a striking antagonism between orientation towards a visual object (fixation), and orientation in the opposite direction (anti-fixation), in the walking fly. Forced choice in a multiple-Y-maze quite definitely elicits fixation in the wild type, and antifixation in the ldquooptomotor blindrdquo mutantomb H31 (Fig. 3). However, these effects cannot be attributed to a continuous predominance of attraction in the wild type and repellence in the mutant. This is shown under comparable conditions of free choice in an arena: The flies of either strain alternate between fixation and anti-fixation of an inaccessible visual object (Fig. 4a), and keep running to and fro between two of these objects in ldquoBuridan's paradigmrdquo (Fig. 4b, c), even if the objects are not alike (Fig. 4d). The sequence of approach, retreat and transition may be repeated a few thousand times to the point of exhaustion (Fig. 5). The process resembles the recurrent alternation of ambiguous figures such as the Necker cube in human perception. The recurrent transition between competitive objects counteracts the accumulation of spontaneous preferences, and is likely to explain the apparent lack of pattern-discrimination under operant and non-operant conditions of continued free choice inDrosophila. The conspicuous dichotomy of fixation and anti-fixation in the same environment is, as yet, incompatible with the phenomenological theory of visually controlled orientation in larger flies. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/recurrent_inversion_of_visual_orientation_in_the_walking_fly_drosophila_melanogaster_854[0].pdf published 10 15017 15423 1265 3 R Hengstenberg 1982-07-00 1-2 6 169 171 Journal of Neuroscience Methods For electrophysiological studies in the fly's optic lobe, a procedure was required to perfuse the brain at controllable rates with oxygenated solutions, while recording intracellularly from visual interneurons. The cornea of the compound eyes has to be left in air, in order not to disturb their optics, and to allow physiological stimulation. When the head capsule is opened to get acces to the brain, air sacs and major tracheal trunks tend to collapse by either of two causes: surface tension may flatten tracheae at too low fluid levels and hydrostatic pressure may flatten tracheae at too high levels. Only within a small range of saline levels ( &lt; ± 30µm ) main tracheae and air sacs were found to remain inflated in the dissected head. Therefore the saline level had to be precisely controlled and stabilized within a few microns. With this procedure, it is possible to maintain the fly's brain (&lt; 5 µl ) alive for more than 24 hrs in a perfused volume of only 15 µl, and to record intracellularly from fibres as small as 2 µm. The same procedure should be applicable wherever small, open volumes have to be precisely perfused, and mechanical or electrical artifacts cannot be tolerated.   no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1265.pdf published 2 15017 15423 1268 3 R Hengstenberg 1982-06-00 2 149 179 193 Journal of Comparative Physiology The common response properties to simple visual stimuli (light impulses, light steps, and movement of simple patterns at different speeds) has been investigated by intracellular recording from Giant Vertical Cells (VS) in the lobula plate of the blowflyCalliphora erythrocephala. The impulse response begins < 10ms after onset of the photoreceptor signal (Fig. 6), and shows several phases which gradually subside within about 0.5 s. Very late events, which would hint at recurrent or far-reaching sidepaths, were not observed. The step response is highly non-linear in that both, the increase and decrease of brightness elicit transient depolarization. The excitatory transients are followed by inhibitory waves (Figs. 7, 8), similar to those observed in impulse responses. The possible significance of this succession of excitation and inhibition is discussed. Vertical movement of arbitrary patterns (dot, edges, bar, and gratings) elicit, invariably and irrespective of contrast polarity, depolarizing responses with downward movement, and hyperpolarizing responses with upward movement (Fig. 10). Both responses increase nonlinearly with contour length (Fig. 11). Possible mechanisms, and the functional significance of such nonlinear summation are discussed. The velocity dependence of movement responses to periodic gratings was investigated at both high and low pattern luminance and contrast. Under these conditions VS-cells respond best at a contrast frequency of ≈ 2 Hz, which corresponds with that of velocity dependent optomotor reactions. These results confirm earlier findings that giant vertical cells have many response properties in common. They are best suited to perceive widefield motion, which occurs when a fly performs rotatory and translatory movements in a resting environment. VS-cells are therefore most likely involved in the visual control of such movements. The present results are not sufficient to indicate which of the VS-cells contribute to which of the optomotor reactions. A subsequent publication will be addressed to these questions. