Table Of ContentInformation Processing
in the Visual Systems
of Arthropods
Symposium Held at the Department of Zoology
University of Zurich, March 6-9, 1972
Edited by
RUdiger Wehner
With 263 Figures
Spri nger-Verlag
Berlin Heidelberg New York 1972
The cover shows a scanning electron micrograph of the compound eye of a desert ant,
Cataglyphis bicolor (University of Zurich).
ISBN-13: 978-3-540-06020-8 e-ISBN-13: 978-3-642-65477-0
001: 10.1007/978-3-642-65477-0
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© by Springer·Verlag Berlin' Heidelberg 1972. Library of Congress Catalog Card Number 72-91887.
Preface
It is now generally accepted for a variety of reasons - morphological as well as physiological -
that the visual systems of arthropods provide a suitable model for the study of information proces
sing in neuronal networks. Unlike the neurophysiology of the visual pathway in the frog and the
cat which is more than adequately documented, recent work on the compound eye and optical
ganglia of spiders, crustaceans, and insects has scarcely been summarized. In order to fill this
void so that others, especially vertebrate neurophysiologists may become familiar with the advan
tages of these systems, our group at Zurich University organized here in March 1972, a European
meeting to discuss the anatomical.! neurophysiological and behavioral knowledge on the compound
eye and the visual. pathway of arthropods. Systems analysis was regarded as the main theme of the
conference, but systems analysis of a network of neurons cannot be done as a mere "black-box"
maneuver. The conference therefore tried to reconcile neurophysiology and behavioral analysis
in order to make predictions about a necessary and sufficient neural structure. The "wiring dia
grams" of such a structure might then be confirmed histologically. Hence the aim of the conferen
ce was not to deal only with the structure and function of the compound eye - i.e. data processing
in the dioptric apparatus and in the layer of the retinular cells - but to apply this knowledge to
the performance of the visual system in color discrimination, pattern recognition, and other cent
ral nervous processes.
Although the official language of the symposium was German, the contributors wrote their manus
cripts in English, also reflecting the extensive discussions that arose both during and after the for
mal sessions. We thus hope to make any insights derived from the discussions available to those
unable to attend.
Besides the authors, we would like to thank all the 77 participants of the symposium for their li
vely contributions both inside the lecture hall and outside. The discussions allowed the contribu
tors to cross-check recent knowledge and made it clear that the visual systems of arthropods have
much to contribute to neurophysiology. Much enthusiasm was generated for planning the further
work along these lines. I wish most of all to express our appreciation to the Swiss National Found
ation for Scientific Research, which provided generous financial support for the meeting, and to
Springer-Verlag for the very rapid publication of the proceedings.
Zurich, June 1972 RUdiger Wehner
Contents
Preface III
List of Participants IX
Opening Remarks
I.
ANATOMY OF THE VISUAL SYSTEM
1. V. BRAITENBERG
Periodic Structures and Structural Gradients in the Visual
Gangl ia of the Fly 3
2. C.B. BOSCHEK
Synaptology of the Lamina ganglionaris in the Fly 17
3. N.J. STRAUSFELD and J.A. CAMPOS-ORTEGA
Some Interrelationships between the First and Second Synaptic
Regions of the Fly's (Musca domestica) Visual System 23
4. J.A. CAMPOS-ORTEGA and N.J. STRAUSFELD
Col umns and Layers in the Second Synaptic Region of the
Fly's Visual System: The Case for Two Superimposed Neuronal
Arch i cetures 31
5. R. MENZEL
The Fine Structure of the Compound Eye of Formica polyctena -
Functional Morphology of a Hymenopterean Eye 37
6. P.L. HERRLlNG
Measurements on the Arrangement of Ommatidial Structures
in the Retina of Cataglyphis bicolor (Formicidae, Hymenoptera) 49
7. H.F. PAULUS
The Ultrastructure of the Photosensible Elements in the Eyes of
Collembola and Their Orientation (Insecta) 55
II.
