A stroll through the worlds of animals and men: A picture book of invisible worlds

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is little monograph does not claim to point the wayto a new science. it should be called a stroll into unfamiliar worlds; worlds strange us but known to other creatures, manifold and varied as the animals "h~·~~"'ves. The best time to set out on such an adventure is on a sunny The place, a flower-strewn meadow, humming with insects, fluttering butterflies. Here we may glimpse the worlds of the' lowly dwellers ;. the meadow. To do so, we must first blow, in fancy; soap, bubble ' each creature to represent its own world, filled with the percepwhich it alone knows. When we ourselves then step into One of bubbles, the familiar meadow is transformed. Many of its colorful disappear, others no longer belong together but appear in new ,,::,~".vU~"Ul-"V. A new world comes into being. Through the bubble we see .world of the burrowing worm, of the butterfly, or of the field mouse; world as it appears to the animals themselves, not as it appears to , This we may call the phenomenal world or the se(f~ll'orld oj the animal. some, these worlds are invisible. Many a zoologist and physiologist, to the doctrine that all living beings are mere machines, denies existence and thus boards up the gates to other worlds so that no ray of light shines forth from all the radiance that is shed over ' " But let us who are not committed to the machine',theory consider nature of machines. AIl Our useful devices, Our machines: only impleour acts. There are tools that help our senses, spectacles, telescopes, ....ione '" which we may call perceptual tools. There are also tools to effect our purposes, the machines of Our factories and of transrr.·","·V'., lathes and motor cars. These we may call effector tools. ow we might assume that an animal is nothing but a collection of






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published in Instinctive Behavior, trans. by Claire H, Schiller (ed.), 5-80. Madi. International Universities Press, 1957. Reprinted by permission the publisher.




0037-1998/92/0089-0319 $2.00 © Walt~r de Gruyter


J..-von Uexkidl The worlds of animals and men

perceptual and ..effector tools, connected by an integrating apparatus
which, though still amechanism, is yet fit to carryon the life 'functions. This is indeed .the position of all mechanistic, theorists, whether analogies are in terms of rigid mechanics or more plastic dynamics. brand animals as mere objects. The proponents of such, theories that, from the first, they have overlooked the most important thing, the subject which, uses the tools, perceives and functions with their aid. .... The mechanists have' pieced together the se~sory and ,motor,organs animals; Iike.somariy parts of a 'machine, ignoring their real' lInf'Tl,nn< of perceiving. and acting, and have even gone 011 to mechanize himself. According to the behaviorists, man's.own 'sensations and are.mere appearance, to be considered, if at all, only as disturbing But: we who .still hold that our sense organs serve our perceptions, " our motor organs our actions.see in 'animals aswell not only. the '11""'''''' cal-structure, but also the operator, who is built into their organs, as are-into pur bodies, We 'no longer regard animals as mere machines, as subjects whose essential' activity consists of perceiving and acting. thus unlock the gates that lead to other realms, for all that: a perceives becomes his perceptual world and all.thathe does, his world. Perceptual and effector worlds together,' form a closed unit, Umwelt. .These different worlds, .which are as -manifold as the themselves, present to natureIovers new landsof such wealth beauty that a walk through themis well worth while, even though unfold not to the physical but .only to the spiritual eye. So" reader, us.as we ramble through these worlds of wonder., ',' , Anyone who lives in the country and, roams through woods and with his dog has surely made the acquaintance of a tiny insect hanging-from the' branches of bushes.Turks for its prey.ibe it animal, "ready to hurl itself at its victim and gorge itself- with his until it swells to the size ofa pea (Fig. I)'. The tick, though not is ;till an unpleasantguest of mammals, including men, Recent tions have clarified many details.of its life story so that we are trace an almost' complete pictureof it. : " , From the egg there issues forth a small animal, not yet fully for it lacks a-pair of legs and sex 9rgans: In this' state it is already of ittacking,cold~blooded animals, such. as lizards, whom it it sits OQ the tip' of abladeof grass, After shedding its skin several it acquires the .missing organs, mates', and starts its hunt ,for blooded' animals, . , . " , After mating, the female climbs to the tip of atwig on some There she clings at such a height tha t she can drop upon small m that may under 'her, or be brushed' off by larger animals.