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1268.pdf published 14 15017 15423 1267 3 R Hengstenberg K Hausen B Hengstenberg 1982-06-00 2 149 163 177 Journal of Comparative Physiology 1. The structure of one class of giant tangential neurons in the lobula plate of Calliphora, the &quot;Vertical System (VS)&quot; has been investigated by light microscopy. Different staining and reconstruction procedures were employed to ensure that all existing VS-neurons are revealed. 2. There are 11 VS-cells in a characteristic, and constant arrangement (Fig .2). Each cell covers a particular area of the lobula plate, i.e., a distinct area of the retinotopic input array (Table 2), and therefore has a distinct receptive field. 3. Although VS-cells in general tend to occupy the posterior surface of the lobula plate, only three of them (VS2-VS5) reside exclusively in this layer. The other cells (VS1 and VS6-VS10) have bistratified dendritic arborizations (Fig. 6), whose dorsal parts are apposed to the anterior surface of the lobula plate. 4. The arrangement, territory and stratification of any given VS-cell is largely invariant in different individuals, whereas, the branching pattern may vary considerably (Fig. 3). 5. The present results provide the framework for physiological studies of the role of individual VS-cells in movement perception, and their involvement in the control of particular locomotor behaviour.&nbsp; no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1267.pdf published 14 15017 15423 1266 7 R Hengstenberg DC Sandeman Hannover, Germany1982-06-00 313 75. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1266.pdf published -313 15017 15423 1262 3 R Hengstenberg 1981-10-00 3 4 249 255 Journal of Neuroscience Methods A small piezolectric device for cell penetration is described. It retracts the micropipette slowly by electrostriction, and pushes it very fast (&lt;5µsec) forward by shorcircuiting the transducer. The design, operation circuit, and performance under test conditions are described. Penetration examples from small nerve fibres (&lt;5µm) show that membrane puncture occurs only with the fast forward push. Cells are not noticeably damaged, even if the device is repeatedly operated after cell penetration. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1262.pdf published 6 15017 15423 860 3 HH Bülthoff 1981-08-00 2 41 139 145 Biological Cybernetics Drosophila melanogaster is able to detect a small visual object hidden in a background of identical texture, as long as there is relative motion between their retinal images. The properties of figure-ground discrimination in the walking fly are studied under experimental conditions where the positions of figure and ground oscillate sinusoidally with similar frequency and similar amplitude but with different phase. The following points have been established. (a) The average turning reaction of the stationarily walkingDrosophila depends on phase; contrary to results obtained with the flyingMusca (Reichardt and Poggio, 1979), antiphasic oscillation of figure and ground does not suppress the attrativeness of the figure. (b) A translatory response has been found which also depends on the phase difference of the oscillatory movements of figure and ground. (c) The time course of the responses and its intra- and inter-individual variability do not seem to fit into a rigid model of figure-ground discrimination. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf860.pdf published 6 15017 15423 1264 3 R Hengstenberg DC Sandeman 1981-00-00 1981 Annual Report of the Research School of Biological Sciences: Australian National University The halteres are unique balance organs which stabilise flies during flight. They are known to elicit compensatory head movements when the flying animals are rotated about yaw. With imposed roll, flying animals perform compensatory head movements, controlled by mechanosensory and visual inputs. Removal of halteres, and of visual cues, abolishes such head movements. Unilateral haltere ablation suggests that, as in other compensatory systems, the inputs from the two sides are poised against each other. The classical analogy between a gyroscope and the fly halteres is revised in the light of these findings, and the halteres are better described as rapidly oscillating pendulums. The physical properties are subtly but significantly different from a gyroscope, and afford the animal a way of monitoring motion about all three axes by one sense organ without confusion. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1264.pdf published 0 15017 15423 1263 7 R Hengstenberg Bremen, Germany1981-06-00 180 74. Jahresversammlung der Deutschen Zoologischen Gesellschaft no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1263.pdf published -180 15017 15423 1261 2 R Hengstenberg B Hengstenberg Springer Berlin, Germany 1980-00-00 308 324 Neuroanatomical Techniques: insect nervous system no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1261.pdf published 16 15017 15423 1084 2 KG Götz Plenum Press New York, NY, USA 1980-00-00 391 407 Development and Neurobiology of Drosophila no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 1258 3 E Buchner S Buchner R Hengstenberg 1979-08-00 4407 205 687 688 Science Adult Drosophila were fed with tritium-labeled deoxyglucose prior to a 5-hour period of visual stimulation. A flickering disk of light and a moving grating were presented to the left and right eyes, respectively. Autoradiography revealed enhanced labeling solely in that part of the second optic ganglion (medulla) whose visual field was stimulated by movement. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1258.pdf published 1 15017 15423 865 3 HH Bülthoff KG Götz 1979-04-00 5705 278 636 638 Nature Since Wertheimer's classic paper1, research in motion perception has been concerned with the study of visual illusions such as phi-motion. Various phenomena of this type are easy to elicit by successive changes of the light flux in spatially distinct photoreceptors, and easy to explain by the specific properties of the motion detectors, although there are reports to the contrary2. The present account deals with a less easily comprehensible illusion which is elicited by simultaneous changes of the light flux in differently illuminated receptors3−5. The phenomenon has previously been ascribed to the prolonged latency of the 'light-on' responses at lower levels of illumination which converts simultaneous stimuli into successive signals6, but this does not explain the illusion satisfactorily3. MacKay and co-workers were the first to attribute the apparent motion to the adaptive properties of the input channels of the motion detectors4. We show that the illusion can be induced in the fruitfly, Drosophila melanogaster, as well as in man. The course control response to motion provides a quantitative assay of the illusion in the fly. The results suggest that the illusion originates in the distortion of the visual signals before motion detection. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/analogous_motion_illusion_in_man_and_fly_865[0].pdf published 2 15017 15423 1259 7 E Buchner S Buchner R Hengstenberg Roma, Italy1979-09-00 S132 Third European Neuroscience Meeting no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1259.pdf published 0 15017 15423 512 3 E Buchner KG Götz C Straub 1978-12-00 4 31 235 242 Biological Cybernetics Optomotor thrust responses of the fruitfly Drosophila melanogaster to moving gratings have been analysed in order to determine the arrangement of elementary movement detectors in the hexagonal array of the compound eye. These detectors enable the fly to “perceive” vertical movement. The results indicate that, under photopic stimulation of a lateral equatorial eye region, the movement specific response originates predominantly from two types of elementary movement detectors which connect neighbouring visual elements in the compound eye. One of the detectors is oriented vertically, the other detector deviates 60° towards the anterior-superior direction (Fig. 5b). The maximum of the thrust differences to antagonistic movement is obtained if the pattern is moving vertically or along a superior/anterior — inferior/posterior direction 30° displaced from the vertical (Fig. 3d,e, Fig. 6). Only one of the detectors coincides with one of the two detectors responsible for horizontal movement detection. This indicates that a third movement specific interaction in the compound eye of Drosophila has to be postulated. — The contrast dependence of the thrust response (Fig. 2) yields the acceptance angle of the receptors mediating the response. The result Δϱ coincides with the acceptance angle found by analysis of the turning response of Drosophila (Heisenberg and Buchner, 1977). This value corresponds to the acceptance angle expected, on the basis of optical considerations, for the receptor system R 1–6. — The movement-specific neuronal network responsible for thrust control is not homogeneous throughout the visual field of Drosophila. Magnitude and preferred direction of the thrust response in the upper frontal part of the visual field seem to vary considerably in different flies (Fig. 6). no notspecified http://www.kyb.tuebingen.mpg.de/ published 7 15017 15423 513 3 KG Götz E Buchner 1978-12-00 4 31 243 248 Biological Cybernetics Optomotor control of course and altitude in the fruitfly, Drosophila melanogaster, requires dense networks of elementary movement detectors (EMD's) which cover most if not all of the visual field. The predominant types of EMD's in these networks represent interactions between neighbouring visual elements along the three main directions of the hexagonal array in the compound eye. — Course control in the walking fly is achieved mainly by pairs of equivalent EMD's which occupy 2 o'clock and 4 o'clock positions with respect to the right eye (Buchner, 1976). Comparison of the turning response and the torque response in the present account confirms the particular properties of this network, and proves the presumed bidirectional sensitivity of its EMD's for the course control responses of legs and wings in the corresponding modes of locomotion. — Altitude control during flight is achieved by a less homogeneous network of EMD's which modifies lift and thrust simultaneously by the appropriate control of the wing beat amplitudes. The predominant types of EMD's in the lateral eye regions occupy 12 o'clock and 2 o'clock positions with respect to the right