OPTICS OF THE COMPOUND EYE
1. K. KIRSCHFELD
The Visual System of Musca: Studies on Optics, Structure and
Function 61
2. N. FRANCESCHINI
Pupil and Pseudopupil in the Compound Eye of Drosophila 75
VI
3. W.P. EHEIM
Directional Intensity Distribution in Single Ommatidia
of the Honeybee, Apis mell ifera 83
4. P. KUNZE
Pigment Migration and the Pupil of the Dioptric Appa-
ratud in Superposition Eyes 89
5. R. HENGSTENBERG
Eye Movements in the Housefly Musca domestica 93
III.
BIOCHEMISTRY OF VISUAL PIGMENTS
1. K. HAMDORF, R. PAULSEN, J. SCHWEMER and U. TAEUBER
Photoreconversion of Invertebrate Visual Pigments 97
2. H. LANGER
Metarhodopsin in Single Rhabdomeres of the Fly, Call iphora
erythrocepha la 109
IV.
INTENSITY - DEPENDENT REACTIONS
l. T. LABHART
The Discrimination of Light Intensities in the Honey Bee 115
2. M. FRISCHKNECHT
Photopositive Reactions of Honey Bees to Circular Areas of
Varying Sizes and Light Intensities 121
3. P. STRECK
Screening Pigment and Visual Field of Single Retinula Cells
of Call iphora 127
4. G. FLEISSNER
Circadian Sensitivity Changes in the Median Eyes of the
North African Scorpion, Androctonus austral is 133
5. E. BUCHNER
Dark Activation of the Stationary FI ight of the Fruitfly
Drosophila 141
V.
WAVELENGTH - DEPENDENT REACTIONS
1. D. BURKHARDT and I:DE LA MOTTE
Electrophysiological Studies on the Eyes of Diptera, Mecop-
tera and Hymenoptera 147
2. R. SCHUEMPERLI
Wavelength-specific Behavioral Reactions in Drosophila
melanogaster 155
VII
3. W. KAISER and E. LISKE
A Prel iminary Report on the Analysis of the Optomotor
System of the Bee - Behavioral Studies with Spectral
Lights 163
4. W. KAISER
A Prel iminary Report on the Analysis of the Optomotor
System of the Honey Bee - Single Unit Recordings du-
ring Stimulation with Spectral Lights 167
5. F. TOGGWEILER
Spectral Sensitivity and Wavelength Discrimination in
Cataglyphis bicolor (Formicidae) 171
6. H. ROTH and R. MENZEL
ERG of Formica polyctena and Selective Adaptation 177
VI.
PATTERN RECOGNITION
1. R. WEHNER
Pattern Modulation and Pattern Detection in the Visual
System of Hymenoptera 1-83
2. B. SCHNETTER
Experiments on Pattern Discrimination in Honey Bees 195
3. H. CRUSE
A Qualitative Model for Pattern Discrimination in the
Honey Bee 201
4. A. ANDERSON
The Ability of Honey Bees toGeneralise Visual Stimuli 207
5. W. REICHARDT
First Steps in a Behavioral Analysis of Pattern Discrimi-
nation in Diptera 213
6. F. ZETTLER and M. JAERVILEHTO
Intraaxonal Visual Responses from Visual Cells and Second-
order Neurons of an Insect Retina 217
7. H.W. MEYER
Ethometrical Investigations into the Spatial Interaction
within the Visual System of Velia caprai (Hemiptera,
Heteroptera) 223
8. M.F. LAND
Mechanisms of Orientation and Pattern Recognition 231
9. F. SCHALLER
Observations on the Yisual Reactiohs of Collembola 249
VIII
VII.
