ALS Advances in Life Sciences

Sensory SysteDls and CODlDlunication in Arthropods Including the First Comprehensive Collection of Contributions by Soviet Scientists

Edited by F.G. Gribakin K. Wiese A.V.Popov

1990 Springer Basel AG

Editors' addresses:

Dr. RG. Gribakin Dr. A. V. Popov Sechenov Institute ofEvolutionary Physiology and Biochemistry Academy of Sciences of the USSR Prospect M. Thoreza 44 194223 Leningrad / USSR

Dr.K. Wiese Zoologisches Institut Universität Hamburg Martin-Luther-King Platz 3 2000 Hamburg 13/ FRG

ISBN 978-3-0348-6412-1 ISBN 978-3-0348-6410-7 (eBook)

Softcover reprint ofthe hardcover Ist edition 1990 Deutsche Bibliothek Cataloguing-in-Publicatlon Data Sensory systems and communicationin arthropods: including the first comprehensive collection of contributions by Soviet ,ecientists / ed. by RG. Gribakin ... - Basel; Boston; Berlin: Birkhäuser, 1990 (Advances in life sciences)

NE: Gribakin, Feliks G. [Hrsg.]

The publisher cannot assume any legal responsibility for given data, especially as far as directions for the use and the handling of chemicals and drugs are concerned. This information can be obtained from the manufacturers.

This werk 18 subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law, where copies are made for other than private use a fee 18 PBiYable to «Verwertungsgesellschaft Wort», Munich.

© 1990 Springer Basel AG

Origina\ly published by Birkhäuser Verlag in 199().

The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.

DOI 10.1007/978-3-0348-6410-7

The Academy of Sciences USSR and

Deutsche Forschungsgemeinschaft

have sponsored two bilateral Symposia

on

Sensory Systems and Communication in Arthropods

This volume reports on the Leningrad symposium october 8.-15. 1989

The editors of this book are indebted to:

Sergej Kapitzkii, Ieningrad,

Barbara Schmitz, Konstanz,

Theo Weber, Seewiesen,

und Rudolf Steinbrecht, Seewiesen.

jor their jriendly and efflCient help in preparing camera-ready print-outs oj the contributions.

At Hamburg,

Torsten Rüting, Petra Skiebe and Bemd I.ii.hr

helped with the final synthesis.

v

VI

PREFACE

Among arthropods one can find enormous numbers of species weH adapted to vastly different environments. A good part of this achievement is based on various elaborate behavioural patterns (some stereotyped. some plastic) which are -as has been learned very recently -realized by relatively simple neuronal circuits. That is why conspicuous arthropod behaviours are widely used as model objects in neurobiology when attempts are made to und erstand behaviour in terms of activity of single nerve cells. or to study information processing in sensory systems.

In the West neurobiologists working in these fields are actively cooperative. Russian scientists were for a long time rather isolated because of political restrictions and due to a language barrier; western scientists are as a rule. not familiar with the development of this field in Russia. With new trends in the political athmosphere throughout the world of last years. it became possible to break down the wall and to start active scientific cooperation between West and East. That is why the suggestion of Prof. Markl. the president of the Deutsche Forschungsgemeinschaft. to organize 2 soviet-german symposia on:

"Sensory Systems and Communication in Arthropods" (the first in the USSR, the second in Germany) was actively supported by the USSR Academy of Sciences and several universities.

The main idea was to create the conditions for free and intensive exchange of experience between scientists especially young ones working on arthropods. to establish personal contacts and to define possible ways of further cooperation.

Since sensory systems can not be understood without knowledge about their role in behavior and not without knowledge of the conditions of their functioning in natural environments. specialists in different fields. (morphology. physiology. zoology. ethology. ecology) were put together with the intention to develop forms of more integrated approach to sensory systems when they are not treated as a black box but as parts with identifiable contributions to the whole system of communication.

The first symposium with about 75 participants (25 from the west) was organized at Leningrad (october 8-15th) in 1989. This book summarizes the information presented during this event and the readers are to decide how successful we were in our efforts to desribe typical systems of communication in arthropods and what we have learned about the design and performance of some of the sensory systems involved.

One result is evident. We better und erstand each other. we trust each other and both sides are open to wide and close co operative contacts in the future.

