The Senses of Fish: Adaptations for the Reception of Natural Stimuli PDF
Preview The Senses of Fish: Adaptations for the Reception of Natural Stimuli
The Senses of Fish Adaptations for the Reception of Natural Stimuli The Senses of Fish Adaptations for the Reception of Natural Stimuli Editors Gerhard von der Emde Joachim Mogdans B G Kapoor Springer-Science+Business Media, B. V. A. C.I.P. catalogue record for the book is available from the Library of Congress ISBN 978-94-010-3779-2 ISBN 978-94-007-1060-3 (eBook) DOI 10.1007/978-94-007-1060-3 In India, Bangladesh, Nepal, Pakistan and Sri Lanka by Narosa Publishing House 22 Daryaganj, Delhi Medical Association Road, New Delhi 110 002, India All Rights Reserved Copyright © 2004 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2004 Softcover reprint of the hardcover lst edition 2004 Ali rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission of the publishers. Foreword FISHANDFOLK V. Benno Meyer-Rochow International University Bremen (IUB), FacultyofEngineering & Science, Bremen,Germany Ilovefishand when asked towritethis Foreword wasveryhappy tobegiventhe opportunityto welcome this exciting tome on "The Senses of Fish: Adaptations for theReception of Natural Stimuli", edited by Profs. Gerhard von der Emde, Joachim Mogdans, and B.G. Kapoor. My enthusiasm was briefly damped when a graduate student of mine, whom I told of the task allotted tome,remarked "Whostillreadsbooks these days?".Temporarilystunned, Ihastened toreply"Smartpeople do,ofcourse!". This volume contains such a wealth of information and presents such a wonderfully balanced variety of contributions byexperts in their own fields, each and all expanding their subject one or another, that I could have added "This book makes smart people smarter still and that's whysmart peoplewillread it".The topicscovered fallneatly intosixcategories.The first twochapters, byBarbara Evansand Arnold Sillman & David Dahlin, provide the reader withexcitingnewdetailson fisheyeadaptationsand the reasonsbehindsuch adaptations. Eyes and vision in fish are finely tuned to the photic environment and as a consequence of this relationship aswellasthe widerange ofhabitats thatfishoccur in,avariety ofmodifications to the eye'soptics,development, retinal organization, photopigments, absolute and spectral sensi tivities have evolved. Without a thorough examination of these adaptations, we would be unable tomake muchsenseofthe visualbehavior offish. Recent yearshaveseenan upsurgeofinterestin the chemosensory abilitiesoffishand three chapters, one each byAnne Hansen& KlausReutter,Tine Valentineie, and GabrielleNevitt& Andrew Dittman, testifyto thisfact.Usingbehavioral responsesand a comparative approach, Tine Valentineie showsthat predatory fishwith functional olfactoryand taste receptors do not necessarilyusethese sensesduring foodprocurement.Visualhunters usethe taste systemsolely during oral food evaluation. The situationfor the detectionofunderwatersmellsissomewhat different as Hansen and Reutter demonstrate. However, that olfaction in migratory species playsamaior rolehas been known foralongtime, but how,forinstance, imprintinginjuvenile vi Foreword salmon might work, in contrast to imprinting in mammalian species, isinvestigated byNevitt and Dittman. The two authors provide evidence that odor memories may also be retained peripherally and thatolfactory receptor neurons could be "selectively tuned" byresponding to specificodors present"during ahormonallylinkedsensitive period". Thisisan exciting concept and one can expect more fascinating discoveries in this fertile field of 'developmental neuro etho-ecology'. A fascinating overview of the sensory systemsand sensory brain areas in deep sea fish by Hans Joachim Wagner allows the reader not only to appreciate the specializations that are present in the denizens of deep water, but also to see the developmental parallels in fishes of otherhabitats (cf.,forinstance,Meyer-Rochow &Coddington, 2003, In: FishAdaptations,eds. ValA.L. & Kapoor B.G., Oxford and IBH Pub!. Mis Sci. Pub!., Enfied, New Hampshire, pp. 339-383). This chapter byWagner, placed in the centre of the book, thus connects the more detailed treatments of selected senses and sensory adaptations, preceding and following Wagner's chapter, with eachotherand, likethe fish'sbrain, has an integrativefunction. Aspects of sound production and hearing in fish are dealt with in another fascinating section of this book. Far from being silent creatures, Friedrich Ladich introduces us to the 'noisy' underwater world and describes how and why fish produce sounds. Little known and poorlystudies isthe