VISUAL CONTROL OF ORIENTATION PATTERNS
1. K.G. GOETZ
Processing of CI ues from the tvbving Environment in the
Drosophila Navigation System 255
2. M. HEISENBERG
Behavioral Diagnostics; a Way to Analyse Visual Mutants
of Drosophila 265
3. P. GOERNER
Resultant Positioning between Optical and Kinesthetic
Orientation in the Spider Agelena labyrinthica clerck 269
4. M.L. MITTELSTAEDT -BURGER
Idiothetic Course Control and Visual Orientation 275
5. P. DUELLI
The Relation of Astromenotactic and Anemomenotactic
Orientation Mechanisms in Desert Ants, Cataglyphis
bicolor (Formicidae, Hymenoptera) 281
6. R. WEILER and M. HUBER
The Significance of Different Eye Regions for Astromeno
tactic Orientation in Cataglyphis bicolor (Formicidae,
Hymenoptera) 287
7. R. WEHNER and I. FLATT
The Visual Orientation of Desert Ants, Cataglyphis bico
lor, by Means of Terrestrial Clues 295
8. A. BURKHALTER
Distance Measuring as Influenced by Terrestrial Clues in
Cataglyphis bicolor (Formicidae, Hymenoptera) 303
VII I.
STORAGE OF VISUAL INFORMATION
1. J. ERBER
The Time-Dependent Storing of Optical Information in
the Honey Bee 309
2. T. MASUHR and R. MENZEL
Learning Experiments on the Use of Side - Specific Infor
mation in the Olfactory and Visual System in the Honey
Bee (Apis mell ifica) 315
IX.
METHODS OF QUANTIFYING BEHAVIORAL DATA
1. O. v. HELVERSEN
The Relationship between Difference in Stimuli and Choice
Frequency in Training Experiments with the Honey Bee 323
List of Partici pants
ANDERSON, A.M., University of Edinburgh, Dept. of Zoology, West Mains Rd., GB-Edinburgh,
EH93JT
BAESSLER, U., Universitat Trier -Kaiserslautern, Naturwiss. -Techn. Fakul tat, D-675 Kaisers-
lautern, Postfach 1019
BAUMANN, F., Universite de Gen~ve, Dept. de Physiologie, CH-1211 Gen~ve 4
BOSCHEK, C.B., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr.38
BRAITENBERG, V., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemann-
strasse 38
BRUNNERT, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
BURKHALTER, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstler
gasse 16
BURKHARDT, D., Universitat Regensburg, Fachbereich Biologie, D-8400 Regensburg, Universitats
strasse 31
BUTENANDT, E., Max-Planck-Institut fUr Verhaltensphysiologie, D-8131 Seewiesen/Obb.
CAMPOS-ORTEGA, J.A., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen,
Spemannstr. 38
CRUSE, H., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
DORNFELD, K., II. Zoologisches Institut der Freien Universitat, D-1 Berlin-41, Grunewaldstr.34
DUELLI, P., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
EHEIM, W.P., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
ERBER, J., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnittspahnstr.3
FLATT, E., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
FLEISSNER, G., Zoologisches Institur der Universitat, D-6 Frankfurt a.M., Siesmayerstr. 70
FRANCESCHINI, N., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemann-
strasse 38
FRISCHKNECHT, M., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstler-
gasse 16
GEIGER, G., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
GOERNER, P., II. Zoologisches Institut der Freien Universitat, D-1 Berlin 41, Grunewaldstr. 34
GOETZ, K.G., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
HAMDORF, K., Ruhr-Universitat Bochum, Lehrstuhl fUr Tierphysiologie, Gebaude ND, 5.0.G.,
D-463 Bochum
HAUSEN, H., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
HEISENBERG, M., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemann
strasse 38
HELVERSEN, O.v., Biologisches Institut I (Zoologie) der Albert-Ludwigs-Universitat, D-78 Frei
burg i.Br., Katharinenstr. 20
HENGSTENBERG, R., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemann-
strasse 38
HERRLlNG, P.L., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
HUBER, M., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
JAERVILEHTO, M., Zoologisches Institut der Universitat MUnchen, D-8 MUnchen 2, Luisenstr. 14
KAISER, W., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnittspahn-
strasse 3
x
KIEPENHEUER, J., Universitat Frankfurt, Zoologisches Institut, D-6 Frankfurt a.M., Siesmayer-
strasse 70
KIRSCHFELD, K., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
KLEIN, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
KRETZ, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
KUNZE, P., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
KURTH, B., II. Zoologisches Institut der Freien Universitat, D-1 Berlin-41, Grunewaldstr. 34
lABHART, T., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
lAND, M., University of Sussex, Biology Bldg., Falmer, Brighton, Suss. BN1 9QG, Great Britain
lANGER, H., Ruhr-Universitat Bochum, Institut fUr Tierphysiologie, Gebaude JC-4/43,
D-463 Bochum, Postfach 2148
LISKE, E., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnittspahnstr. 3
MASUHR, T., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnitt
spahnstr. 3
MENZEL, R., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnitt-
spahnstr. 3
MEYER, L, Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
MEYER, H., Zoologisches Institut der Universitat, D-6 Frankfurt a.M., Siesmayerstr. 70
MITTELSTAEDT, H., Max-Planck-Institut fUr Verhaltensphysiologie, D-8131 Seewiesen/Obb.
