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INTERNATIONALSOCIETY FOR

NEUROETHOLOGY Newsletter

November, 2001 International Society for Neuroethology c/o Panacea Associates 744 Duparc Circle Tallahassee FL 32312 USA

phone/fax: +001 (850) 894_3480E-mail: ISN@panassoc.comWebsite: www.neurobio.arizona.edu/isn/

©2001 Int’l Soc. Neuroethology. Authors may freely use the material they have provided.

ISN OFFICERSPRESIDENT: Albert S. Feng, Dept. Molecular &Integrative Physiology, Univ. of Illinois, Urbana IL 61801USA Phone:1_217_244_1951, FAX: 1_217_244_5180,a_feng@uiuc.edu.

TREASURER: Sheryl Coombs, Parmly Hearing Inst.,Loyola Univ. of Chicago, 6525 N. Sheridan Rd., Chicago,IL 60626 USA. Phone: 1-713 508-2720; FAX: 1-713 508-2719; scoombs@wpo.it.luc.edu

SECRETARY: Arthur N. Popper, Dept. of Biology, Univ.of Maryland, College Park, MD 20742 USA.Phone:1_301_405_1940, FAX: 1_301_314_9358,ap17@umail.umd.edu

PAST PRESIDENT: Malcolm Burrows, Dept. ofZoology, Cambridge Univ., Downing St., Cambridge, UKCB2 3EJ. Phone: 44 1223 336628, FAX: 44 1223 330934;mb135@cus.cam.ac.uk

PRESIDENT ELECT: PLEASE VOTE!

FROM PRESIDENT ALBERT FENG

Dear Colleagues I would like to take this opportunity to thank theoutgoing officers (Malcolm Burrows, Arthur Popper andSheryl Coombs) who have done a marvelous job over thepast three years. During their term, the professionalismthat John Hildebrand had established for the ISN wasstrengthened significantly. We are all indebted to themand their hard work. I hope to continue to lead the societyin the direction they have set, as well as new directionsthat you would wish to pursue. I very much look forward

to hearing from you on things we should be doing topromote our discipline. The Sixth congress in Bonn was a great success! Thecongress attendance and the scientific presentations weresuper, and the congress location was ideal. I hope youenjoyed the meeting as much as I did. The Congressorganizers led by Horst Bleckmann and the studenthelpers did a splendid job organizing the meeting. TheCongress Program Committee, under the stewardship ofHermann Wagner, put together a very exciting scientificprogram. We very much appreciate their fine work! Ofcourse, your s

strong participation in the congress was the ultimaterecipe for the success. The organizers and chairs of thesymposia and plenary lectures of the Bonn congress havekindly prepared summaries of highlights of these sessions- these reports are presented in this Newsletter for yourperusal. Many of you have responded to the congressquestionnaire that I sent out to you earlier, and some havemade excellent suggestions for further improvements forfuture congresses. To them I wish to express my sincerethanks. I have recently appointed an ad hoc Long RangePlanning Committee (Arthur Popper as chair, AnsgarBüshges, Ichiro Fujita, Ron Harris-Warrick, CathyRankin), with specific charge and duties. One of theduties is to evaluate the members’ responses to congressquestionnaire, and to prepare a report based on thefindings. This report will be presented in an upcomingnewsletter. The post-congress election is now underway! The mailballots have been sent to you by surface mail. We have a

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slate of excellent nominees/candidates. Please take amoment to vote! Finally, there are seven councilors (Kioshi Aoki, AvisCohen, Sten Grillner, Ed Kravitz, Peter Narins, GerhardNeuweiler, and Catherine Rankin) whose term of officehas expired. On behalf of the society, I wish to thankthem for their important contributions.

With best wishes, Albert Feng

ELECTION OF OFFICERS

ou will shortly receive, or may already have gotten, aballot for the new officers for ISN along with abiographical sketch of each candidate. The list of

officer candidates is below. Please be sure and make yourselections and return your ballot right away. The futureof the Society is in your hands and so it is important thatall members contribute to the voting.

President_elect: Ed Kravitz USA, Axel Michelsen DenmarkSecretary: Brian Mulloney USA, Janis Weeks USATreasurer: Sarah Bottjer USA, Sheryl Coombs USA

Council: Shelley A. Adamo Canada, Horst Bleckmann,Germany, Alison Doupe USA, Martin Giurfa France,Ryohei Kanzaki Japan, Harvey Karten USA, EricKnudsen USA, Frederic Libersat Israel, DorothyPaul Canada, Claire Rind UK Sakai Japan,Mandyam Srinivasan Australia, Annemarie SurlykkeDenmark, Hermann Wagner Germany, HaroldZakon USA

PESENTATIONS AT 6th CONGRESS

n order to help those who attended the 6th Congressremember the many symposia, and in order to providemembers who could not attend with some overview of

the many exciting symposia and talks at the meeting, wehave asked each of the organizers to provide a shortdescription of the highlights of the organized program.Needless to say, there were also numerous wonderfulposters at the meeting, but there is no way that we caninclude a description of each in the newsletter, as much aswe would have liked to have done this. We hope,however, that this “taste” of the meeting will be useful,and wet appetites for attendance at the meeting that willtake place in Denmark in 2004.

Plenary lectures 8-9(Session chair: C. Rind, Newcastle, UK)

Alexander Borst (Berkeley, USA) delivered animpressive lecture on Neural Computation of VisualMotion Information in the Fly. He provided a clear andinsightful account of the way the fly is able to extract anduse motion over its eyes for course guidance. Thedirection of motion is not explicitly represented at thelevel of the retina but rather motion information has to becalculated by the nervous system from the changingimages on the retina. Axel Borst examined this process inthe fly, a fast-flying vision specialist in which it is

possible to study the performance of single cells andsmall networks of cells as the fly makes visualdiscriminations. The neurons Borst and his lab haveexamined are located only a few synapses away from thephotoreceptors of the eye and yet they respond in aselective way to motion of the fly in a particular direction.Borst described his experiments using calcium sensitivedyes to image segments of the flies motion detectingneurons to test if processing in a small segment wasdependent on the structure of the image, as predicted bythe model of directional selectivity proposed by Reichardtand Hassenstein. The dependence of the signal on thestructure of the image was found at the level of a smallsegment of dendrite however it did not appear in theoutput of the neuron as a whole. This was because thedendrites of the cell were oriented along the direction ofmotion ensuring integration removes any ripple visible atthe local level. Borst and co-workers have also usedinformation theory and discovered that the informationrate carried by a motion detecting neuron is limited eitherby the noise of the visual pattern (photon noise) or theneural hardware, but not the stimulus entropy rate (e.g.stimulus frequency). The neurons were found to adapt toparticular stimulus conditions so that they were ablecover the dynamic range of a given stimulus. Thisadaptive rescaling of their response acts to keepinformation transmission constant in the face ofdynamically changing visual inputs.Small networks of neurons extract motion cues andinteractions between neurons enhance the tuning of thecells for particular movements of the fly- for examplerotation versus translational movements. Axel predictedthat the cellular organization of the flies motion circuitrywould be understood in his lifetime. Luckily for us he isstill in his prime! In his lecture on Fly Memories-What, Where and How,Martin Heisenberg (Würzburg, Germany) gave anoverview of research on learning and memory in flies. Hebegan by revealing that a male Drosophila can rememberbeing rejected in love for up to 10 days after the event.Flies can remember unprecedented consequences of their

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actions (operant conditioning) as well as novel groupingsof cues in the world (classical conditioning). Both typesof memory have been shown to improve their ability tomake decisions. The cells responsible, at least forolfactory memory are the 700 neurons in the mushroombodies within the fly brain. Flies do not remember ifactivity in the 700 neurons are blocked, and it is only inthese cells that cAMP dependent synaptic plasticity isnecessary for the laying down of short-term olfactorymemory. An important point made by Martin Heisenbergis that memory is not dependent on circuitry but can bemanifest in single cells. Memories can be stored in aninstant and can last for many days. In visual memory,features that are close together are stored as a singlefigure. Drosophila visual memory can hold at least fourfigures in parallel. Surprisingly visual memories caninitially be erased by changes in the context the fly findsitself in. Martin Heisenberg detailed how visual memory,in contrast to olfactory memory is not stored in the cellsof the mushroom body but the mushroom bodiescontribute to the robustness of visual memories so that thememory is no longer dependent on the context in whichthey were acquired. Memory is not simple even in a flythe size of Drosophila.

Plenary lectures 13-14(Session chair: C. Rankin, Vancouver, Canada)

The first plenary talk on Friday morning was by Dr.John Hildebrand (Tuscon, USA) who spoke on NeuralProcessing and plasticity underlying odor-modulatedbehavior in moths. The data from this talk ranged fromthe level of behavior to neuroanatomy toneurophysiology. John gave us a marvelous story abouthow the moths' brain processes pheromones. The storywas one of a dynamic neural complex with the temporalfiring pattern of afferent cells providing information forhigher centers to process changing olfactory inputs. The second plenary talk was by Dr. Barbara Webb(Scotland, UK) who spoke about using robots to modelanimals. Barbara gave an informative history of robotsbuilt to resemble animals. She talked about ways thatrobots could help us to understand biology and offered usvideos of several fascinating robots that are modeled onbiology, including her own robot cricket. This was anexcellent talk to introduce people to the area of robotmodeling.