Leningrad and Hamburg. spring 1990 The editors

TABLE OF CONTENTS

SYSTEMS OF OLFACTORY COMMUNICATION

MORPHOWGICAL AND ELEC1ROPHYSIOWGICAL CHARACTERISTICS OF INSECT SENSILIA

VII

Tatyana A. Belousova & Alexey Ye. Redkozubov 3

NEURAL BASIS OF CHEMICAL COMMUNICATION BE'IWEEN SEXES IN THE AMERICAN COCKROACH. PERIPLANETA AMERICANA Jürgen Boeckh 10

CONFORMATIONS OF AMERICAN COCKROACH SEX PHEROMONE AND ITS MIMIC Maria B. Bykhovskaya & Boris S. Zhorov 16

PREY SPECIFICI1Y AND PREY RECOGNITION IN THE DIGGER WASP URIS NIGER FABR.: THE IMPORTANCE OF CWSE RANGE CHEMICAL CUES Wemer Gnatzy. Sylvia Anton and Susanne Thier 20

INTRA- AND INTERSPECIFIC CHEMICAL SIGNALS IN THE TERMITE SCHEDORHINOTERMES -Production sites. chemistry and behaviour Manfred Kaib 26

EFFECT OF WWERED p02 ON ELECTRICAL PROPERTIES OF THE ANTENNA IN MALES OF PERIPLANETA AMERICANA Sergej V.Kapitzkii and Felix G. Gribakin 33

TERMITE TRAIL PHEROMONES: SPECIFICIlY AND BIOSYNTHESIS Sergej G. Klochkov & Dmitrij P. Zhuzhikov 40

FINE STRUCTURAL FEATURES OF SENSORY SYSTEMS IN TICKS AND MITES: EVOLUTIONARY AND ETHOLOGICAL ASPECTS Sergej A. Leonovich 44

RECEPTOR POTENTIALS RECORDED FROM INTACT OLFACTORY SENSILLA OF THE MALE SILKMOTH ANTHERAEA PERNYI Alexander V. Minor & Tatyana A. Belousova

SOME CHARACTERISTICS OF INSECT PHEROMONAL COMMUNICATION Algirdas Skirkevicius

49

55

VIII

FINE STRUCTURE OF ANTENNAL CONTACT CHEMORECEPTORS AFfER CRYOFIXATION Rudolf Alexander Steinbrecht and Jong-Kyoo Lee 59

FINE STRUCTURE OF INSECT THERMO-/HYGRO-SENSITIVE SENSILLA AFfER CRYOFIXATION: STRUCTURAL CHANGES AFfER MOIST- AND DRY- ADAPTATION Rudolf Alexander Steinbrecht 62

MORPHOPHYSIOLOGICAL PROPERTIES OF AN INSECT PHEROMONE SENSORY SYSTEM IN LEPIDOPTERAN AND HYMENOPTERAN SPECIMENS Grazina Vaitkeviciene, Aukse Karaliene & Zuzana Skirkeviciene 69

MORPHOPHYSIOLOGICAL PECULIARITIES OF AN INSECT PHEROMONE SENSORY SYSTEM IN LEPIDOPTERAN AND HYMENOPTERAN SPECIMENS Grazina Vaitkeviciene, Aukse Karaliene & Zuzana Skirkeviciene 75

SYSTEMS OF VISUAL COMMUNICATION

''WHITE EYE" AS A MODEL FOR TIfE STUDY OF OPTICAL PROPERTIES OF VISUAL PIGMENTS IN INSECTS Felix G.Gribakin

NATURAL AND GENETICALLY ENGINEERED VISUAL PIGMENTS OFTIfEFLY

81

Kuno Kirschfeld 86

LUMINOUS COMMUNICATIVE BEHAVIOUR OF THE LUCIOLA MINGRELICA FIREFLIES (COLEOPTERA, LAMPYRIDAE) Sergej B. Landa, Vadim Yu. Bolshakov, Evgenij A. Drobchenko, Semyon I. Peimer 9 1

CALCIUM-CONTAINING OMMOCHROME GRANULES IN OUTER PIGMENT CELLS OF THE HONEYBEE EYE Andrej D. Polyanovsky & Kirill Yu. Ukhanov 96