MITTELSTAEDT, M., Max-Planck-Institut fUr Verhaltensphysiologie, D-8131 Seewiesen/Obb.
c.,
MONDADORI, Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstler-
gasse 16
MOTTE de la, J., Universitat Regensburg, Fachbereich Biologie, D-8400 Regensburg, Univer
sitatsstr.31
NOWAK, H., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
PAULUS, H., I. Zoologisches Institut der Universitat Wien, A-10lO Wien-1, Dr. Karl Lueger
Ring 1
PERRELET, A., Universite de Gen~ve, Dept. Histologie, CH-1211-Gen~ve 4,20, rue de
I' Ecol e de Medeci ne
PICK, H., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr. 38
REICHARDT, W., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemannstr.38
REISSLAND, A., II. Zoologisches Institut der Freien Universitat, D-1 Berlin-41, Grunewaldstr.34
RIBI, W., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, Kunstlergasse 16
ROTH, H., Zoologisches Institut der Technischen Hochschule, D-61 Darmstadt, Schnittspahnstr.3
ROTHENBACH, D., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstler-
gasse 16
SCHALLER, F., I. Zoologisches Institut der Universitat Wien, A-10lO Wien-1, Dr. Karl Lueger
Ring 1
SCHIFF-SERTORIO, H., Laboratoire d'Anatomie physique compare, Universite de Gen~ve,
CH-1200 Gen~ve, 4, rue Candolle
SCHNEIDER, V., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
SCHNETTER, B., Zoologisches Institut der Universitat, D-87 WUrzburg, Rontgenring 10
SCHOENENBERGER, N., Laboratoire d'Anatomie physique compare, Universite de Gen~ve,
CH-1200 Gen~ve, 4, rue Candolle
SCHUEMPERLI, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
SEGESSER v. BRUNEGG, S., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich,
KUnstl ergasse 16
SPATZ, Ch., Institut fUr Biologie III (Genetik und Molekularbiologie) Universitat Freiburg,
D-78 Freiburg i.Br., Schanzlestr. 9-11
STRAUSFELD, N.J., Max-Planck-Institut fUr biologische Kybernetik, D-74 TUbingen, Spemann
strasse 38
STRECK, P., Universitat Regensburg, LS Biologie II, D-8400 Regensburg, Universitatsstr. 31
TOGGWEILER, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
XI
VIRSIK-STRAUSFELO, R., Max-Planck-Institut fUr biologische Kybernetik, 0-74 TUbingen,
Spemannstr. 38
VOGT, K., Max-Planck-Institut fUr biologische Kybernetik, 0-74 TUbingen, Spemannstr. 38
WEHNER, R., Zo:>logisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
WEHRHAHN, Ch., Max-Planck-Institut fUr biologische Kybernetik, 0-74 TUbingen, Spemannstr.38
WEILER, R., Zoologisches Institut der Universitat ZUrich, CH-8006 ZUrich, KUnstlergasse 16
WIEBKING, H., Springer-Verlag, 0-69 Heidelberg-1, Neuenheimer Landstrasse 28-30
ZETTLER, F., Zoologisches Institut der Universitat MUnchen, 0-8 MUnchen-2, Luisenstr. 14
ZIMMERMANN, G., Max-Planck-Institut fUr biologische Kybernetik, 0-74 TUbingen, Spe-
mannstr. 38