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The eyeless tick is directed to thi ' of her skin, The a r ~s watch~ower by a general photohighway woman by h pp caching prey IS revealed to the blind and er sense of smelI Th d ' emanates from the skin glands f II ' e 0 Or of butyric acid , loa mammals act the ti ' to eave her watchtower a d h I h ,s on e tick as she lands on something w~ ~ a erself downwards. If, in 'so this to her - she has re h d h fine sense of temperature I ac e er prey the b ' , t only remains for her to find hai I ' warm- looded into the skin of her prey and I a lv: ess spot. There she burrows" .: ' sow y pumps herself"fulI of warm . with artificial membran d fl ' proved that the tick lacks all esfan uids other than blood sense 0 taste Once th b h e will drink any fluid of the ri h . s e mem rane is , after the stimulus of butyric acid h g t, temperature, ,' cold, she has missed he as fundctlOned, the tiGk falls upon . r prey an must again clirrih to her


tick's abundant blood repast is also her I ', left for her to do but drop t hI ast meal. Now. there is , tick's life history rovides s 0 eart , ay he: :ggs and die, the heretofore cu~omary p~:~ot ~or Ithe validity of the ,biological' every living creature is an obiec~lOth°gIt ap!:,roach. To the physioloca. J a exists III his hum Id t h e organs of living thi d an WOr . He technician would examine a st;:n:n the. way they work together, hand, takes into account each' gd' n:dachllne. The b,lOlogist, on the f .' m IVl ua as a subie t 1"' .' o ItS own, of which it is the J C, rvmg II) a Uiu'alC:J' to a'machine but ani t center, ~t cannot, therefore, be . If we ask whether the t" kY' 0 the e?gmeer who operates the IC IS a machine or a ' o~ a SUbject, the physiologist will reply that h findoperator, a mere , ens receptors, that

and effectors'.1 the traits given operati I an . heat. In 0 smaller or larger clusters Th I .tick' S body has 'the 'a machine. M. . of external objects This m~a? Into units that become the __ .serve only. ':"subjectiv~ factor. t e sensatIOn' the objects: blue the sky. which admits only "'influences such as' butyric acid and warmth. eseharle the Cues by which-we g reen t e aw · '1 " . is fundamentally different from this. and screens out all ends with a muscle which moves an effector. such as cold. integrating device inthe central nervous system. ~electric currents. ~ . mmands only one receptor sign and and acting of the whole animal signs (Wirkzeichen) . ~. t fact. But since all of the traits of '. and has perceptual or receptor signs. it "not occur in objects'.n. a leg or proboscis. undeterred.~. which approach ~h es~ c us~ers correspond to groups e am a1 · other half of the brain cells is used b : In th.object WIth two arms of e cue or perceptual me~ning 'tehonh It Invests the object with .. ..' but with the . too are elementar at. and the clusters f fiis t e receptor organs' (Merk'its' e ff ector organs' (Wirk . each of which com t us be reduced to the cooperation effector sign.t-ndepend~nt of the spatial o tor signs of a group of p SSl 1 rty does.c'. as does any machine.~~.Impulse melodies act .n R.entirely physical waves of excitation (produced in the nerves by . No single part· of the . meet the 'everywhere. every animal rasns i . The clusters of receptor cell fill he ' c s answers to the of the brain. perception whatever the source of stimulation. whether ether waves.everywhere operators are at work'.ls~lated: If It were not possible for lc lar:~rt'f'rC. meanmg.. h .n . And the stimulus must be "perceived" by a subject. Motor cell. ~hlch repJ:esent Our specific of external objects and (Melkmal) which constitute the sensation 'blue' becomes th s:brvjeas t~e real basis of Our actions. .onal significance (Wirkmal). n. d' .0bJect are structurally' rona meanmg m t' ff .:: ". _wharever their activity . The 'physiologist . ·.' . .~. . Figure 2.c. The action of living .not merely machine parts. ) 0 e ector cells mak e up t he contents . f the receptor organ. Effector order to achieve an orderly collabor . . war . muscles of the effectors. The isotated imp 1 g ups acc~rdlng to their effector self'contained motor I'm I uses are Coordinated into units and' pu ses or rhyth .its effect only if it is swung back and forth in a certain : To all other' agents. alkalies. the biologist will counter. no engineer or~ngineers appear anywhere -process'. e units of infer ti h ' convey would also remain' " ma IOn w ich they to be fused into new units Wh.. visual sensory " duce the' same. organ. The receptor cells are bi organ.Ill. we may conclude that each living cell is an engineer who ~"T'''''''''''. e Impart t h e subje r .WI t e other an eff .-. . e greenness' of the law Th y. is. von' Uexkiill . ' as spatially separate units TPh .and the reflex arc is the foundation of all 'machines :O:ig. and so change the object . Since the time of ] ohannes · we know that a .e form of questions. that is.~' : acts. . In lVld~~1 cells of the perce tor or . E. Any machine part. x erna o· jects their tively speaking. Sensory cell.:{ ". E. . Impulse cells. 2). are ?rganized into well-articulated ro organ. '. with the transfer ofmotion. He finds no trace of ppenltor:.initiate the' nervous excitation and the motor cells elicit the motor impulse. gan. em nd the lb' t e d b y the separate muscles' .. . im s or other organs rmprmr Upon the e t I bi c~e Or functi. Receptor..'.:the stimulus... indeed outside the a' I' com ined . For all the cells of the : are concerned. 322 'r. as connecting links to transmit .' perceptual meaning through th bi us a ect those' e 0 ject. acids.. Simi ar process takes place in the effector .muscle responds' to all external agents in one · same way'by contraction. Johannes 'Muller-showed also that all _'. ector cue or' r"'l1.e. organs of action. receptor. ~he isolated effector or Impulses. electric · it responds as would any other piece of metal. --". cells (these.). or perceptive external stimuli. the orgamsm uses 'the 'receptor cells' in the stimulus-r ~ n:echamcs) and groups half of them . ":continue.influences affecting'. : entire reflex arc works by transfer of motion. not.-t '.: S. and is grouped i t I Y t e o~ganrsm as 'effector cells' n of the effectors Thes ? c usters WIth which' it controls the ld . '. M.. Ph CI c to It. 'sensory cells that.: . unite into perceptual cu:. connected by' . th ' e ueness of the sk . The manifQld perceiv·all the tiny cells.' To this the' biologist will reply.. ":receptor upon' external stimulation) to the.lOn. . (Merkzeichen) and The worlds of animals and men . It begins with a receptor. s. It transforms all external i . exist..outside the "~o. 323 which are s e 'fi ..ielicit a' sensation of light: Our. This 0 a~~. . I' " the muscles subordinated to th A mica ... suchas the clapper of a produces. Reflex arc.completely. : 'O~ the contrary. 'You mistake the character of organism . and effector With ~h ps ItS. actions of the tick reflex' in' character .rand responds to it with the same "resulting in contraction. 'we. 'We can show that all the. III fact..' the same effective stimulus..~. the optic nerve. All Our hu~an se:sr~~ectlOn ~f sensory impressions is' signs. " ' .t ecelVlng part of the bra' . sense organs.c.ons.