abilityofsome speciesoffishto detectsounds ashigh as 180kHz.Adapta tions responsible for this remarkable feat, in particular within the Clupeiformes, and possible ecological consequencesofultrasound detectionare being reviewed byDennis Higgs.But how the acousticsignalsin the end are 'heard'and processedbythe fish'snervoussystemisthe topic ofZhongminLu'schapter.Concentratingon what has beendiscovered on the goldfish'ssound detecting pathways, Lu alsoexamines acoustic communication and directional hearing in the mormyrid Pollimyrus adspersus, the midshipman Porichthys notatus, and other species. Hong YoungYanthensummarizesthe currentstateofknowledge with regard to hearinginfishesand emphatically draws attention to the fact that despite considerable progress, many aspects of sound perception in fishare still poorly understood. A very comprehensive treatment byJohn Janssen is given to the role of the lateral line, especially in species that inhabit dark environments or turbid waters. Yet,how the different species of fish integrate the signals coming from the lateral line with those being received through channels of the other senses is still largely unknown. Joachim Mogdans, Sophia Krother, &Jacob Engelmannin theirchaptertouch upon this problem, butdeal primarilywith recentfindingson the lateral linebrainstem's responses to avariety ofstimuli.Twosubsystems, consisting of superficial neuromasts whose function is impaired in running water, and canal neuromasts, whose function remains unaffected in runningwater,were discovered and led the researchers to conclude the presence of a clear form-function relationship for the lateral line. Finally,the enigmaticelectricsenseofsomefish:Thattopicformsthe subjectofchaptersby LonWilkens, Tim Tricas&JosephSisneros,Clifford Keller, and MaryHagedorn. Wilkens intro duces us to the paddlefish, a fish that rarely features as an experimental subject. However, as Wilkens ever so ablydemonstrated with his remarkable research, the paddlefish usesrostrally located electroreceptors to detect plankton and to investigate its environment. A form of Foreword vii "passive electroreception", and what exactly this term entails, is revealed in the chapter by Tricas and Sisneros. The two researchers have chosen skates as their experimental subjects, marine species in other words, which, unlike the weakly electricfish (dealtwith in the chapter by Keller), are not at all famous for communicating electrically. Keller leads us into an alien world of senses, into a world in which signals familiar to us humans have become largely replacedbyelectricalstimuli.Aimed primarily at conspecifics,suchsignals,likeMary Hagedorn explains, can also be apowerful incentive to those adventurous humanbeings, wishing to work with fish in remote junglesettings. Although the contributors to this book hail from several different countries and presum ably were not brought up in the same cultural environments, their fascination for fish unites them all. As someone who wasborn under the sign ofPisces, Itoo, have a"softspot" forfishof allkinds: Ilike themfried, cooked, and smoked, and Iregularlyenjoyed 'gefilte fish' asa boy in my grandmother's home. But I am equally fond of them when I find them cut open in the dissection dish, under the microscope, or placed in some apparatus designed to reveal their physiological secrets.Ilove watching themdartaround in myaquariumathomeand havegone diving to see them in their natural habitats. Ieven have a collection ofcoins that feature fish and Iknowpeople, who possessthousands offishstampsfrom allover the world. Obviously,as the enthusiasm has shown, with which this book's contributors have written their chapters, there are manydifferent waysto express one's adorationforfish. What makes fish so attractive to humans? Where does this age-old fascination for the slippery creature of the aquatic domain come from? Isit because we all pass through fish-like stages in our embryological development? Is it because the class Pisces outnumbers all other vertebrate classes with regard to the number of species (at least 25,000 species have been described to date)?Orisit,becausefishare seenasinhabitantsand messengersofaworld alien, and maybe even threatening, to humans? Iguess there isn't a single reason why humans since ancient times have found fishcaptivating, butfact isthat the humblefish, as asymbol, has had averylong historyand occursin cultures and religions throughout the world.So fond must the earlyEnglish havebeenoffish (orwere theysimplybad observers?) thattheyevenlentthis term to animals, which have little to do with real fish (witness