Symposium #1 – Venom Cocktails and theOrchestration of Prey Paralysis

Organizers: M. Adams (U.S.A.) & F. Libersat (Israel) Venomous animals employ diverse biochemicalstrategies to subdue and exploit their victims. Predatorsgenerally aim to immobilize prey rapidly for direct

consumption, whereas parasitoids exploit PREY forextended nourishment of developing progeny. Thissymposium examined venom cocktails from predatorscorpions, spiders, and snakes, as well as parasitoid insectwasps. Scorpions have evolved several classes of toxinstargeting different binding sites on sodium channels andconsequently different prey specificity. How does thisdiversity of toxins evolve? Michael Gurevitz (Israel)described C-terminal tail wiggling, whereby the C-terminus varies considerably in its attachment to the toxinsurface via the 4th disulfide bond. The variable locationof this attachment has led to a striking diversity ofbinding faces optimized for different ion channel receptorarchitectures. Michael Adams (USA) described simplevs. complex spider venom cocktails and synergismbetween components. The orb weaver spiders inject acocktail, which consists of polyamine-like antagonistswith high concentrations of the neurotransmitter L-glutamate. The presence of glutamate maximizesavailability of toxin binding sites, enhancing the efficacyof the polyamine toxins. Funnel web spiders also employuse-dependent polyamine toxins, but with sodium channelactivator toxins, which lead to excessive release ofendogenous glutamate. A third class of toxins inducesslower, but irreversible antagonism of presynapticcalcium channels, thus blocking transmitter release. AlanHarvey (UK) described joint actions of krait, cobra andmamba venom toxins. Asian kraits combine alpha-bungarotoxins, blockers of nicotinic receptors, with beta-bungarotoxins, which act presynaptically to decreasetransmitter release. The two classes of toxin produce amore rapid onset of flaccid paralysis than either onealone. African mambas instead produce excitatoryparalysis through the actions of dendrotoxins targetingpresynaptic potassium channels and fasciculins, blockersof acetylcholinesterase. Fred Libersat (Israel) wove astory of voodoo rituals and zombies. The mainprotagonists are a parasitoid wasp and a cockroach host.The wasp injects a cocktail of dopamine and proteintoxins directly into the central nervous system, the desiredeffect being behavior modification instead of paralysis.Two stings are observed, one into the thorax to produce atemporary paralysis, the second into the subesophagealganglion and the brain to induce frantic groomingfollowed by lethargy of the host for 5 weeks. Thecompliant host cooperates with the wasp, allowing itselfto be led into a burrow, whereupon it acquires a singlewasp egg and subsequently serves as the sole food sourceof the developing parasitoid. Libersat showed astupendous movie of the entire sequence - it was a hit.Overall, the symposium covered a number of themes,including toxin evolution, joint actions of venom toxins,and behavior modification by venoms.

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Symposium #2 – Visual Ecology and Physiology ofInvertebrate Color Vision

Organizers: K. Arikawa (Yokohama, Japan) and D.Stavenga (Groningen, Netherlands)

The theme of the symposium was covered by fivepresentations on a few exemplary cases where recentlyconsiderable progress has been achieved on therelationship between ecology and color vision ofinvertebrates. In the first talk by Cronin it was shown that the stronglyvarying chromatic environment of marine animalscorrelates with the spectral characteristics of thereceptors. The visual system of mantis shrimps appears tobe spectrally adapted to the local habitat. Hempel deIbarra demonstrated with intricate behavioral experimentson the honeybee, using colored patterns, that processingof chromatic and achromatic contrast depends on thedistance of the pattern and the spatial distribution of thecontrast in the patterns. This will play a crucial role inflower detection by honeybees. Behavioral experimentson nocturnal hawkmoths presented by Kelberdemonstrated that color vision is present at extremely lowlight intensities, where humans are completely colorblind. Arikawa discussed the amazing complexity of theretina of papilionid butterflies. The modular array of theretinal ommatidia is composed of three classes, withunique combinations of photoreceptors and screeningpigments, yielding highly tuned sensitivity spectra. Somephotoreceptors express more than one type of rhodopsin.The papilionid butterflies have now shown to possessexcellent color vision, including color constancy.Stavenga further emphasized the striking heterogeneity in

the ommatidial characteristics of butterfly eyes, that canbe visualized with a novel optical method. The retinal heterogeneity and its possible function wasalso highlighted in a perfect sequel to the Bonn meeting:a conference on ‘Invertebrate Vision’, held near Lund(Sweden) in the week following Bonn. Several of theparticipants were in both meetings. In his Swedenpresentation, White (Boston) revealed by molecularbiological techniques that a surprising retinalheterogeneity is present in moths. Briscoe (Arizona)showed related findings in diurnal butterflies. Croninadded to his Bonn presentation an extensive overview ofthe stunningly colorful eyes of stomatopods.ISN members enjoying a comfortable evening on the Rhine Although color vision is a classical subject ofneurobiology, the neuronal mechanism underlying colorvision is hardly understood in any animal includingvertebrates. This is largely due to the enormouscomplexity of the brain. As demonstrated in thesymposium in Bonn and the subsequent conference inLund, we have accumulated a substantial amount ofknowledge about the function of the input device, thecompound eye, and the basic features of color visioncapacity by a number of behavioral experiments. Basedon the present state of the field, one of the obvious trendsin the next decade would be to ask how and where in thebrain wavelength information is processed into therecognition of ‘color’. Such attempts in the small brain ofinvertebrates would provide a major breakthrough to ourunderstanding about color vision mechanisms in general,and thus would expand our insight in the basic design ofbrain.

Symposium #3 – Constancy of Perception:Neuroethological and Comparative AspectsOrganizer: V. Bastakov (Moscow, Russia)

For proper spatial orientation, animals utilizeinformation of the physical parameters of externalstimuli. However the brightness of an object, its size,shape and color largely depend on the characteristics ofthe environment. The aim of this symposium is to discusswhat is known about mechanisms of constancy perceptionin various sensory systems and at in animals of variousevolutionary levels of development. Justin Marshall andTom Cronin presented both electrophysiological andmicrospectrophotometric evidence showing that thestomatopod eye has 16 anatomically distinct types ofphotoreceptor. They discussed whether thesephotoreceptors are connected with color constancy or not.Vadim Maximov and Oleg Orlov described color-dependent mate choice in common toads Bufo bufo andexperiments that indicate that relative attractiveness ofcolor stimuli was independent of illumination color in a

wide range of tested conditions and a linear function ofcolor coordinates (R, G and B) of the light reflected from

them. Galina Rozhkova discussed the space constancy

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mechanism in cricket cercal system - this systemprovided adequate processing of inputs from hairreceptors in consideration of positional information thatcould be used to provide an invariant estimation of signaldirection. The theme of constancy of perception wasdeveloped further by Vladimir Bastakov, who discussedsize constancy mechanisms in frogs and toads anddescribed that, without visual structural background,prey-catching or escape responses of frogs and toads tostimuli moving around on fixed distances were triggereddepending on the perceived size of the object andperceived distance. Inappropriate reactions (i.e. escape ofa prey-size object and trying to catch a very large one)were the consequences of the wrong estimation ofmoving object distance, i.e. functionally corresponding tohuman visual illusions. Constancy perception tasks couldbe solved in the nervous systems of simple animals(cricket cercal system, stomatopod eye) and vertebrates(frog, toad, man). The comparative method of constancyperception studies is considered to be perspective due tosimilarity of constancy perception mechanisms inanimals of different classes.

Symposium #4 – Avian Models of ComplexInformation Processing

Organizers: M. Bateson (Newcastle, UK) and S. Healy(Edinburgh, Scotland)

Birds have provided us with some of the best paradigmsfor understanding the connections between brain andbehavior, especially in the realm of complex informationprocessing tasks that involve perception, learning,memory and decision-making. Our speakers were chosento represent four different problems in bird behavior, andto reflect the different levels of analysis at whichbehavioral mechanisms can be explored. Candy Rowe (UK) proposed that the evolution ofcomplex, multimodal animal signals might best beunderstood by studying their psychological impact onsignal receivers. Her experimental evidence highlightedthe importance of studying sensory integration at alllevels in receivers in order to appreciate fully the designof multisensory communication. Anna Gagliardo (Italy) reviewed the evidence for anodor-based navigational map in homing pigeons. Shedescribed the ontogeny of this map and presentedevidence that it can only be acquired if pigeons areexposed to odor information during a sensitive period indevelopment. Lesion studies suggest that the avianhippocampal formation is involved in learning thenavigational map, although young hippocampal birds canstill learn a map if they are subjected to training flightsfollowing surgery.

Melissa Bateson (UK) presented the hypothesis that thecomputation of rate of energy intake while foragingrequires an internal clock capable of timing short timeintervals in the seconds to minutes range. She presentedevidence that foraging starlings have such a timingability. Pharmacological manipulations have revealed thatthe neurotransmitter dopamine is involved in the controlof the speed of the pacemaker underlying this clock. The relationship between variation in the size of a brainregion and its putative cognitive output was discussed bySusan D. Healy (Scotland). She described recentbehavioral experiments that show that the largehippocampus of food storing tits may be specificallyassociated with increased duration of memory rather thanimproved accuracy of memory or higher number oflocations remembered.