MECHANISMS OF PHOTOTRANSDUCTION IN INVERTEBRATES Sergej A. Shukolyukov 99

SPECTRAL SENSITIVITY OF PHOTORECEPTORS AND SPECTRAL INPUTS TO THE NEURONS OF THE FIRST OPTIC GANGLION IN THE LOCUST (LOCUSTA MIGRATORIA) Tamara M. Vishnevskaya, Tatyana M. Shura-Bura 106

SYSTEMS OF SENSORY CONTROL OF INSECT FLIGHT

PROPRIOCEPTIVE CONTROL OF FUGHT IN INSECTS

IX

Valerij S. Gorelkin, Yurij A. Karelin & Vladlmir L. Svldersky 115

PERIPHERAL INHIBmON IN THE SYSTEM RESPONSIBLE FOR INITIATION AND MAINTENANCE OF LOCUST FUGHT Vladimir L. Svldersky 122

VISUAL CONTROL OF COMPENSATORY HEAD MOVEMENTS IN THE SPHINX MOTH Uwe J. Dombrowski, Jochen J. Milde and Gernot Wendler 127

BEHAVIOURS DESCRIBING SYSTEMS OF COMMUNICATION

SYSTEM ANALYSIS OF INSTINCTIVE BEHAVIOUR IN INSECTS Peter M. Filimonov, Eugenij A. Faidysh 137

BEHAVIOURAL CHOICE OF CONSPECIFIC MATES IN CLOSELY RELATED DRAGONFLY SPECIES Leonld I. Frantsevltch, Pyotr A. Mokrushov 142

BEHAVIOURALFEATURESANDCOMMUMCATIONINSOCIAL WASPS VESPULA (VESPIDAE, HYMENOPTERA) Vladlmir P. Ivanov 149

VISUAL SEARCHING OF LOCAL FOOD SOURCES IN SOCIAL HYMENOPTERA Vladimir M. Kartsev 154

AUDITORY AND MECHANOSENSORY SYSTEMS OF COMMUNlCATION

SPIDER COURTSHIP: MALE VIBRATIONS, FEMALE RESPONSIVENESS AND REPRODUCTIVE ISOLATION Frledrich G. Barth 161

ACOUSTIC BEHAVIOR OF MOLE CRICKETS OF GRYLLOTALPA GENUS Vitalij S. Chukanov, Dmitrij N. Lapshin 167

AUDITORY INTERNEURONS IN THE FREELY MOVING BUSHCRICKET TEITIGOMA CANTANS FUESS.(ORTHOPTERA, TETTI GO NIl DAE) Vitalij S. Chukanovand Rustem D. Zhantiev 173

x

RESPONSES OF MOTHS TO ULTRASOUNDS Marina V. Fyodorova. Dmitrij N. Lapshin 178

STRIDULATORY INTERNEURONS IN THE METATHORACIC GANGLION OF THE GRASSHOPPER OMOCESTUS VIRIDULUS L. Berthold Hedwig. Siglinde Gramoll and Norbert Elsner 183

THE INTERSEGMENTAL NETWORK UNDERLYlNG STRIDULATION IN THE GRASSHOPPER OMOCESTUS VIRIDULUS L. Berthold Hedwig

MODULATION OF AUDITORY INFORMATION PROCESSING BY MOTOR ACTIVITY AND MECHANICAL STIMULATION IN GRASSHOPPERS AND LOCUSTS Berthold Hedwig. Friederike Lang and Norbert Elsner

BlOPHYSICAL AND NEUROPHYSIOLOGICAL EFFECTS OF RESPIRATION ON SOUND RECEPTION IN THE MIGRATORY LOCUST LOCUSTA MIGRATORIA Axel Michelsen. Berthold Hedwig and Norbert Elsner

LIFE HISTORY AND COMMUNICATION IN PHANEROPTERID BUSHCRICKETS Klaus-Gerhard Heller

PATTERN RECOGNITION AND DIRECTIONAL ANALYSIS: ROUTES AND STATIONS OF INFORMATION FLOW IN THE CNS OF A GRASSHOPPER Dagmar v. Helversen and Otto v. Helversen

ACOUSTIC COMMUNICATION IN PERIODICAL CICADAS: NEURONAL RESPONSES TO SONGS OF SYMPATRIC SPECIES Franz Huber. Hans-Ulrich Kleindienst. Thomas E. Moore. Klaus

189

193

199

204

209

Schildberger and Theo Weber 217

A SIGNAL PRODUCED BY A DIGGER WASP PREYlNG ON CRICKETS Günter Kämper and Wemer Gnatzy 229