shellfish, starfish, crawfish, jellyfish, cuttlefish, crayfish, and silverfish). Christiansrevere thefishasasymbolofChristand inJewish lore thefish,beingcustomarily consumed on the Sabbath, symbolizes prosperity. Pre-Christian Germanic tribes ate fish in honorofthe greatmothergoddess Freya,ahabitthataccordingtosome scholarspersists to this day in the traditionofmanya Europeanto have ameal offishonFridays.In India areincarna tionofthe goddess Durga, knownasMeenakshi, wasrespectfullycalled'thebeautiful fish-eyed one'and the firstIndiancoinageof600-300BC has manyexamplesoffishmotifs. Severalother non-IndiandeitieslikeIsis,Apollo,and Poseidon (theGodoftheSea) were alsoassociatedwith fish.Amongstthe Maya oftheNew World differentspecies offishhelddifferentmeanings: hags were seenasspies and destroyers, catfish represented cleaners and renewers, and otherspecies were invoked in local systems ofcounting. InJapan, on the day known as 'kodomo no hi' (May 5th each year), even today tall bamboo poles with carp kites and streamers are erected in the viii Foreword gardens ofmanyahome,fortotheJapanese the carpembodiesstrengthand determination.On the otherhand,fearand panic have nowadays,unfortunately, become almost synonymouswith sharks around the world. However, actual shark-worshippers stillinhabit some remote islands likeMalaita ofthe Solomons in the WestPacific. Clearly,fish have accompanied humanity and human culture from their beginnings, and this has alsofound numerous reflections in the arts. Poemsin praiseoffishhave been written and set to music (ESchubert's composition "Die Forelle"comes to mind), fishpaintings (e.g., PaulKlee's"GoldenFish"),fishsculptures (the OlympicfishofBarcelonabeatingthemallwith its gigantic size), and nowadays even fish movies, fishing competitions, and fish games exist. Stories, fairy tales, and fables involving fish, often rooted in antiquity (who would not have heard ofAesop's 'Fishermanand the Little Fish'?) have along history,butthey are stillpopular today and they are being told in virtually allparts ofthe world. This book, although anything else but a collection of mere fishystories and fables, does follow that long tradition of human fascination with fish and there is not doubt that the scientific research, reported in this volume, represents the fruits of years of patient and dedi catedstudy.Theauthors, editors, and publishersmustfeelprivilegedtoshare in the production ofsoworthy a text asIdo to have been called to welcome it on its appearance. As wegather moreand moredataon the wondrous waysthe fishes'function',weslowlystarttograspwhy,for at least400millionyears,these animalshave managed toflourishand beenable tolivethrough gee-historic crises and environmental changes too profound for us to comprehend. We are reminded of the fact that almost three quarter of the Earth's surface are the domain of our friends, the fish.Andwearerealizingthatweareinthe processofunderstanding, what kindsof adaptationshave allowedthemtosucceed inoccupyingeveryconceivablenicheoftheirwatery realm and what roles their sensory capacities have played to guarantee their survival through the ages. Much ofwhat weknow today about the sensesoffish,represents abilities and adaptations almost unimaginable to us'mere humans'onlyafewyearsago.This isafact and there isnoth ing'fishyabout that. However, none of the exciting discoveries can diminish the deep respect wehold forour 'scientific quarry'. Onthe contrary, the discoveries have served toenhanceour appreciation of this formidable creature ofNature and sometimes mayeven cause usto take a deep breath and reflect on our own imperfections. BeforeIend this Forewordwith a translationofthat fitting and famous poem "Die Forelle/ TheTrout" byChristianSchubart, Ishould liketo emphatically state: This book cannot failto givegreat pleasure to allthose people smart enough to pick it upand read it thoroughly! Foreword ix The Trout Beneaththe limpid water Isawajollytrout As swiftas arrow darting And flashing in and out. Istood upon the margin, All in the morning cool, And watch'd the fishdisporting Downin the crystal pool. Nearbythere stood an angler, With rod and line and hook, And saw the trout a-swimming Downin the crystal brook. Withshine upon the water Unbroken,so Ithought, In vainwillbe his angling, The fishwillnot be caught. Atlast the thievish angler Impatientgrew.Withguile The waterclear he ruffled, Andin a little while He jerk'd hisrod in triumph. And struggle, struggle asit may, The poor outwittedvictim Upon the greensward lay. From:GarranR.R. 1946(reprinted1971) "SchubertandSchumann".MelbourneUniversity Press Preface Animals have to be able to perceive