Symposium #5 – Multi-modal sensory guidance ofcomplex behaviors

Organizers: S. Coombs (Chicago, USA) & J. New(Chicago, USA)

Animal behaviors rely upon the integration ofinformation from a number of sources of sensoryinformation. The assembly of this information into acoherent and useful “gestalt” upon which appropriatebehavioral responses can be generated is a fundamentalrole of the brain. This symposium brought togetherinvestigators who have used a variety of approaches toaddress questions of how sensory systems extractdifferent kinds of information and how the relativecontributions of different systems might change duringthe time course of a given behavior. At least five, overriding principles have emerged fromthis symposium. One, different submodalities of thesame sensory system can underlie different behaviors.For example, the ability of fish orient to uniform flowfields (rheotaxis) or to sources (e.g. prey) of spatiallynon-uniform fields, is mediated by separate classes oflateral line receptors. Two, different sensory systemssimultaneously extract fundamentally different types ofinformation to accomplish a single behavioral objective.The ability of many invertebrates and fishes to track odorplumes involves a mechanosensory-based rheotaxis to airor water currents. Three, the contributions of differentsensory systems to a behavior can change as a function ofdistance to the stimulus. The prey capture behavior ofsome fishes can be divided into two phases: an initialapproach phase that is guided by visual cues and a strikephase guided by either visual or lateral line cues. Four,the relative contributions of sensory systems may dependon changing environmental factors: electroreceptiondominates the prey capture behavior of weakly electricfish at low water conductivities, but at higher

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conductivities the mechanosensory lateral line may play asignificant role. Five, simultaneous use of differentsensory modalities can enhance information transfer andcontent. In wolf spiders, for example, females are mostreceptive to courtship by males when both visual andvibratory cues are present. It was apparent from these presentations that animalmodels provide a rich source for examining multimodalinformation processing and integration in nervoussystems. This field of investigation offers great promisefor a better understanding of the means by which brainsguide behaviors.

Symposium #6 – Auditory Information Processing andEcholocation: From Neurons to Robot Bats

Organizers: E. Covey (Seattle, USA) & C. Moss(College Park, USA)

Echolocation in bats is one of the foremost areas inwhich it is possible to link neural circuitry to specificbehavioral patterns. Although neuroethological studies ofecholocation have traditionally emphasized sensoryprocessing of echolocation signals, this symposiumpresented information about aspects of the neural basis ofecholocation that have not received extensive publicity.Elisabeth Kalko discussed the relative contributions ofphylogeny and environmental conditions to echolocationcall design in species that hunt in open versus clutteredspace, and those that engage in aerial prey capture versusthose that glean prey from surfaces. Thomas Park andAchim Klug showed that single neurons in the midbrainof the free-tailed bat respond to both echolocation andcommunication calls, and proposed that these responsesare largely based on frequency tuning combined with thetemporal properties of monaural and binaural excitatoryand inhibitory inputs. Nick Fuzessery presented evidencethat the pallid bat uses echolocation for generalorientation, but relies on passive sound localization todetect prey. It must therefore attend simultaneously totwo streams of acoustic information while hunting. Itsauditory system contains a high percentage of neuronsselective for its echolocation pulse, a feature not observedin dedicated echolocators like the free-tailed bat. Theseresults suggest that this response selectivity helpssegregate the parallel, dedicated pathways required forthis dual-stream processing. Walter Metzner discussed theneural circuitry used for auditory control of vocal outputin bats that adjust their vocalizations to compensate forDoppler-shifts in the returning echoes. Ghose and Mosspresented animated data from studies of bats takingtethered insects in a laboratory flight room. Theydescribed the temporal patterning, direction and focus ofthe bat's sonar beam during target selection andinterception, and proposed that echolocation signals can

be used as an index of attention and selective tracking.Timothy Horiuchi described neurobiologically detailedVLSI circuit models of the bat echolocation system testedin a flying bat robot. Behaviors simulated include targetlocalization in three dimensions, tracking of targets usinghead movements, and control of vocalization.

Symposium #7 – Polarization VisionOrganizers: T. Cronin (Baltimore, USA)

& C. Hawryshyn (Victoria, Canada) It has been recognized for more than half a century thatanimals are sensitive to and respond to the polarizationproperties of natural light fields. Recent research on thispolarization sensitivity has progressed well beyond thehistorical perspective, where its utility has primarily beenrecognized as contributing to orientation and navigation.In this symposium, we featured a variety of presentationsillustrating new results in the developing field ofpolarization vision (PV). Marie Dacke presented work done at the Univ. of Lund,Sweden, on PV in the spider Drassodes cupreus. Thisanimal has eyes with polarization sensitive receptors thatintegrate over a large part of the sky. The spiders areprimarily active after sunset and that they use polarizationcues to find their way back to the nest. Justin Marshall(Univ. of Queensland) followed, discussing the use ofspecialized polarization signaling systems in thestomatopod crustaceans. These animals have numerouspolarization receptor classes, and may even be capable ofthe analysis of circular polarization as well as ofvisualizing species-specific signals only used duringintraspecific encounters. The theme of polarizationsignaling was developed further in the next talk, byNadav Shashar (InterUniv. of Eilat), in his powerfulpresentation on cephalopod PV. Squids and cuttlefishesalso identify prey using this visual modality. The symposium then turned to vertebrate PV,concentrating on the salmonid fishes. Daryl Parkyn(Univ. of Florida) presented a suite of results on the useof polarized-light cues for orientation, as well as coveringproperties of visual and central neurons that arespecialized for polarization analysis. Craig Hawryshyncontinued this theme by introducing new results onmultidimensional polarization analysis by the visualsystems of marine damselfishes. Attendees of this symposium were treated to a newwindow into the ways animals view their worlds, as thesymposium provided strong evidence for theincorporation of PV and polarized-light cues into visualdomains traditionally assigned to spatial and color vision.The extension of this research avenue into new phyleticgroups and new environments suggests that exciting newaspects of PV are yet to be discovered.

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Symposium #8 – Dendritic Computation inNeuroethological Relevant Systems

Organizer: R. Eaton (Colorado, USA) Dendritic computation is of critical importance inunderstanding all cellular and network functions from thesimplest reflex to the most advanced cognition. Technicaladvances in imaging, electrophysiological recording, andpowerful numerical simulation methods have led to arapid expansion of knowledge and interest in this area.This symposium covered dendritic processing in fourdifferent systems: A. Borst and J. Haag: In most neurons dendrites processthe incoming signals of presynaptic nerve cells byconverting them into a single, time varying signal that ispassed along the axon onto one or many postsynapticpartners. In addition, the dendrites of some neurons,mostly involved in local processing, have been found topossess output synapses, too. In their talk, Borst and Haagdescribed in a fly visual interneuron a type of dendritewith a dual mode of operation: for neurons postsynapticto the axon terminal the dendrite operates as an integrator,while for neurons postsynaptic to the dendrite it acts as aspatial filter preserving the retinotopic information of themotion image. G. Cummins, J. Casagrand and R. Eaton: Mauthnerneurons of the fish brainstem process acousticinformation and trigger sudden avoidance maneuvers.High levels of input convergence to these cells, and otherneurophysiological data, suggest that the Mauthner lateraldendrite can compute sound source localization. Thisprocess requires a non-linear phase comparison of thesound pressure and acceleration of the sound wave. Thetalk by this group presented data from modeling studiesthat suggest testable cellular mechanisms for thediscrimination. These mechanisms involve voltage-sensitive conductance changes paired with appropriatelocation of afferents on the dendritic membrane. D. Edwards: A lateral excitatory network was found toexist among the terminals of primary mechanosensoryafferents that mediate excitation of the lateral giantcommand neuron for escape in crayfish. When a criticaldensity of afferents is excited, the network recruitsunstimulated afferents to provide additional excitation oflateral giant and of mechanosensory interneurons thatexcite the lateral giant. Depolarization of lateral giants bycurrent injection facilitates the recruitment of afferents,and thereby of the mechanosensory neurons and thelateral giant itself. Edwards suggested that thedepolarizing EPSPs would have the same effect, thuscreating a situation where the lateral giant enhances itsown excitation when its input exceeds a critical value.

G. Jacobs: In the cricket cercal sensory system, primarysensory interneurons are tuned to both the direction andfrequency of air currents. Sensory input creates spatio-temporal patterns of activity within a neural map ofstimulus direction formed by the sensory neurons. Jacobspresented evidence suggesting that interneurons decodethese patterns of activity by virtue of the shape andposition of their dendritic trees within the neural map.The decoding mechanism involves a shape-matchingalgorithm in which the cells respond preferentially tothose patterns of activity that most closely match theshape of their dendritic trees. It was hypothesized thatfrequency tuning could emerge from frequency-dependent phase alignment of synaptic inputs fromdifferent classes of afferents. Jacobs showed that at low stimulus frequencies (30Hz)the spatio-temporal patterns among the sensory neuronswere in-phase, providing the maximum amount ofsynaptic input to some sensory interneurons. In contrast,higher stimulus frequencies (100Hz) produced spatio-temporal patterns among the afferents that weresignificantly out-of-phase with each other, providing alower synaptic drive to the interneurons.