eye (Buchner et al., 1978). The present evaluation of the optomotor responses of thrust and wing beat confirms the preferred orientation of these EMD's and discloses a pecularity of their internal structure. The movement detectors of this network lack the bidirectional sensitivity of the EMD's in the course control system. At least the fronto-lateral network of the altitude control system seems to consist mainly of pairs of equivalent unidirectional EMD's. The detectors in 12 o'clock position increase wing beat in response to movement of the visual surroundings from inferior to superior. The opposite effect is produced by the detectors in 2 o'clock position which respond to movement from anterior-superior to posterior-inferior. These properties qualify unidirectional EMD's as the functional units of the optomotor control system in the fruitfly. Pairs of unidirectional antagonists would be sufficient to establish the bidirectional sensitivity found in the movement detectors of the course control system. no notspecified http://www.kyb.tuebingen.mpg.de/ published 5 15017 15423 1186 3 KG Götz 1978-06-00 53 167 167 Drosophila Information Service no notspecified http://www.kyb.tuebingen.mpg.de/ published 0 15017 15423 1174 KG Götz 1978-00-00 no notspecified http://www.kyb.tuebingen.mpg.de/ published 15017 15423 1257 3 R Hengstenberg 1977-11-00 5635 270 338 340 Nature Our understanding of information processing in nerve nets has been modified by the concept of graded signal transmission. Descriptions of non-spiking interneurons in insects, and the demonstration of graded synaptic transmision have contributed to this development. In the fly visual nervous system, second and higher order interneurones are known, which apparently do not produce action potentials. We show here that at least eight individually identifiable movement-sensitive cells, which have the characteristic properties of non-spiking interneurones,&nbsp; will generate spikes with imposed hyperpolarisation. Their graded mode of operation is due to maintained refractoriness. This applies selectively to neurones, which belong to either of&nbsp; two anatomically, and physiologically distinct classes. Ohter cell types in the same preparation generate spikes spontaneously. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1257.pdf published 2 15017 15423 511 3 KG Götz 1977-00-00 32 125 132 Zeitschrift für Naturforschung C no notspecified http://www.kyb.tuebingen.mpg.de/ published 7 15017 15423 1083 7 T Poggio E Bizzi RE Eckmiller AM Graybiel KG Götz M Ito MF Land FA Miles WE Reichardt DA Robinson DC Sandemann PH Schiller G Westheimer Berlin, Germany1977-03-00 309 327 Dahlem Workshop on Function and Formation of Neural Systems no notspecified http://www.kyb.tuebingen.mpg.de/ published 18 15017 1542315017 15423 1082 7 KG Götz Aarau, Switzerland1977-00-00 10 33 155. Jahresversammlung der Schweizerischen Naturforschenden Gesellschaft 1975 no notspecified http://www.kyb.tuebingen.mpg.de/ published 23 15017 15423 1173 KG Götz 1976-11-00 A mechanical-electrical transducer for the simultaneous contact-free and reaction free determination of a plurality of translatory and/or rotary components of the motion or position of an object coupled to the transducer, with a computer circuit which generates signals corresponding in sign and magnitude to the components of the deviation of the object from a predetermined initial position. no notspecified http://www.kyb.tuebingen.mpg.de/ published 15017 15423 508 3 KG Götz 1975-10-00 10 62 468 475 Naturwissenschaften no notspecified http://www.kyb.tuebingen.mpg.de/ published 7 15017 15423 510 3 KG Götz 1975-09-00 3 99 187 210 Journal of Comparative Physiology In his discussion of the optomotor behaviour Kalmus claimed in 1964 that the visual systems of insects continuously resolve horizontal displacements relative to the surroundings into rotatory and translatory components, each associated with optomotor feedback of particular quality and sign. The feedback is supposed to achieve, simultaneously, minimization of the rotatory movements and maximization of the translatory movements, a behaviour repeatedly observed with actively moving insects such as the fruitflyDrosophila melanogaster. The present approach takes into account that the output of movement detectors in the visual system of insects is necessarily equivocal with respect to the speed of the stimulus (e.g. zero output at both zero and infinite speed). Decomposition of the stimulus is not feasible under these conditions. It is obviously the composite stimulus to which the insects respond. Moreover, there is experimental evidence that optomotor feedback on the translatory movement is not necessarily a response-determining factor in insects. The optomotor behaviour of the walking fruitfly is sufficiently described by the sum of itsrotatory responses to the composite stimuli on either side. A diagram representing the expected rotatory response of the walking fruitfly as a function of both the rotatory and the translatory stimulus component is used to derive the prevailing traits of the behaviour in resting, rotating and floating environments, respectively. Most conspicuous is the inversion of the course-control