external sensory stimuli and generate appropriate behavioral responses in order to live successfullyin their environments. On an ontogenetic scale,behavioral responses to sensorystimuli are eitherinnate or learned. Learning enables an animal torespond tochangingstimuli throughexperiences madeduring life.Onaphylogenetic scale, organisms also 'learn' to adapt to changed signals in their environment. Evolutionary learning works in a different time frame, however, that is from generation to generation. Its motor is selection, and its result is an adaptation of the sensory apparatus and the nervous system.During this adaptation process,'communication' with the environment ismaintained, enabling the animal to gather important signalscoming either from the surroundings or from otheranimals, likeprey, predators, or conspecifics. In this book we use fishas model systemsto demonstrate how evolutionary learning has shaped various sensorysystems.Why fish? There are three main reasons for this: First, 'fishs' comprise a huge and diversegroup ofanimals.There are about 25000 extant fishspecies that live in such diverse environments like the deep sea or shallow freshwater puddles in the Himalayas.Closelyrelated taxa mayliveindifferent habitats and thus undercompletely differ, ent environmental conditions. They have adapted to these diverse environmental conditions not onlyin their externalbodyfeatures, but alsobymodifyingtheir sensoryapparatus. Second, fish possessmore senses they many other animals. They are equipped with the well known senses that humans have, but in addition they possessat least three more senses. Third, fish stand at the basisofvertebrateevolution.While shaping the sensesoffish,evolutionworkedon the very 'blueprint' of vertebrate organisation, which forms the basisof all other vertebrates, including humans. Fish thus can serve as model systemsfor vertebrate evolution in general, including the evolutionofsensorysystemsand their adaptations to environmental conditions. Webrought together scientific experts fromdifferent areas, who report about the sensory systemsof fish and unravel their physiologicaland ecological evolutionary adaptations. The first two chapters deal with vision,one of the most important sensesofvertebrates. However, vision in different aquatic habitats has to cope with quite diverse photic conditions and thus requires fundamental adaptations of the eyes and their sensory receptors. Chemoreception, dealt with in chapters three to five,maybeone ofthe "oldest"sensesofvertebrates. Yetagain, xii Preface environmental conditions differ in the various habitats fish live in, and this requires adaptations ofthe olfactoryand taste systemsofthe different groups.Aninterestingcasestudy ofolfactoryspecializationisolfactoryimprinting,whichcanbefound inseveral groupsoffishes. The best-known example may be olfactory imprinting in the service of homing in salmons as described inchapterfive.Chaptersixdoesnotdealwithaparticularsensorysystem,but takesa lookat several groupsoffishlivingin an extreme environment:the deep sea.Here, the chemi calsenses,vision,and the laterallinesystemareespeciallydevelopedand adapted to the differ ent niches in this ultimate environment. Thatfishare not deafiswellknownsince the daysof Karl von Frisch, and chapters seven to ten discuss several aspects of underwater hearing. Related tohearing, sound productionbyfish(chapter 10),mainlyin the service ofintra-specific communication, isan important evolutionary factor shaping fish hearing organs. The lateral line systemoffishcompromises two main sub-systems: the mechanosensory lateral line, dealt with in chapters 11and 12,and the electrosensory lateral line (chapters 13 to 16).The latter canbe divided stillfurther, dependingon the typeofstimuli that are perceived:lowfrequency electric signalsfor'passive' electrolocationthroughampullary electroreceptororgans (chapters 13 and 14), and high frequency signals, actively produced and thus involved in 'active' electrolocation (chapter 15).Finally, chapter 16stressesan important'human' point:scienceis made bypeople,sometimes underextremeconditions.Especiallywhen studying environmental adaptations ofanimals, it isnecessaryto gooutinto the fieldand get informationdirectly from the source. This canbedangerous at timesand mayrequire personal sacrificesbythe scientists that are often forgotten and not mentioned in scientific reports, thus making scientific results look simple and 'easy' to obtain. That this is often not the case is highlighted by our final essay-chapter. Gerhard von der Emde Joacbdm ~ogdans B G Kapoor
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