'itself.q .~ .. the extent that It . only three become stimuli.objects fO. whose blood she needs before she can bear her young. Why these three and no others? we are dealing with is not an exchange of forces between two but the relations between a living subject and its object.'effector cue or meaning. which are valid for all animals. e . ' linked by a connecting cou~~erstructu~:c't ~re best shown by the The relations between S~bJect.: " '. are fitt~~. . falling on the hairs of the mammal.object .earer . . iects the effector cue of shock onto them. Out of 'whole environment.' ualities that can serve " possess certam . corresponding specific effector Cues.l: theory: a ~:~ wo~lds with equal COl' nnietenes ' complex. 'We are concerned with the chemical stimulus of butyric acid. to constitute a systematic .'ctionai 'cycle Umwelt. . Each elicited by objectively demonstrable physical or chemical stimuli.slder th. mewo-i. ~ ti (the nature of which we do not .' b s on the other. "I f butyric' acid releases 'specific receptor ' cycle~ SInce the stimu us 0 these receptor signs are' p:ojected :.. . ! _' ~ '. and . namely.: cue ~eal~m~ich the receptors let through.IOf' lis which . certainty of her actions. of the f~nct~onal. . Figure .. . now. The tick. . tern. and serve as to lead her unerringly to her goal. tick hangs motionless on the tip of a branch in a forest clearing.' F.at a SUbJ~: we shall gain insight into the .~ 0 ."..particrpates in 'as 'perceptual cue-bearers ' " .'as an olfactory cu~. And these 'perceptual prescribe the course of her actions so rigidly that she is only able to .' The . .324 .1 :_ . faCt~: :~I~st~h~~~. We are admittedly confronted here with three successive reflexes.guishes t~e ricepto. ' ' functiona cyc es hb rs of perceptual meanmg m " '( h ammal are t e eare. or the temperature of the skin.flustrates' how the subject .consisting. This in turn releases a tactile which extinguishes the olfactory stimulus of the butyric acid.G._. something quite wonderful happens. mto u~~ple animal a well-articulated . Out of the vast world which surrounds the three stimuli shine forth from the dark like beacons. !)~noi~e~. furnish the .~cl~ (Fig. which starts the boring response. -The receptor cue elicits running about. . . sUl'?I~ t e s~~e 0 . ' position gives her the chance to drop on a passing mammal.her Umnwelt. h ' bjects with effectot cues. simple world co~respon s to a Sl .3.. the tick. . To accomplish this. By l~ uc ion . . \S tid ·. " ' d b.th~ aid of their ' : andarrang~ment of r~cep f the Umwelt with receptor 'cues. ' . . signs. We are concerned solely with the fact that. and " Besidetheselection 0 s~~: I iables the effectors to function in arrangement of mu~~l~s w IC en. is related fa the same or to If we further c?n.' principle'. These on an altogether different plane. f an action is the ~. no stimulus affects her until a mammal U"". or~a~. the : And nowlet us ~et mto bi t It shows at a glance that : subject.by means of their ' " number and arrangement 0 euector ce ..". .. r: :: a complex on~.' J. of.and the mammal aSh er 0. . object 'are dove~al~ed into on.:. field Receptor Perceptual-cue bearer (releaser) . and security is more important than the example of the tick we can deduce the basic structural traits' . which she can use as sign stimuli..uc. the tick's receptor. This is best expresse . out. her body with its receptors and effectors. But' who is content with this statement and assumes that it solves the' proves only that he has not grasped the basic question.' r processes that take place in the receptor organ initiate corresponding. Umwe. Motor field whole rich world around the tick shrinks and changes into a scanty' in essence. of three receptor cues and three effector" ... Central :.n.ue.' ~ecw' ll-planned succession.'. and these ' one hand and as functional c. However. i' '. ' . objects by severalfunctional Il a " als f~om the simplest to the .. the tickpossesses . von Uex~ull The worlds of animals and men ' 325 ' . and in a definite sequence. ' signs. only three become the bearers of receptor cues for the tick... themost dec+slve.r~i~h .ways.'. until it in turn is replaced by the 'on' of heat.. '" th che~a of the functional cycle. But the very poverty of this world guarantees the. Of all the influences emanate from the mammal's body. has been given three' signs. in the effector organ. any more than the mechanical stimulus (released by the hairs). the action only to'. and these impulses induce the tick to.: glands?. between the receptor sign subject and the stimulus from the Object.. Functional Effector cue bearer effector . let with her legs and drop. I I follow eac ot her m . of " hundreds of stimuli radiating from the qualities of the mammal's y. iefly as: The . Perceptual Central receptor. l'Uuu.