INFORMATION PROCESSING IN THE CERCAL SYSTEM OF CRICKETS: IMPLICATIONS OF GROWTH Günter Kämper 232

COMPARATIVE INVESTIGATION ON THE MORPHOLOGY AND PHYSIOLOGY OF THE AUDITORY RECEPTOR ORGANS OF SEVEN SPECIES OF BUSHCRICKETS Klaus Kalmring. Rüttger Ebendt. Janak Ahi. Judith Hellweg. David Young. Heribert Halex. Reinhard Lakes. Wolf gang Rössler and Jürgen Schröder 241

IMPORTANCE OF TOOnI IMPACT RATE IN ACOUSTIC COMMUNICATION IN BUSHCRICKETS Klaus Kalmring. Wolf gang Roessler. Jürgen Schroeder. Otto Stiedl. UIf Bickmeyer and Winston J. Bailey

SOUND LOCALIZATION IN INTACT AND ONE-EARED CRICKEfS

XI

248

Hans-Ulrich Kleindienst. Klaus Schildberger. Franz Huber 254

MECHANORECEPTORS AND PECULIARITIES OF LOCOMOTOR BEHAVIOUR OF INSECTS Natalya I. Knyazeva 259

INTERACTION OF MECHANORECEPTOR SYSTEMS AS A BASIS FOR ACOUSTIC COMMUNICATION IN INSECTS Alexander N. Knyazev

HORMONAL REGULATION OF ACOUSTIC COMMUNICATIONS IN CRICKEfS

265

Alexander N. Knyazev and Irina V. Chudakova 271

DEVELOPMENT OF SENSORY CELLS OF THE LABELLA AND THE LEGS OF THE BLOWFLY PHORMIA REGINA Reinhard Lakes and Gerald S. Pollack 275

PLASTICITI OF nIE NERVOUS SYSTEM OF ORTHOPTERANS Reinhard Lakes 280

TEMPORAL RELATIONS BElWEEN THE PATTERN OF THE CALLING SONG AND ACTIVITI OF AUDITORY INTERNEURONS IN THE FREELY MOVING CRICKET GRYLLUS BlMACULATUS (ORTHOPTERA. GRYLLIDAE) Dmitrij N. Lapshin. Rustem D. Zhantiev 285

THE ACTION OF SENSORY INPUTS ON MOTOR CENTERS EXCITABILITI IN INSECTS Victor P. Lapitzki. Vladimir V. Kovalec. Anna I. Grigoryeva

TRANSFER OF INFORMATION DURING HONEYBEE DANCES. STUDIED BY MEANS OF A MECHANICAL MODEL Axel Michelsen and Bent Bach Andersen

CO-EVOLUTION OF SOUND-PRODUCTION AND HEARING IN INSECTS Andrej V.Popov

INFORMATION THEORY APPROACH TO COMMUNICATION IN ANTS Zhanna I. Reznikova. Boris Ya. Ryabko

289

294

301

305

XII

CUES FOR PHONOTAXIS IN BUSHCRICKETS Jürgen Rheinlaender

INSECT ACOUSTIC COMMUNICATION OUTDOORS: A MULTIDISCIPLINE APPROACH Heiner Römer

CONTRIBUTIONS OF BRAIN AND THORACIC GANGLIA TO THE GENERATION OF THE STRIDULATION PATTERN IN CHORTHIPPUS DORSATUS Bernhard Ronacher

THE CERCAL SYSTEM: CONSTANCY OF SPATIAL LOCALIZATION AND ITS MECHANISMS

308

312

317

Galina I. Rozhkova 324

ANTENNAL SWEEPS ELICITED BY WATER VIBRATIONS IN THE CRAYFISH (PROCAMBARUS CLARKIl). DIRECTIONALITI BEFORE AND AFTER OBSTRUCTING INPUT OF TAILFAN MECHANORECEPTORS. Barbara Schmitz 329

ONTOGENESIS OF PHONOTAXIS IN FEMALE CRICKETS GRYLLUS BlMACULATUS Maria V. Sergeeva & Andrej V. Popov 335

PLASTICITI OF PHONOTAXIS SPECIFICITI IN CRICKETS Victor F. Shuvalov 341

IS HISTAMINE THE INHIBITORY TRANSMITTER OF THE OMEGA NEURONES (ON1)? Petra Skiebe. Brian J. Corrette and Konrad Wiese 345