Symposium #9 – Auditory Brain Maps and theirRelation to Sound Perception

Organizer: G. Ehret (Ulm, Germany) What else is mapped besides sound frequency(tonotopic map) in centers of the auditory system ofanimals, from orthopteran insects to vertebrates up tohumans? How do further maps relate to and are adjustedby specific requirements of acoustic communication andthe acoustic environment of a given species? How maybiologically significant information be generated by andextracted from neural activity distributions in maps? Howplastic are the maps? The seven contributors to thesymposium presented exciting evidence about a map ofsound intensity in the cricket CNS (Gerald Pollack), amap of periodicity pitch in the primary auditory cortex ofthe Mongolian gerbil (Holger Schulze), place maps forsound objects calculated from interaural time differences(Albert Feng), adjustment of space maps in the midbrainof owls and rats by attention (Bernhard Gaese),approaches to maps of combination sensitivity of neuronsin the primate higher-order auditory fields to representphonemes (Josef Rauschecker), circuits of corticofugalmodulation of auditory brain maps in bats (Nobuo Suga),and the general significance of auditory maps for soundperception and recognition (Günter Ehret). Fourteen types of maps (orderly or clustered spatialrepresentations) of sound properties or neural responseproperties have been described to exist in centers of themammalian auditory system so far, most in the auditory

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midbrain and/or fields of the auditory cortex. Auditorymaps, similar to maps in the visual system, reflect generalstrategies to process and represent acoustic information inthe brain. It appears that auditory perceptual maps, suchas maps of auditory space, are all based on combinationsensitivity of neurons in centers of the midbrain or ofhigher levels of the auditory pathways (of vertebrates).We have just begun to recognize that auditory maps areplastic and subject to modulation and adjustment duringthe ontogeny of an animal and by mechanisms ofattention and learning. Auditory brain maps in adultanimals will tell us not only about adaptations of speciesto their acoustical niches but also about individualizedbrains reflecting adaptations to individual acousticexperiences and to specific constraints of the acousticenvironment.

Symposium #10 – Cognitive Abilities in InvertebratesRobot Bats

Organizers: M. Giurfa (Toulouse, France)& E. Capaldi (Lewisburg, PA, USA)

The main aim of this symposium was to illustrate themost recent research demonstrating that invertebrates arecapable of rather complex learning tasks that go beyondelementary associations. Research on invertebratelearning and memory has focused mainly on this kind ofassociations, that is, on the building of simple associativelinks between, say, two stimuli or between a stimulus anda response. Recent research demonstrates new ways ofthinking about the study of invertebrate behavioralplasticity. In particular, it has been shown thatinvertebrates are capable of complex spatial learning andof non-elementary, configural learning. In addition, some

invertebrates can categorize stimuli and learn identifycomplex relationships between stimuli. The talks of Carolyn Sherff (UC Irvine) and DanielTomsic (Univ. of Buenos Aires) addressed the problem ofmemory dynamics in Aplysia and in the crabChasmagnathus, respectively. Makoto Mizunami(Hokkaido Univ.) and Stimson Wilcox (BinghamtonUniv.) reported new findings showing non-elementalassociations in crickets and spiders, respectively. Finally,Paul Graham (Univ. of Sussex) focused on learning visualcues for route guidance in ants. One of the main conclusions of the symposium is thatthere are indeed learning performances in invertebratesthat cannot be explained on the sole basis of simpleassociations. The mechanistic basis of such complexlearning forms is still unknown but as invertebrates haveaccessible and relatively simple nervous systems, there isa fair chance in addressing this question at the cellularlevel. The main highlight of the symposium was causedby a technical difficulty encountered in the symposiumroom, which affected the delivery of the last two talks.This problem had, however, an unexpected advantage: thesymposium was the only one in the conference in which a45 min general discussion took place. In such adiscussion some of the points raised above whereintensively discussed by the audience. The use of the term“cognition” was subject of debate. It was concluded that apossible dividing line is not one between cognitive vs. notcognitive abilities, but one between elementary vs. non-elementary forms of learning.

Symposium #11 – Neural Mechanisms of SoundProduction: A Comparative Approach

Organizers: B. Hedwig (Cambridge, UK)& U. Jürgens (Göttingen, Germany)

Sound production and acoustic communication systemshave evolved in vertebrate and invertebrate taxa and playa major role in the behavior and lifestyle of manyanimals. For this reason the symposium presented acomparative approach to the neural mechanismsunderlying sound production. For the first time it broughttogether speakers working on completely different modelsystems, from invertebrates to primates. Besides thegeneral concepts, the species-specific neural mechanismsof vocalization and sound production and the differentmethods to approach them were emphasized. Matthias Henning and Paulo Fonseca (Berlin/Lisboa)and Berthold Hedwig (Cambridge) discussed neuronalpattern-generating mechanisms and their control by singleidentified higher order interneurons in cicadas and in

crickets/grasshoppers, respectively. In these invertebratemodel systems by intracellular recording and stimulationthe functional significance of single identifiedmotoneurons and interneurons for sound production canbe demonstrated. Wolfgang Walkowiak (Cologne) reported on theemerging principles of the central neural organization ofvocalization in amphibians, which is closely coupled withrespiration. Vocalization related moto- and interneuronscan be characterized by intracellular recording even inisolated brainstem preparations. The importance oflearning and of auditory feedback to acquire song patternswas addressed by Daniel Margoliash (Chicago)demonstrating the precise temporal activity patterns ofneurons in awake and sleeping zebra finches. The datasuggest a model whereby efference copy during singingand rehearsal premotor activity during sleep contribute tosong learning.

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The mapping of auditory feedback on the motor patternin bats was addressed by Gerd Schuller (Munich). Heshowed that audio-vocal feedback behavior can bemodulated by prevocal structures, which may reflectspecific adaptations of bats. Finally Uwe Jürgens(Göttingen) discussed the neuronal organization ofvocalization in monkeys. Single unit recordings in awakemonkey provide strong evidence for the existence ofvocalization-initiation and vocalization-pattern generationin different brain areas.

Symposium #12 – Modulatory Signaling: ConveyingInformation is Not Always Exciting (or Inhibiting),

but Can Be Mind-AlteringOrganizer: P. Katz (Atlanta, USA)

This symposium dealt with the implications thatmodulatory signaling may have for how information istransferred between neurons and even how information istransferred between animals. In the nervous system,neuromodulatory signaling is often mediated by Gprotein-coupled receptors and has widespread effects,changing the *cellular or synaptic properties of neurons.The roles of neuromodulation in sensory systems, motorsystems, and complex social behavior were presented inthe symposium. Using biogenic amine neuromodulation in insects as anexample, Alison Mercer discussed the plasticity thatconfers richness and complexity on modulatory messages.She made the point that one of our greatest challenges isnot that there are multiple forms of neural plasticity, butthat the mechanisms underlying neural plasticity arethemselves are remarkably plastic. Vladimir Brezina discussed how modulatory signalssuch as neuropeptides allow a simple neuromuscularsystem in the mollusc, Aplysia, to optimize multipleparameters, giving the animal a greater repertoire ofmovement amplitude and frequency. Paul Katz showed that modulatory signals could playmore than a subsidiary role; the dynamics of secondmessenger signaling may form the basis for motor patterngeneration in the escape swim of the mollusc, Tritonia.Modulatory signaling may also be a target of naturalselection in the generation of species-specific behavior. Thomas Insel discussed how differences in thepromoter region of the vasopressin receptor gene mayunderlie its differential distribution in two closely relatedspecies of vole and how this may contribute to differencesin affiliative behavior in those species; a small change inthe distribution of this G protein-coupled receptor maycause a species to exhibit pair-bonding. Finally, Stanley Schneider introduced the notion of amodulatory behavior: a vibration signal produced byworker bees produces a contingent action on the state of

other bees that depends upon the location of the workersin the hive and the nature of the task they were involvedin. Thus, modulatory signaling may be a ubiquitous meansof communication in systems that require complex action.This symposium was the first of three symposia at themeeting that dealt directly with the neuromodulation,suggesting a rising awareness of its importance in thecontrol of behavior.

Symposium #13 – Modulating the NeuromodulatorsOrganizer: K. Mesce (Minnesota, USA)

In this symposium, participants discussed how potentchemical modulators of behavior are themselves'modulated' by a variety of influences including: socialand developmental events, steroid hormones, and networkconfigurations. By addressing how the physiology andexternal world of an animal shape the suite of neuroactivesubstances within the nervous system, the view waspromoted that a clearer and more comprehensiveunderstanding of behavioral mechanisms can be achieved.Karen A. Mesce (Univ. of Minnesota) in collaborationwith Herman K. Lehman (Hamilton College, NY)discussed how the insect steroid hormone 20-hydroxyecdysone (20-E) was shown to influenceoctopamine (OA), a biogenic amine similar tonorepinephrine. In the insect Manduca sexta, 20-E wasdemonstrated to affect the developmental plasticity of OAsynthesis and the production of new OA-immunoreactive(ir) neurons not previously identified. These newlydescribed OA-ir neurons likely contributed to the steroid-induced elevations of OA observed at the end ofmetamorphosis. The developmental and hormonalregulation of newly identified dopamine-ir neurons wasbriefly discussed and suggested also to be linked tofluctuations in 20-E during metamorphosis. The neuromodulator OA was also reported by David S.Schulz (Univ. of Illinois) in collaboration with Gene E.Robinson (Univ. of Illinois) to influence the transition ofhoney bees from hive workers to foragers. Social factorsthat influenced this shift also influenced levels of OA inantennal lobes of worker bees, suggesting an interactionbetween social environment and the neuromodulation ofbehavior. However, flight experience did not affect OAin antennal lobes; octopamine increased as beesdeveloped towards foraging regardless of whether theywere allowed to fly. These results demonstrate that brainamine systems in the honey bee are sensitive to a range ofboth intrinsic and extrinsic influences. A continued discussion of social interactions and howthey alter the biogenic amines was conducted by DonaldH. Edwards (Georgia State). Don outlined how theabrupt decision of one of two fighting crayfish to escape

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determines the social hierarchy of two crayfish, with theescaping animal identified as the new subordinate and itsopponent as the new dominant. Maturation of thehierarchy was shown to be marked by a reduction in thefrequency of aggressive interactions as the subordinatecomes to avoid the dominant. These behavioral changesappeared to result from neuromodulatory changes thataltered the relative excitability of neural circuits thatmediated discrete behaviors. Two serotonergicneuromodulatory changes were identified in crayfish:changes in patterns of serotonin release that affectedwalking and changes in serotonin receptor distributionthat affected escape. The presentation by Michael P. Nusbaum (Univ. ofPennsylvania) brought the focus of neuromodulation tothe level of identified circuits. It was stressed how notmuch information is available regarding the extent towhich there is uni- or bi-directional communicationbetween descending projections and the neural circuitsthey control. Using the stomatogastric nervous system ofthe crab Cancer borealis, it was documented thatneuronal circuits do indeed provide functionallyimportant synaptic feedback that regulates the activity,and therefore the influence, of modulatory projectionneurons. Michael found that one function of thisfeedback was to enable one neural circuit (the pyloriccircuit) to regulate and coordinate the activity of a distinctbut behaviorally-related neural circuit (the gastric millcircuit). Specifically, this feedback enabled the pyloriccircuit to control the speed of the gastric mill rhythm andits coordination with the pyloric rhythm.