response in about one half of the possible states of stimulation. This effect gives rise to at least some of the apparently spontaneous turns of actively moving insects which have been ascribed by v. Holst and Mittelstaedt to efferent commands from higher centres of the brain, according to their principle of reafference. The present results merely disprove the necessity of these commands. Inversion of the response is also an inherent property of the course-control systems of the optomotorically active insects. The expected increase of these inversions with closer proximity of the visual environment is found by observation of walking fruitflies. The relation between the rotatory and translatory movements of the freely walking fly and its state of stimulation in a given environment is used to describe the expected behaviour in terms of the most probable transition of state. The approach is based on estimates of the power required by the fly in order to maintain a given state against the torque that is produced by its course-control system in response to the optomotor stimulation. The most probable transition of state is apparently determined by the tendency of the fly to decrease the power requirement by appropriate adaptation of its rotatory movement. The transition may come to an end in one of the states of minimu no notspecified http://www.kyb.tuebingen.mpg.de/ published 23 15017 15423 509 3 M Heisenberg KG Götz 1975-09-00 3 98 217 241 Journal of Comparative Physiology Partially blind mutants can be used to investigate the processing of visual information in the fruit flyDrosophila. This approach requires (1) procedures for the selection of a variety of partially blind mutants, and (2) a strategy for the identification and coordination of visual malfunctions by comparison of interrelated traits of behaviour. The two selection techniques so far employed to recover partially blind mutants use either the fast phototaxis or the optomotor response as selection determining behaviour. The second method is described here and is applied specifically to select mutants in which one of the two autonomous subsystems of vision designated asHigh Sensitivity System andHigh Acuity System is defective. (The mutants obtained are apparently normal with respect to their HAS whereas the HSS is blocked.) Two sets of experiments have been developed in order to test interrelated traits of behaviour in a comparatively large number of flies. One set of experiments measuresslow phototaxis as a function of light intensity. The other is to determine the optomotor response to moving patterns of different spatial periods as functions of both the average brightness and the speed of the movement. Further techniques such as electroretinography and optical inspection of the eyes are used to complement the behavioural approach. By combination of the different tests a first step has been made in the characterization and classification of partially blind mutants with neuronal disorders obtained by different selection procedures and in different laboratories. no notspecified http://www.kyb.tuebingen.mpg.de/ published 24 15017 15423 1256 7 R Hengstenberg 1975-00-00 Experimental Brain Research no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1256.pdf published 0 15017 15423 1254 3 R Hengstenberg 1973-09-00 9 28 593 597 Zeitschrift für Naturforschung C Action potentials of some of the 3578 nerve fibres in the cervical connective of Drosophila melanogaster can be recorded with extracellular electrodes. The spike rate increases if moving striped patterns are presented to the compound eyes, especially with horizontal front-to-back movement. The response is small or absent with the reverse (or with vertical) direction of movement. The main properties of this response are described, and briefly discussed. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1254.pdf published 4 15017 15423 507 3 KG Götz H Wenking 1973-09-00 3 85 235 266 Journal of Comparative Physiology To investigate the optomotor leg responses ofDrosophila melanogaster the free walking fly is kept in stationary orientation and position on top of a ball. The stimulus consists of continuous pattern movement in the equatorial zone of the visual field. The rotatory and translatory responses are derived from the signals of a servo-system which maintains the stationary state of the walking fly by appropriate rotations of the ball. Course control, or the tendency to follow rotatory displacements of the visual surroundings, is established in the wild type and in a behavioural mutant with the eye colour markerw a.The movement detecting systems in the complex eyes on either side respond to the horizontal component of the stimulus and control, predominantly, the thrust of the ipsilateral legs in the wild type, and the thrust of the contralateral legs in the mutant. As a result, front-to-back movement is decreasing the walking speed of the wild type and increasing the walking speed of the mutant, andvice versa. The course control responses of flying and walking fruitflies depend on the signals of movement detecting systems which are equivalent with respect to the horizontal orientation, the dynamic range, and the resolving power. Leg