'." we move our limbs freely with our eyes shut. meaning . which frames ~ll happening. ~~e se~ that the ~"bJ~\6:~rstime. e~e d i ection with the doctrine of Kant.I:7~~t:en~.-. the slgn~ of butyric acid arouses her. ~:yv~~:~ ~fc~: :t~~~~\~~v~e~~~~e wI itho~t f~?tdthmeuzsotobl~ga!~~d. a living su ject. . he icks the raisins out of a cake. " Nor do we ask how butyric acid smells or tastes to the tick. h II e that the same IS true of space. he tick's world. we register the fact that butyric acid. because they have specIa. . too. .. because it is biologically mean.r . remar k'"ble faculty ".~/. but in part also contradic't ther.i >. ar~ interested solely in the fact t~alt~~elral. stimuli . .p"'p'~n""'". : ti With this " . . .. . d 0 time is made up of a series of moments. WIthin w . ~tabl~ thirig in co~tr. sequence. the next chapter we s a . In some way. whereas the tion of butyric acid is of eminent importance to her. .. h For the '..: .it intends to . ur 1 . pt. -. every subject spins his relations to n characters of the things around him. not of' butyric is accented." '" with a space that is valid for all beings.ears is beyond the realm of a..e! ~~~!~~~ii%::~~~sbad been starving fOT:ighteenhr~a~~~~ewe ·r e 3 A tick can wait eighteen' years. von Uexkidl . Inste~d o~ . in 'his world. look for the perceptual Cues. Th~t ISso met g. <. tick IS III a s .sm..temporal plane as our own relations with the objects in our human .~~fhiS o!n world. many years a . . . to iznal function agal!l until .ast to.tIV.J. these are Iy fixed and. What have we game .326 .. n the dr~wback sufficiently to. i . b' th can be neither space nor lme.. his World.a most . . t~~ere can be no .explolt' in the Umwe t eo . t the world stan s sti . instead of . and weaves them into a firm which carries his existence.' · u y~ t d in knowing what taste sensations the.ht less' state of the sort that interrupts time f~r . This gives rise to the conviction that there is only one space and one time for all things. '. The raisin stimulus leaves the tick quite cold.now have to say . which all living creatures are pigeonholed. a or ' activity. the ability to But whatever ~umber Irf. 0 P ... another . 'Space in the Umwelt. .. hich 'the world' shows no c ange. at~:.: d by realizing this? Something extremely .' ::. they always take effect outside tever the relations between a subject and the objects in his environthe subject. robability of a prey K .se. the spider spins its threads. d '11 Man's moment last 1/18 of a .!e S. Umwelt of any animal that we wish to investigate is only section out of the environment which we see spread around it _ and this ronment is nothing but our Own human world. Only recently have physicists begun to doubt the existence.'l(. .. n. the stimulus of raisins. ~a~sms ~.. • ' " are easily deluded into assuming that the relationship between a subject and the objects in his world exists on the same spatial . units. .'.. The worlds of animals and men 327 these -worlds.:.. it to say that in the tick's receptor organ there must be receptor which send out their signs. 'a' t rnomen .' u to be the only Time. on the other hand.nce w I I Nor does the large n i~e tick sits abV.lOU~Y ::~~:: ticks ambushed III t. becomes a receptor cue for her. .to the tick. which interpenetrate and complement. without a living subject. since they follow each other in a certain. .. time. u . 10 ogicai . d " hi ch brings a mammal under the twig on .. as hereto~ore. as we assume that the gourmet's receptor send out theirs. t~~. which affords a further' insight . . The tick's receptor signs give the stimulus of butyric a meaning in her OWn world. the tick has Likea gourmet VI. there can be no livingsubject.. exist is evident from the fact that all men live in three distinct . I h f a moment vanes In differen We shall see later th~t the ~ngt 0 de t for the tick. t.:n~:rt~r. . That 'such a space . ~ever-c~an~~~h:~:fore assume that during her period ~Iltty.Of the s.~" .~~r:~~~~:.eep i . On y In t e :' t a time and does not h stops for. tandmg s 'our case. In the gourworld. has ultimately esta?lI~he Its conn I th ry by stressing the decisi . In.. 'b t ': c acid alone from among . . task of research is to identify each animal's perceptual Cues among all stimuli in its environment and to build up the animal's specific world' them. : . """h~. d · to an unusua egree. and that is wherewe. by transforming it into a perceptual' as the gourmet's receptor signs give the raisin stimulus a..' mere o~s': . The first. This fallacy is fed by a belief in the existence of a single world. the things in her environment.. "rl'PI"~I. see~~~~~ ~ of its contents. are' time-bound as well. we know the exact and extent of these motions. '..ubject. The lucky COl~Cl ~.:e ~:. or cannot 0". therefore. Our hands trace paths in a space i" .