Symposium #14 – Perception of Nearfield Flow and ItsImportance for Animal Behavior

Organizer: J. Mogdans (Bonn, Germany) Near-field medium flow provides important sensoryinformation which allows animals to detect and localizeprey, to avoid predators and to find and communicatewith mates and other conspecifics. Thus, animals frommany different phyla have developed sophisticatedsensory system for flow perception. The principalmechanisms of flow detection are similar in air and water.This became evident in the first talk given by Friedrich G.Barth who showed that some spiders species can haveabout 900 delicate so-called trichobothria on their bodieswhich are the most sensitive movement detectors amonga broad spectrum of cuticular hair sensilla in arthropods.The interaction between the air and the hair follows to alarge degree the principles that are known in fluidmechanics. That air movement detectors are enormouslysensitive was further elucidated by Tateo Shimozawa. Heshowed that the cricket wind receptor achieves asensitivity which is limited only by Brownian motion of

single molecules and is thus comparable to the sensitivityof hair-cells in the inner ear of vertebrates. Amongaquatic animals, one of the most striking sensorycapabilities is found in copepods, small crustaceans ofonly 1-10 mm body length. Jeannette Yen demonstratedin a spectacular video presentation that copepods can usetheir highly sensitive setal arrays to detect, analyze andtrack hydrodynamic signals produced by conspecifics.The fish lateral line is perhaps the best-known exampleamong hydrodynamic receptor systems. JoachimMogdans presented electrophysiological data indicatingthat the lateral line consists of at least two subsystemswhich can only be distinguished when the system isexposed to running water. That hydrodynamic receptionis also found among mammals was demonstratedimpressively by Björn Mauck who showed that seals,using only their vibrissae, can find and track the watermotions generated by a miniature submarine. Thus sealvibrissae can function as hydrodynamic receptors. Thepresentations given at this symposium were highlightingbehavioral capabilities that depend on flow perceptionand addressing neural mechanisms that underlie theprocessing of nearfield flow. The examples presentedclearly demonstrate the importance of flow perception,whether in air or in water, for the daily life of animals.

Symposium #15: Designs for signaling: from sense toaction

Organizer: P. Simmons (Newcastle, UK) The main aim of this symposium was to make sense ofthe various types of signals that neurons use, starting withthe encoding of sensory stimuli and progressing throughthe nervous system to consider motor control. First, JohnBirmingham described how a stretch-sensitive sensorycell in the crustacean stomatogastric system can switchbetween two different modes of code, regular spiking andbursting, which enables it to signal extremely slowchanges in stretch as well as fast changes. Next, BernhardRonacher (with Astrid Franz) revealed new features of aNeuroethology Classic, how grasshoppers identify songsof conspecifics. Surprisingly, at least in one species,successful recognition requires as few as two soundbursts to be heard, and even in the presence ofbackground noise is accomplished in less than half asecond. By the feat of recording from pairs of receptors,he showed that this is probably accomplished by an arrayof parallel receptors producing distinct signals. Deeper inthe nervous system, Peter Simmons described intrinsicvariability in transmission at two types of synapse inflight control pathways in the locust, and proposed thatinsect synapses can be clever at minimising noise, butprofligate with neurotransmitter. Ansgar Büschges (with

11

Turgay Akay and Gabriel Knop) considered the ways thatsensory signals from different types of proprioceptor caninfluence walking in locusts and stick insects. Heproposed that each joint is controlled by its own centralrhythm generating mechanism. Finally, Brian Mulloneytalked about modelling studies that compared the effectsof input from graded potential neurons with the effects ofinput from impulse producing neurons onto swimmeretmotor neurons in crayfish. These demonstrated thatsmooth regulation of spike frequency in the motorneurons can be is achieved by the smoothly graded input

provided by non-spiking interneurons, coupled with ashort membrane time constant. The symposium showedthat, in the post-genomic era, there are plenty of long-standing issues about signalling in the nervous systemstill requiring research. All the talks were aboutarthropods, which are excellent animals for this, becausesignals in identified neurons can be related directly to thecontrol of behaviour.

Symposium #16 – Timing is Everything: The Behaviorand Modulation of Neural Oscillators

Organizer: H. Zakon (Austin, TX, USA) Behaviors must be temporally regulated on many timescales. In this symposium we considered oscillatorsunderlying rhythmic behaviors in vertebrates andinvertebrates. The periodicites of these oscillators varyfrom milliseconds (pacemaker neurons of electric fish), tohundreds of milliseconds/seconds (STG/Leechheart/thalamocortical circuits), to days (circadianpacemakers). Talks focused on how oscillators wereconstructed, how much flexibility or plasticity they show,and how mechanisms of plasticity or intrinsic variation isregulated in systems that must be highly precise in theiroutput.Participants in Neurobiology of Electrosensory Organisms The stomatogastric system of crustacea is a simplenetwork which switches among different behavioraloutputs. Eve Marder showed that neurotransmittersmodulate this circuit in complex ways: similar

transmitters may act on multiple ionic currents or,conversely, different transmitters act on the same current.She emphasized the importance of maintaining outputstability in a complex circuit and showed that there maybe various paths to reach the same point of stablity. MarkA. Masino, showed that in a circuit oscillator that controlsheartbeat in the leech, one of the cells switches betweenbeing rhythmically active and inactive, how thisreconfigures the heartbeat circuit, and how this statechange might occur. Such plasticity at the circuit levelmight be expected in vertebrate circuits. Thierry Baldiscussed the influence of cortical feedback showing howactivation of descending cortico_thalamic axons is gatedby the ascending activity of thalamocortical relayneurons, allowing dynamic, endogenous reorganization ofthe artificially coupled network. The pacemaker neuronsof electric fish, which control the electric organdischarge, are the most precise oscillators known. Yet,Joerg Oestreich showed that their oscillation frequencymay be shifted to a new value following synaptic input

that occurs during a socialinteraction called the jammingavoidance response. On acompletely different time scaleGene Block illustrated howcircadian rhythms ininvertebrates and vertebratesare similar: single cells may actas cellular oscillators withcrude rhythmicitiy with theprecision of the whole circadianoscillator depending on theinteractions of the cells in a"neuronal democracy" asWalter Heiligenberg used tosay.

NEUROBIOLOGY OFELECTROSENSORY

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ORGANISMS 27- 29. August 2001, Bonn/Germany

Gerhard von der Emde and Kirsty Grantgerhard@u.washington.edu, Kirsty.Grant@iaf.cnrs-gif.fr

From an informal beginning in Thomas Szabo's lab inGif sur Yvette in 1989, it has become a tradition to holdan electrosensory satellite forum accompanying thetri_annual International Congress of Neuroethology :Gif_sur_Yvette 1989, Montreal 1992, San Diego 1998.All have been exciting and rewarding events, providing amuch appreciated opportunity to bring together thoseinterested in electrosensory systems. This year's meetingheld in Bonn on July 27_29th, immediately preceding the6th ICN congress, was organized by Gerhard von derEmde, Kirsty Grant, and Hans Meek and broughttogether more than 90 participants from 14 differentcountries and 5 continents. The 17th century Poppelsdorfer Schloss and botanicalgardens made a beautiful setting for 2 1/2 days oflectures, poster sessions, and discussion and thanks aregratefully extended to our hosts Prof. Horst Bleckmannand the University of Bonn Department of Zoology fortheir hospitality, and to Dr. Michael Hofmann for his helpwith onsite organization. Twenty_two lectures and 50 posters covered all aspectsof electrosensory systems from molecular mechanismsserved at the cellular level, peripheral electroreceptors,central electrosensory and electromotor networks, thesensorimotor interface and multisensory integration, totheoretical modeling and fish behavior. Such a range ofapproaches around a common theme gives an exceptionalopportunity to formulate an integrative overview of awhole brain system and to obtain a better understandingof fundamental brain mechanisms, how they mightgovern behavior and in what ways they are useful to theanimal. This field of research has always been pleasantlycooperative and the satellite forum was the occasion formany exchanges between laboratories and planning offuture collaborative projects. A number of major advances showed significantprogress since the meeting in San Diego three years ago.A great effort has been made in the field of evolution:what can the classification of species tell us about how,and how many times, the electric senses and activeelectric organs have appeared? Carl Hopkins showed howthe discovery of new species and modern molecularanalysis is generating a number of surprises in ourperception of phylogenetic trees. Ad Kalmijn's overviewof the physical mechanisms of electroreception andorientation within electric fields provided an excellentbase for a better understanding of how fish (both marineand freshwater species) use their electric sense and