responses show that the orientation of the movement detecting systems is independent of their position in the eye, and is invariant to the direction and velocity of the stimulus. The lift control response to the vertical component of the movement stimulus is a quality of the flight system. The response has no counterpart in the optomotor behaviour of the stationarily walking fruitfly. Functional specialization of the different pairs of legs is not detectable in the present experiments. The rotatory response as well as the translatory propulsion are almost equally accomplished by the fore legs, the middle legs and the hind legs of partially amputated fruitflies. The optomotor reactions of the fruitfly are accompanied by at least two side-effects of the visual stimulus: The flicker effect acts on the walking speed. The effect is elicited by the temporal sequence of bright and dark stripes in the receptive fields of the visual elements. It is independent of the direction of the pattern movement, and can be produced by a stationarely flickering light source. An after-effect of the pattern movement appears at the end of the visual stimulation. The effect depends on experimental parameters. An initial magnitude of -1/16 of the preceding course control response and a decay time of about 7 min were observed in the present experiments. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/R11_507[0].pdf published 31 15017 15423 1081 7 KG Götz Cambridge, MA, USA1973-00-00 491 507 Third International Biophysics Congress of the International Union for Pure and Applied Biophysics (IUPAB 1969) no notspecified http://www.kyb.tuebingen.mpg.de/ published 16 15017 15423 1255 7 R Hengstenberg 1973-12-00 149 Drosophila Information Service no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1255.pdf published -149 15017 15423 1031 3 KG Götz 1972-00-00 82 411 417 Bibliotheca Ophthalmologica no notspecified http://www.kyb.tuebingen.mpg.de/ published 6 15017 15423 1253 2 R Hengstenberg Springer Berlin, Germany 1972-00-00 93 96 Information Processing in the Visual Systems of Anthropods “Clock-spikes” in Musca are produced by a motoneurone which lies in the subesopha-geal ganglion. It innervates a small muscle, attached to the inner frontal margin of the retina. In the frontal eye region, muscular activity causes the distal rhabdomere tips to move perpendicular to the ommatidial axes. Slow angular movements of the optical axes of the retinula cells result herefrom. The spike rate can be slightly changed by a variety of gross visual stimuli. The adequate visual stimulus as well as the functional significance of the eye muscle system with respect to the behaviour of the unrestrained insects are still unknown. no notspecified http://www.kyb.tuebingen.mpg.de/ published 3 15017 15423 1032 2 KG Götz Springer Berlin, Germany 1972-00-00 255 263 Information Processing in the Visual Systems of Anthropods The properties of the navigation system are derived from the open loop optomotor reactions of the fruitfly Drosophila which is either flying or walking under stationary conditions. The horizontal component of a movement stimulus controls the difference of the propulsive forces of legs and wings on either side and enables the freely moving fly to counteract involuntary deviations from a straight course. The vertical component controls the sum of the propulsive forces of the wings and enables the fly to maintain a given level of flight. Different properties of the navigation system are found in normal and mutant fruitflies. Comparative studies of open loop and closed loop behaviour disprove the separation theorem of KALMUS according to which the fly is always minimizing the rotatory component, and maximizing the translatory component, of a movement stimulus. There is evidence that inversion of the rotatory response can be elicited by the translatory component of the stimulus. Interaction of the stimulus components must be considered in order to determine the closed loop responses of the freely moving insect. no notspecified http://www.kyb.tuebingen.mpg.de/ published 8 15017 15423 1185 7 KG Götz 1972-03-00 62 Drosophila Information Service no notspecified http://www.kyb.tuebingen.mpg.de/ published -62 15017 15423 1252 1 B Katz F-W Bentrup R Hengstenberg Thieme Stuttgart, Germany 1971-00-00 no notspecified http://www.kyb.tuebingen.mpg.de/ published 164 15017 15423 1237 3 R Hengstenberg 1971-08-00 2 9 56 77 Kybernetik It is possible to record spontaneously occurring impulses in the housefly's optic lobe region. These closely resemble “clock-spikes”, as described for Calliphora by Kuiper and Leutscher-Hazelhoff (1965). The repetition rate of these impulses—here called “C-spikes”—is about 45/s at 20° C and increases with temperature. Between 15 and 35° C the temperature coefficient of the repetition rate is close to Q 10=2. At constant temperature the mean rate is constant for many hours, the individual intervals appear to be gaussian-distributed about the mean interval τˉ . The standard deviation of the interval lengths in samples of >10000 impulses is approximately ±2.5% of the mean. The fluctuation corresponds to a slight modulation of the mean spike frequency by a noise