TRANSMISSION AND FILTERING OF BEHAVIOURALLY RELEVANT SONG PARAMETERS BY AUDITORY NEURONS IN A GRASSHOPPER Andreas Stumpner. Bemhard Ronacher. Otto v. Helversen

CAN THE DIRECTIONAL SENSITIVI1Y OF SINGLE MECHANOSENSORY NEURONS IN ARTHROPODS TELL THE ANIMAL ANYrHING ABOUT STIMULUS DIRECTION? Jürgen Tautz

RESPONSES OF SYMPATRIC ACRIDID SPECIES TO NATURAL AND ARTIFICIAL SOUND SIGNALS

351

359

Varvara Yu. Vedenina 366

CHARACTERISTICS OF SENSORY INPUTS IN CRICKET CERCAL SYSTEMS Varvara Yu. Vedenina and Alexej L. Byzov 371

PHONOTAXIS AND VISUAL ORIENTATION IN GRYLLUS CAMPESTRIS L.: BEHAVIOURAL EXPERIMENTS Theo Weber

PATrERN RECOGNITION AND LOCALIZATION IN CRICKET PHONOTAXIS. Gemot Wendler

A TIME CONSTANT IN THE AUDITORY PATHWAY OF THE CRICKET GRYLLUS BlMACULATUS RELATED TO THE NATURALLY USED PERIOD OF CHIRP REPETITION Konrad Wiese and Simone Albrecht

SOUND COMMUNICATION OF PHANEROPTERIDAE (ORTHOPTERA) Rustem D. Zhantlev & Olga S. Korsunovskaya

AUDITORY INTERNEURONS IN MOLE CRICKETS (ORTHOPTERA: GRYLLOTHALPIDAE) Rustem D. Zhantlev & Olga S. Korsunovskaya

INDEX OF SPECIES

INDEX OF SUBJECTS

INDEX OF AUTHORS: following this page

XIII

377

387

395

402

407

417

419

XIV

INDEX OF AUTHORS

Ahi. Janak 241 Albrecht, Simone 395

Andersen. Bent B. 294

Anton. Sylvia 20

Bailey. Winston J. 248

Barth. Friedrich G. 161 Belousova. Tatyana A. 3. 49 Bickmeyer. Ulf 248 Boeckh. Jürgen 10

Bolshakov. Vadim Yu. 91 Bykhovskaya. Maria B. 16

Byzov. Alexej. L. 371 Chukanov. Vitalij S. 167. 173 Chudakova. Irina V. 271

Corrette. Brian J. 345

Dombrowski. Uwe J. 127

Drobchenko. Evgenij A. 91

Ebendt. Rüttger 241 Elsner. Norbert 183.193.199 Faidysh. Evgenij A. 137 Filimonov. Peter M. 137 Frantsevich. Leonid I. 142 Fyodorova. Marina V. 178 Gnatzy. Werner 20. 229

Gorelkin. Valerij S. 115

Gramoll. Siglinde 183

Gribakin. Felix G. 33. 81

Grigoryeva. Anna I. 289

HaIex. Heribert 241

Hedwig. Berth. 183.189.193.199

Heller. Klaus-Gerhard 204 Hellweg. Judith 241

Helversen v .• Dagmar 209

Helversen v .. Otto 209. 351 Huber. Franz 217.254 Ivanov. Vladimir P. 149

Kämper. Günter 229. 232

Kaib. Manfred 26 Kalmring. Klaus 241. 248

Kapitskii. Sergej V. 33

Karaliene. Aukse 69. 75 Karelin. Yurij A. 115 Kartsev. Vladimir M. 154 Kirschfeld. Kuno 86 Kleindienst. Hans-Ulrich 217. 254 Klochkov. Sergej G. 40

Knyazeva. Natalya I. 259

Knyazev. Alexander N. 265. 271 Korsunovskaya. Olga S. 402. 407 Kovalec. Vladimir V. 289

Lakes. Reinhard 241. 275. 280 Landa. Sergej B. 91 Lang. Friederike 193 Lapshin. Dmitrij N. 167. 178. 285 Lapitzki. Victor P. 289 Lee. Jong-Kyoo 59 Leonovich 44