several speakers dealt with the extraction of weak sensorysignals from background noise and the advantagesbrought by multisensory integration in orientation tasks.Our guest speaker, Sheryl Coombs, gave a stimulatinglythoughtful (and graphically beautiful) analysis of thesimilarities and differences between hydrodynamiclateral line and electrosensory systems, dealing inparticular with the problems of depth perception and thevarious neuronal mechanisms available for eliminatingunwanted sensory reafference generated by bodymovement or self_generated noise interference linked todifferent behavioral contexts. The need for the study ofsensory systems in a natural context was also underlinedby Horst Bleckmann, who illustrated the very differentresponses of lateral line mechanoreceptive systemneurons evoked in static and flowing water. Thephysical nature of stimuli seen at the electrosensorysurface, generated by objects in the near environment,was a subject that required many of us to stretch ourimagination in an effort to see as the electrosensorysystem sees, and it became evident that the 'Mexican hat'profile of the stimulus was a property common toexternal, peripheral, and central levels of the imageformation process, and essential to perception of spatialcontrast and depth. This was beautifully illustrated in aposter of "Gnathonemus petersii's electric imagegallery.” Study of the dynamic properties of sensory neuronalnetworks is an on_going area of important progress, nowsuccessfully bringing together experimental approachesand theoretical modeling. It is clear that the theoreticalapproach will be increasingly necessary to understand ingreater depth the algorithms used by the nervous systemfor sensory image representation. Several studies arecurrently exploring the neuronal mechanisms of temporaland spatial contrast, oscillatory behavior and dendriticintegration, and the functional description and role ofsynaptic plasticity. Finally, much progress has been made since the lastmeeting in the area of neuronal networks governingmotor control. Bruce Carlson's elegant dissection ofmidbrain pre_motor network in mormyrid fish, coupledwith in depth study of electromotor signaling behaviorhas brought knowledge of this system close to that of thegymnotid fish. The latter is now being extended by majorcontributions to the study of long_term hormonal orsensory_induced modulation of electromotor behavior,both at the synaptic and cellular level and in theenvironmental context associated with reproductivebehavioral. The proceedings of the satellite meeting will bepublished as a special volume in the Journal ofPhysiology _ Paris (Elsevier) in Spring 2002. We hope

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that this will serve as the current reference until a futuremeeting in Denmark in 2004!

LISTSERV ANNOUNCEMENT

We have found that our membership e-mail list is notcompletely accurate. Keeping up a list is difficult sincepeople sometimes move or change e-mail addresseswithout notifying all their correspondents, and especiallythe listservs they may be using. Thus, we would like toask that all ISN members check their ISN E-mailaddresses directory athttp://www.neurobio.arizona.edu/isn/isn.memdir.htm. Ifyou have any corrections, please send them toJohnHildebrand at jgh@neurobio.arizona.edu.

“FIELD” BEHAVIOREd Kravitz

special columnist to the ISN newsletteredward_kravitz@hms.harvard.edu

© Copyright 2001 Edward A. Kravitz. All rights reserved.

SCIENCE,SOCIAL CONSCIENCE AND SOCIAL POLICY

PART 1-CBW

Prologue: How is one to reconcile the magnitude of theevents of September 11, 2001 in New York, Washingtonand rural Pennsylvania with the day to day routines ofour scientific lives? Shouldn't we be doing somethingmore important? Shouldn't we be doing something toensure that things like this will never happen again?How can we go into our laboratories and watch flies andlobsters fight, or try for the 80th time to record from thatdifficult to find neuron, or go after that elusive gene wehave been trying to clone for the past six months? Theissue for me on the morning of the eleventh was how can Iand my co-organizers go on with the scheduledconference on "Pathologies of Mind" that we hadassembled for Harvard undergraduates. Little did we

realize in ourplanning sessions sixmonths earlier, howrelevant the themewas to be to the eventsof that day. But go onwe did, all the whilefeeling strongly thatwe were doing theright thing--to ensurethat madmen andfanatics cannot dictatehow we spend our

lives. The science we do is important, and can impactdramatically on society. We can cure diseases and

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unravel the "Pathologies of Mind" that allow people tocarry out actions like those of September 11. We trainthe next generations of scientific leaders and influencethe course of the institutions we are part of. The societywe directly impact on may be small but the consequencesof our actions can be large. These were to be the themesand ideas of a several part essay I had planned to writeconcerned with the social consequences of our activitiesas scientists before the events of the 11th. I feel an evengreater urge to write on these themes now. I am often asked "why do you study aggression"? Myanswer invariably has three parts to it. First I say, thestudy of animal behavior is a trap. Once you startobserving animals, and truly watching what they do, youcannot stop. I add that I am continually amazed by whatanimals are capable of in their lives and am fascinatedwith and challenged by trying to understand thatcapability. I sometimes tell a story I heard from EdWilson of a woman, who after one of his lectures, cameup and asked "Dr Wilson, I have a serious problem withants in my kitchen. I can't get rid of them. Do you haveany suggestions of what I should do"? Supposedly Edreplied "Watch them!" Second, I say, aggression is afundamental component of the behavior of all animals(well, I've asked my colleagues whether nematode wormslike Caenorhabditis elegans fight, and they claim they donot). Aggression is necessary for access to territory, foodand mates in what for most animals is a hostile world.While manifested in forms given names like maternalaggression, territorial aggression, or predator/preyaggression, all forms of aggression may actually involveclosely related or identical final common pathways. Tome, sorting out and understanding those pathways is anenormous challenge, and I enjoy enormous challenges.And finally I comment that violence is a serious problemin our society. It's a problem with a biological basis, Inote, but we have little knowledge or understanding ofthe underlying mechanisms. Therefore, I finish with, ourhope is that we can contribute something to helpingpeople understand and deal with the origins of violenceand possibly, just possibly, contribute to its treatment. The question that almost always follows, however, is"If you do gain knowledge of the biological basis ofaggression, couldn't that be used by governments tocontrol people, or by the military to generate super-soldiers"? To this my reply used to be that the morepublic and widely disseminated information is, the lesslikely it is to be useful for the generation of weapons. Isay "used to be" because much has changed sinceSeptember 11, and now we must ask whether thosechanges of necessity influence the ways in which scienceis and should be used in the future. It's interesting thatprecisely the same argument I used to offer was used injustifying the publication of a six-volume series of booksin 1973 called "The Problem of Chemical and BiologicalWarfare" by the Stockholm International Peace Research

Initiative.1 The authors of these volumes state that "Adifficult dilemma has confronted us in this task. Incollecting together and publishing material necessary fora clear understanding of the problems of CBdisarmament, might we not be compromising ourobjective by disseminating information that couldpromote proliferation of CB weapons"? They add,however, "Elsewhere we have argued that the service wecould do by improving the level of public discussionwould be greater than any disservice in transmittingdangerous knowledge." In fact these authors too felt thatnations were less likely to produce weapons that couldjust as easily be used against them and threaten them withextinction. That fear and a small moral compunctionagainst the use of these weapons led to the generation andsigning of a comprehensive international treaty in 1975,The Biological Weapons Convention,2 banning theproduction and use of weapons of biological warfare.Article one of the Biological Weapons Convention reads"Each State Party to this Convention undertakes never inany circumstances to develop, produce, stockpile orotherwise acquire or retain: (1) Microbial or otherbiological agents or toxins whatever their origin ormethod of production, of types and in quantities that haveno justification for prophylactic, protective or otherpeaceful purposes." While the Convention was a clearstatement of intent to ban these materials as sources ofweapons, no means of verification was included in theoriginal document and modern technology has producednew categories of organisms that nations and individualsargue are not included. In the present climate, however,the most important new element introduced by the eventsof September 11 and its aftermath are that we no longerare dealing with States and well defined borders and weno longer are dealing with individuals concerned aboutself-preservation. In 1993 a Chemical WeaponsConvention banned the development, production andpossession of chemical weapons, and this has beenratified by 135 countries including the United States. My first introduction to what chemical and biologicalwarfare was all about began innocently enough. In themid-1960s I was visited by four generals from the UnitedStates Army Chemical Corps, just at the time our work onGABA as a transmitter compound was beginning to berecognized. Frank (F. O.) Schmitt3 had called me fromMIT that morning to ask whether these officers couldvisit me. Frank was familiar with our work on GABA asa transmitter compound and the single cell approaches wewere taking to explore neuronal function, as I hadpresented the work at several Neurosciences ResearchProgram workshops in the early 1960s. I had no ideawhy military people wanted to talk with me, but beingyoung and easily flattered by the attention, I agreed to thevisit. A short time later, four distinguished, highlybraided individuals showed up in my shared little office.They asked first about my work and then started