signal, comprising slow as well as fast components. The time course of extracellularly recorded spikes in combination with evidence from simultaneous recordings at different sites shows that typical C-spikes are produced by the subsequent activity of at least two distinct sources: “Prespikes” originate in the midbrain and are centrifugally conducted with about 2 m/s at room temperature to a peripheral site of C-spike-activity, where they induce a strictly event-correlated impulse activity of a “postspike” source. Decapitation shows that all elements that are necessary to produce and to maintain the regular C-spike activity are located within the head. Under constant conditions no interaction is observed between C-spike sources on the left and right side of the head. Intracellular recordings show that the membranes of the postspike sources on either side are of the electrically unexcitable type. Each of the postspike sources is formed by a cluster of at least two cells. Electrophysiological localization experiments indicate that the postspike sources are located outside of the optic lobes, but close to the lower frontal margin of the left-and right-hand medulla. The sources of C-spike activity could be identified by histological localization of the recording sites. The anatomical correlate of the electrophysiologically determined C-spike system has been reconstructed by means of silver impregnated serial sections: In the lateral perikaryon layer on either side of the subesophageal ganglion lies a single large motoneurone, which is spontaneously producing the regular impulses, most probably during the entire life time of the fly. These impulses are centrifugally conducted along a thin peripheral nerve, which only contains a single motor axon of 6 μm diameter. The nerve runs to a very small muscle, consisting of 14–20 tubular skeletal muscle fibres of 7–10 μm diameter. These fibres are innervated by numerous grape-like neuromuscular endings. From this unineuronal, multiterminal innervation it is concluded that the muscle acts as a functional unit. Extracellular and intracellular recordings under microscopic observation prove the identity of the muscle fibres with the source of the postspikes. The muscle has not been previously described for Musca. It is shown that one end of the muscle is inserted at the inner margin of the orbital ridge, i.e. at the base of the frontal ommatidia in the vicinity of the equator of the compound eye. The other end is fixed to an apodeme which originates near the foramen occipitale on the ventral occipital ridge and which most probably is homologous with the tentorium of other insects. Hence the muscle is denoted as Musculus orbitotentorialis. Similar muscles with comparable insertions are found in Calliphora and Drosophila. The orbito-tentorial muscle also exists in Eristalis, where the tentorium is well developed. Here the muscle inserts on the anterior tentorial arm and at the inner margin of the orbital ridge. This muscle also produces continuously regular spikes. The structure of the head skeleton of Musca shows that the tentorial insertion of the muscle is relatively rigid. Since antagonistic muscles are obviously missing, it is concluded that the orbito-tentorial muscle acts against the elastic forces of the eye tissue and of the orbital skeleton. It is conceivable that the muscular action causes displacements of the optic axes of the visual elements in the compound eyes. The physiological meaning of these displacements is still obscure and deserves further investigations. no notspecified http://www.kyb.tuebingen.mpg.de//fileadmin/user_upload/files/publications/pdf1237.pdf published 21 15017 15423 506 3 KG Götz 1970-04-00 2 52 419 436 Journal of Experimantal Biology 1. The optomotor control of orientation and locomotion in the fruitfly Drosophila melanogaster requires the conveyance of information from distinct movement detectors in the visual system to distinct movement effectors in the motor system. Abnormalities of the optomotor control system have been found occasionally in Drosophila. 2. The abnormal flies can be isolated from population samples by appropriate fractionation according to the magnitude and the sign of the optomotor responses. A cyclically operating machine was used to fractionate two inbred strains, w+ and wα, which possess different alleles on the white-locus of their X-chromosomes. 3. Movements of an artificial visual environment elicit similar orientation-control responses, but antagonistic locomotion-control responses in the two strains. The responses depend on various parameters and may even change with habituation to the stimulus. However, the application of selection pressure through eight generations has little if any effect on the different optomotor behaviour of the inbred strains. no notspecified http://www.kyb.tuebingen.mpg.de/ published 17 15017 15423 1184 7 KG Götz 1970-02-00 105 Drosophila Information Service no notspecified http://www.kyb.tuebingen.mpg.de/ published -105 15017 15423 1030 7 KG Götz Varenna, Italy1969-00-00 494 509 International School of Physics "Enrico Fermi": Course XLIII, 1968 no notspecified http://www.kyb.tuebingen.mpg.de/ published 15 15017 15423 1029 41 KG Götz