Michelsen. Axel 199. 294

Milde. Jochen J. 127

Minor. Alexander V. 49

Mokrushov. Pyotr A. 142

Moore. Thomas E. 217

Peimer. Semyon I. 91

Pollack. Gerald S. 275 Polyanovsky. AndreJ D. 96

Popov. Andrej V. 301. 335

Redkozubov, Alexej Ye. 3 Reznikova, Zhanna I. 305 Rheinlaender, Jürgen 308

Römer, Heiner 241, 312 Rössler, Wolf gang 248 Ronacher, Bernhard 317, 351 Rozhkova, Gal1naI.324

Ryabko, BonsYa. 305 Schildberger, Klaus 217, 254 Schmitz, Barbara 329 Schröder, Jürgen 241, 248

Sergeeva, Mada V. 335 Shukolyukov, Sergej A. 99

Shura-Bura, Tatyana M. 106 Shuvalov, Victor F. 341

Skiebe, Petra 345 Skirkeviciene, Zuzana 69, 75 Skirkevicius, Algirdas 55 Steinbrecht, Rudolf A. 59, 62 Stiedl, Oliver 248 Stumpner, Andreas 351

Svidersky, Vladimir L. 115, 122

Tautz, Jürgen 359 Thier, Susanne 20 Ukhanov, Kinll Yu. 96 Vaitkeviciene, Grazina 69, 75 Vedenina, Vanrcrra. YU.366, 371 Vishnevskaya, Tamara M. 106 Weber, Theo 217, 377

Wendler, Gemot 127, 387

Wiese, Konrad 345, 395

Young, David 241

Zhantiev, Rustem D. 173,285,402,407 Zhorov, Bons S. 16

Zhuzhikov, Dmitrij 40

xv

XVI

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h Jl J 1.1, I I I 5.-• •

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IH n J1-n 1. )'1 ~)nJ~@ ~rl'a'J'1' ,,@ » F .. ., ..... • ;

Conjerence anthem.: The text is said to be typically russian. it tells about a case of attempted communication between a human and a group of arthropods. which. at that time. failed (crude methods. lack of patience on behalf of the human). Nevertheless. the overall mood of this popular song is positive. encourages continued dialogue between man and arthropod.

y ~eAYmxH aa neqKOm KOMnaHHH CH~~ H,pacneBSH neceHKB, ycauH meB8nH~, noymHHSDT ~PYEHO H nO&8~cR H8 OOqOK qeTHpe HepaanyqHHX TspaKSHS • CBepqOK. KSK-TO aB Bcm 0PSBY H~ C~SPHK ~oOHn, BCHna~ aa neq~ OTP8BY, qTOO OxnS~BT~ HX nY3. Hoq~1l UH oßsn cnoKoäHO, YTpOU llvnea Ha n8q~, A TAMS BecenaR KOMß8HBH aa ßeqKoro CB~HT H.pacneBa nec8HKB, YC8MB meB8nHT. COEPSnH c SßneTH~OM R~OBB~HH nopomOK qeTHpe HepssnyqHHX TspaxsBs,. CB8pqOX. nnmsyn Hsm ~e~ cep~~o H ßepe~ T8U, K8K n8q~, B3Rn OH B ~HSUHTOU PasBopoTHn Bom ßeq~. Y!POII CTapHK aa~an Mycop y~pST~ B yrny ••• A TAM: BecenSR Koun8HHR BS K811eWK8X CH~~ ~,pAcneE8H neceHKH, YCSMB meBenHT, Y~SCHHä B3phlB ßepeHeCnH, K8K naCK~BH~ menqOK. qeTbIpe HepaanyqHHx fSPSKSB8 1'; CBepqOK. 3noll cy~~Oon yOHfhtä npOKnRn Bec~ ~enYä CBef, be~an OH ~O AHfSPKTB~ Of KBapfßpSHfOB ~e~. npß6hIn Ha ~~HhtH nonDO, C'Tl\pHn CBOM CYH~yqOK ••• A TAM: Becen8H KCMnaHHH no~ xpHmKOE CH~f, Mi,pacneBaa ßeceHKH, ~caLW me:aenHT, nO~J.ißr~fBalOT Beceno: '3~OPOBC, CTapMiqOK 1" qeThIpe HepaanyqHbIX TapsKaHs Mi CBepqOK.

XVII

XVIII

Hotel Morskaya Woksal, Leningrad, Site of the conference SSCA