15

questioning me about people who might be doing similarwork in vertebrate systems. It slowly dawned on me whatthis was all about. They were there to find out whetherthere was military potential in the work I was doing forwe were working on what turned out to be the majorinhibitory transmitter compound in the nervous system ofall species, including man. A few years later, at the time of the Vietnam War, Iwould have refused to see these military officers. Butwould I have been right in doing so? After all, myresearch was entirely supported by U. S. governmentfunding, these men dedicated their lives to the protectionof our nation, and here, they felt, was a possible newsource of weapons to defend our nation. Even more -what would I do now, in the face of horrific terroristactions that threaten the entire civilized world? Would Inow be willing to meet with a contingent of militaryofficers intent on using information I supplied them withto develop anti-terrorist weapons based on my research? My lectures to Harvard Medical students in the late1960s and early 1970s focused on acetylcholine as atransmitter compound, and included pointed commentsabout the development of nerve gases, their stockpilingand their potential use as weapons in new kinds of warsenvisioned by military minds in books like "Tomorrow'sWeapons."4 Since the commonly available nerve gaseswere substrates for, and then covalently-linked essentiallyirreversible inhibitors of acetylcholinesterases, theenzymes required for inactivation of acetylcholine atneuromuscular junctions, they had the potential to serveas highly potent weapons of war killing people withinminutes of exposure. In "Tomorrow's Weapons," theauthor J. H. Rothschild discusses the wonders of usingweapons of this type to incapacitate or kill people withoutdamaging property, a concept that chilled me. In onescary scenario Rothschild describes tests using the drugLSD-25 on a military command center and incapacitatingthe human subjects involved. His comment on this resultwas "Think of the effect of using this type of materialcovertly on a higher headquarters of a military unit, orovertly on a large organization. Some military leadersfeel that we should not consider using these materialsbecause we do not know exactly what will happen and noclear cut results can be predicted. But imagine wherescience would be today if the reaction to trying anythingnew had been 'Let's not try it until we know what theresults will be'." The journal Military Review in 1970 featured an articleby Carl A. Larsen entitled "Ethnic Weapons"5 dealingwith polymorphisms that naturally exist in enzymeswithin different ethnic groups and the possibility ofexploiting these differences to produce selectivevulnerabilities to drug attack. Then there was the plan bythe military, uncovered in 1969, to make a "routine"shipment by freight train of 27,000 tons of chemicalwarfare agents across the United States for dumping in

the ocean, because the weapons were leaking andbecoming dangerous to stockpile. Half of that shipment,that was to go through major population centers likeIndianapolis, Dayton, Knoxville, Cincinnati, Philadelphiaand Elizabeth, N.J., contained the nerve gas GB. Anarmy officer supposedly told a newsman at the time that"the gas from a single bomb the size of a quart fruit jarcould kill every living thing within a cubic mile,depending on the wind and weather conditions."6 Thearmy also initially claimed they were not responsible, andthen admitted responsibility in the death in 1967 of 6400sheep in Utah, in farmland close to the Dugway ProvingGrounds where secret airborne tests of chemical warfareagents were being carried out. Was it any wonder that Iwas lecturing to future doctors about the horrors ofchemical warfare and about the callous way thesematerials were being discussed and handled by themilitary? What also was distressing to me at the time wasthat scientists had to be involved in all of these activitiessince many scientific, technical and technologicalproblems had to be solved to make effective weapons ofchemical and biological agents. And now we have anthrax -- weapons-grade anthrax.While the United States relentlessly bombs an invisibleenemy in a far-away country, a very few envelopescontaining a deadly form of what originally was abacterial pathogen of animals have caused great concern,even panic within the population at home. It impacts onall of us and on our daily lives in ways that would havebeen inconceivable just a few short weeks ago. Toillustrate, I received an envelope yesterday from SantaBarbara, California without a return address. I don'tknow anyone in Santa Barbara, nor was I expecting mailfrom that part of the country. Should I throw the letterout unopened? Deciding instead to open the letter, I puton gloves and a face mask and carefully slit the envelopeopen at the top. Of course, there was nothing of danger inthe envelope - it contained an unsolicited reprint sent outin a general mailing. In our departmental business officetwo days ago an assistant found dust on her desk afteropening the mail (the office had received razor blades inan earlier mailing leaving people on high alert) and calledthe police. This led to a quiet closing of the floor and anexamination by the bomb squad and decontaminationagents of what turned out to be just dust. The anxietylevel is high in the American populace at the present timeand it is not reassuring to know that our government'spresent policies of retaliation are generating even largernumbers of future enemies of our country and its policies.Wouldn't it be better if we were making as much effort atfeeding the starving refugees in Afghanistan as we are atdestroying the very small amount of infrastructure thatexists in a country that has been at war for decades?

The anthrax mailings must involve scientists andengineers who have been trained in the handling ofdeadly organisms and in generating forms suitable for

16

transmission through the air. There is a reasonably highlikelihood that these individuals learned the care andhandling of toxic organisms in our own universities andresearch laboratories. Moreover, from the media andgovernment sites thusfar targeted, the perpetrators couldjust as well be any of a number of right-wingorganizations in the United States as foreign international

terrorists. Once again, the question surfaces though: Howcan scientists act in ways that pervert their science toinhumane actions of these sorts? Is there any way toensure that, just as doctors have their Hippocratic Oathpromising the delivery of their art to all in need of it,scientists could be persuaded to use their science only forthe betterment of mankind?

Actually there is no single form of theHippocratic Oath, as the original is very outdated in amodern world, but most doctors in the United States onreceiving their medical degrees swear to some form ofHippocratic Oath (98% of U.S. Schools administer anoath). All versions include phrases dedicated to usingmedical knowledge for helping, not hurting people andfor offering medical aid to all in need of that aid. Arecent issue of the Harvard Medical Alumni Bulletin(Summer 2001) featured a section called "What did youdo in the war, Doctor?" that brought home the strength ofswearing to an oath of this sort in reminiscences of WorldWar II by Harvard Medical School-educated physicians.Most of the articles talked about how battlefield medicalstations treated American and German wounded soldiersequally as the casualties of war mounted. Surprisingly onresearching the Hippocratic Oath on the web, I discoveredthat there have been numerous attempts to develop aHippocratic Oath for Scientists as well. The goal is tohave all who receive an advanced scientific degree swearan oath to use their scientific knowledge for thebetterment of mankind. Here's one version that Iparticularly like. "The Hippocratic Oath for Scientists,Engineers and Executives proposed by The Institute forSocial Inventions.7 I vow to practice my profession withconscience and dignity; I will strive to apply my skillsonly with the utmost respect for the well-being ofhumanity, the Earth and all its species; I will not permitconsiderations of nationality, politics, prejudice, ormaterial advancement to intervene between my work andthis duty to present and future generations. I make thisOath solemnly, freely, and upon my honor." Would this work? Would this prevent science andscientists from using their art in perverse ways? I doubtit, but it just might make someone about to embark on acontrary route pause before moving in that direction.

References and footnotes:1. Almqvist and Wiksell, Stockholm; Humanities Press, NY;and Paul Elek, London.2. The Biological Weapons Convention was signed in 1972,entered into force in 1975 and ratified by 143 State Partiesincluding the United States. A previous international ban onchemical and biological weapons, called the Geneva Protocolof 1925, was never ratified by the United States.3. F. O. Schmitt was the founder of the Neurosciences ResearchProgram (NRP), whose board was a group of distinguishedinvestigators from many disciplines who viewed their role as

interpreting the implications and future directions of research inthe nervous system.4. "Tomorrow's Weapons-Chemical and Biological", by J.H.Rothschild, retired U.S. Army Brigadier General, published in1964 by McGraw-Hill Book Company.5. Military Review (November 1970), pp 3-11.6. "Germs and Gas as Weapons" by Seymour M. Hersh in theNew Republic, June 7, 1969.7. The Institute for Social Inventions, 20 Heber Road, LondonNW2 6AA, UK.

MEETINGS AND COURSES

Marine Biological Laboratory Summer CourseNeural Systems and BehaviorJune 16 - August 10, 2002Directors: Catherine Carr, Univ. of Maryland and Richard Levine, Univ. of Arizona An intensive eight-week laboratory and lecture coursefocusing on the neural basis of behavior, from the cellularand synaptic levels to the analysis of complex systems.Intended for graduate students, postdoctoral students, andindependent investigators. Limited to 20 students. The central theme of the course is the investigation ofhow neurons and neural circuits produce behavior andplasticity. Laboratory and lecture components combinestate-of-the-art neurobiological techniques withbehavioral and developmental analyses. The lecture seriesbegins with a consideration of electrophysiological andanatomical principles of neuronal function. Topics thenmove on to how properties of individual neurons cometogether in simple neural networks for behaviors such aslocomotion, escape, and the generation of rhythmicpatterns of activity. Modulation of neural activity andneural circuits by transmitter and hormone action, long-term potentiation, and the biophysical basis of sensorytransduction are covered. Finally, emphasizingcomputational approaches, we consider questions such ashow animals process complex auditory stimuli oraccomplish spatial and vocal learning. Weeklyseminars are given by invited lecturers and distinguishedScholars-in Residence. The heart of the course is the laboratory, whereadvanced techniques in cellular neurobiology are broughtto bear on neural systems. Methods taught includeintracellular recording; single cell dye-injection; singleand double-electrode, patch, and whole-cell voltageclamp; analysis of synaptic transmission and plasticity;

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brain slice; computational and behavioral approaches. Avariety of vertebrates and invertebrates serve asexperimental systems. For further information and application forms, see thecourse web site: http://courses.mbl.edu/

Summer course in "Chemosensory Neurobiology in theMarine Environment" at the Bermuda BiologicalStation for Research, 16 June 5 July 2002 (3 weeks). Wewill study chemosensory neurobiology in the marineenvironment at the physiological, biochemical, andmolecular levels. This course emphasizes experimentaltechniques and approaches to the study of chemosensorybiology, and is designed to benefit graduate students andadvanced undergraduates who have interests inorganismal, systems, cellular, or molecular biology.Laboratory exercises and research projects will use theolfactory system of the spiny lobster, Panulirus argus, asthe main teaching and research tool. Receptor cellelectrophysiology, immunocytochemistry, BrdU labelingof cell proliferation, biochemistry of receptor andperireceptor phenomena, and molecular biology (PCR,sequencing, and other techniques) will be taught. Formore information, contact: Charles Derby, Dept. ofBiology, Georgia State Univ., P. O. Box 4010, Atlanta,GA 30302_4010; cderby@gsu.edu, (404) 651_3058(office), http://www.gsu.edu/~biocdd/

NEW BOOKS BY ISN MEMBERS

A SPIDER'S WORLD _ SENSES AND BEHAVIOR.Friedrich G. Barth, 2001, Springer_Verlag (ISBN3_540_42046_0, Hardcover, approx. 400 pp., 309 figs.,16 color plates, US$ 69.95, BP 48._, DM 129.90, FF524). In this book the technical perfection of spidersensory systems for different forms of stimulus energy isput into a broad biological and neuroethological contextwhere the senses serve as links between the speciesspecific environment and behavior.

AUDITORY WORLDS: SENSORY ANALYSIS ANDPERCEPTION IN ANIMALS AND MAN, edited byGeoffrey A. Manley, Hugo Fastl, Manfred Kössl, HorstOeckinghaus and Georg Klump. XIX + 359 pages, 76Figures, 6 Tables, DM 198,_ ISBN 3_527_27587_8,Wiley_VCH Verlag,Weinheim 2000. Design plasticity inthe evolution of the amniote hearing organ; Comparativeanatomy and physiology of hearing organs; Cochlearfrequency maps and their specializations in mammals andbirds; Models of the human auditory system; 5 Activemechanics and otoacoustic emissions in animals and man;Neural processing in the brain; Comparative animalbehaviour and psychoacoustics; The human aspect I:Human psychophysics; Hearing impairment: Evaluationand rehabilitation

POSITION ANNOUNCEMENTS

ASSISTANT PROFESSOR ARTHROPODBEHAVIOR/NEUROBIOLOGY Univ. of California,Berkeley. The Division of Insect Biology, ESPM Dept., invitesapplications for a tenure_track 9_month appointment at theAssistant Professor level in any area of insect or terrestrialarthropod behavior or neurobiology, starting July 1, 2002. Thesuccessful applicant must have a Ph.D. in the biologicalsciences, an excellent record of scientific accomplishment,strong commitment to undergraduate and graduate teaching,and an interest in being part of a community of insectbiologists.Applicants should submit a curriculum vitae, copiesof recent publications, statements of research and teachinginterests and experience and three letters of references sentto:Professor Wayne M. Getz, Chair, Behavior/NeurobiologySearch Committee, ESPM_Division of Insect Biology,201Wellman Hall, Univ. of California, Berkeley, CA 94720_3112.The closing date for applications is January 4, 2001. Thedeadline for the letters of reference is January 18, 2002.

The Psychology Dept. at the Univ. of Maryland, CollegePark, has a tenure-track position at the Assistant Professorlevel for a neurobiologist who uses animal systems, preferablyinvertebrate, to study learning, neural plasticity, or adaptivechanges in the nervous system that occur during development.Research using molecular and genetic techniques is welcome,as long as it remains strongly linked to behavior. Stronghistory of research productivity and active scientific programrequired. Teaching will include graduate and undergraduatecourses in behavioral neuroscience, neuroethology, andneuropharmacology. For more information visit:http://www.bsos.umd.edu/cebh/INSjob.html. Applicantsshould send CV, statement of research interests, and arrange forthree letters of recommendation to be sent to: Dr. RobertDooling, Chair, Neuroscience Search, Psychology Dept., Univ.of Maryland, College Park, MD 20742-4411. Univ. ofMaryland actively subscribes to a policy of Affirmative Actionand Equal Educational Employment Opportunities. For bestconsideration, materials should be received by 15 November2001.

Tenure track faculty position in Cellular and MolecularNeuroscience in the Dept. of Biology at the Univ. of Maryland.The successful candidate will direct an innovative andcompetitively funded research program, and be part of theneuroscience program (http://www.life.umd.edu/NACS/). Thecandidate will also participate in the instructional program atboth the graduate and undergraduate levels(http://www.life.umd.edu/biology). UM is the flagship campusof the University of Maryland System, and is located in theheart of the Baltimore_Washington research corridor.Applications should be received by November 15, 2001 toreceive full consideration but the search will continue until the

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position is filled. Materials, including a letter of application,curriculum vitae, statement of research and teaching interests,and the names and E-mail addresses of four references shouldbe directed to: Neuroscience Search, Dept. of Biology, Univ. ofMaryland, College Park, MD 20742_4415. The Univ. ofMaryland is an Affirmative Action/Equal OpportunityEmployer.

The Dept. of Biological Sciences at Vanderbilt Univ. seekscandidates to fill a rank-open, tenure-track or tenured facultyposition in cellular/molecular neurobiology. It is part of thegrowing neuroscience community at Vanderbilt with stronggraduate and undergraduate programs. Women and minoritycandidates are especially encouraged to apply. Applicantsshould send a letter of application together with a curriculumvitae, a statement of current and future research interests, andselected reprints to Neurobiology Search Committee, Dept. ofBiological Sciences, Vanderbilt Univ., VU Station B 351634,Nashville, TN 37235-1634 U.S.A. Junior faculty applicantsshould arrange for three letters of recommendation. Seniorfaculty applicants should provide a list of at least six references.Review of applications will begin October 29 and will continueuntil the position has been filled.

TRAINING OPPORTUNITIES

Seeking Masters of Science Student. Our current researchemphasis is on the mechanisms of water_flow sensitivity in thenudibranch sea slug Titania diomedea, the integration ofsensory information, and motor control of crawling and turning.Training in brain cell recording, computer analysis of video,microscopy, and underwater animal observation is available.Financial support is available during the academic year and thesummer. Applications will be accepted from biology andcross_disciplinary students (physics, chemistry). The BiologyDept. contains faculty including two neurobiologists, a sensorybiologist, and another electrophysiologist with advancedcourses in animal physiology, molecular biology systematics,and neurobiology. Please send a curriculum vita, unofficialtranscript, a 1_page statement of research interest, and thenames and contact information (preferably e_mail) of 2references to: James A. Murray, Ph.D. Assistant Professor ofBiology 156 Lewis Science Center Univ. of Central ArkansasConway, AR 72035, (501) 450_5923,E-mail: jmurray@mail.uca.eduhttp://www.uca.edu/divisions/academic/biology/faculty/jmurray.htm

Postdoctoral Position in Sensory Physiology A postdoctoralposition to study auditory processing and plasticity in barn owls

is available in the laboratory of Prof. Eric Knudsen at StanfordUniv.. We focus on neural mechanisms underlying soundlocalization and how they are shaped by auditory and visualexperience. We use neurophysiological, pharmacological,anatomical and behavioral techniques. Experience withelectrophysiology preferred. Please contact Dr. Eric Knudsen,Dept. of Neurobiology, Stanford Univ., Stanford, CA. 94305-5125. e-mail: eknudsen@stanford.edu

Postdoctoral Fellowships in Neuroethology. TheNeuroethology Training Program at the Univ. of Maryland isseeking qualified applicants for postdoctoral fellowships tostudy the relationships between neural circuits, behavior andevolution. Opportunities to conduct research on soundlocalization (Carr), comparative psychoacoustics (Dooling),locomotion (Cohen), evolution of hearing (Popper),echolocation (Moss), auditory physiology (Yager), visualsystem (Hodos), reproductive behavior (Ottinger), biologicalmodels of learning (Halperin), evolution of developmentalmechanisms (Jeffery), development of auditory CNS (Hall),behavioral genetics and sexual selection (Shaw), and songdevelopment (Troyer). Specific interests of the faculty and acomplete description of the program can be found athttp://www.bsos.umd.edu/psyc/neuroethology/. Applicantsmust be U.S. citizens or permanent residents, and should sendtheir CV, a brief description of research interests and the namesof three references to: Dr. Cynthia Moss, Dept. of Psychology,Univ. of Maryland, College Park, MD 20742-4415. Univ. ofMaryland is an Affirmative Action and Equal OpportunityEmployer

Postdoctoral Fellowships in Auditory Neuroscience. TheComparative and Evolutionary Biology of Hearing TrainingProgram at the Univ. of Maryland seeks applicants forpostdoctoral fellowships to study auditory neuroscience. Theprogram (www.life.umd.edu/biology/cebh) has recentlyexpanded and includes 11 faculty on the College Park campusas well as 12 additional faculty from the intramural program ofthe National Institute on Deafness and CommunicationDisorders of NIH. Research organisms extend from insects tohumans, and research approaches from molecular biology towhole brain imaging. Specific interests of the faculty and acomplete description of the program can be found on our website. Applicants must be U.S. citizens or permanent residents,and should send their CV, a brief description of researchinterests and the names of three references to: Dr. Arthur N.Popper, Dept. of Biology, Univ. of Maryland, College Park,MD 20742. Univ. of Maryland is an Affirmative Action andEqual Opportunity Employer

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Meeting organizer Horst Bleckmann (Bonn) relaxes with TedBullock (San Diego).A Postdoctoral Research Scientist position is availableimmediately to investigate the cellular and molecular control ofpost-embryonic development in the olfactory system of a modelanimal, the spiny lobster. Neuronal proliferation,differentiation, and turnover will be studied using techniquesthat may include in vivo and tissue culture,immunocytochemistry, in situ hybridization, gene expression,and electrophysiology. Salary is $27,000-40,000, depending onexperience. GSU is part of a well-equipped, well-funded, andhighly interactive Atlanta Neuroscience community thatprovides opportunities for collaborative investigations,including the NSF-funded Atlanta-wide Center for Beh