i( . However.ed'the a. the static plane's ' firm 'scaffolding that: ensures order in functi~nal. which is the basis.iat right angles to the forehead.space. up and down.. ina tic"arch as. his operational space with a t: framework for his directional .of each step perfectly: through kinesthetic sensationsi direction signs. the f coordinate system.. I' :1·" . 0/ man "J to t~ry' e:~lanes of oper~tional space.. 5). this boundary is at 'eye level. By one's hand horizontally and moving . The semicircular canals . height of the upper lip. head. A}. right and ieft.' the tip of the nose. It.·1 "I I i .cannot give the effector space any solidity. most people. forward' and backward. This relationship so dearly merous expenments that we can k . and moving it right left With' eyes . These steps evidently do not provide :a very precise .rthe boundary between the two becomes 0 coincides approximately with the' median plane of thebody. others indicate the .by . · . B'y holding handvertically.ssert]lon: all space. It is found by holding up' one's hand forward and moving it~back and forth at the side of the.trying to bring together the forefingers of · hands with his eyes' closed. ImenSlOna opera- is 'i Ii •-. .anything up to 2 em.. . It fs imperative for anyonewho deals with the problem of space become aware of thi~ fact. possessing the three canals also have a th~:e. whose position roughly 4. coordinate system composed 'of these three planes and firmly with his head (Fig. Into the' changing throng of directional steps which.can easily be ascertained. It is of t~e utmostimportance to us that paths once traced are very easily.closed. for the space in which they are taken. Cyon's great contribution is.. though many-people at the. This is called kinesthesia.the border plane.. The coordinate system 0/ man j . Anyone may easily find out : inaccurate they are . We distinguish six directions. of all spatial definitions. since we the direction . · . Every . This is what makes writing iITthe dark possible. called se$ci~ctihir canals' (Fig. . " . . ". which we shall call directional steps. contains and is ruled system of planes 'placed perpendicularly to one another.':. thus providing. as motor . that he traced the three-dimensional acter of ourspace a sense.I. Comprehensive experiments have shown that the shortest steps we can measure with the index finger of an outstretched 'arm are of : 2cm. the boundarybetween above and below ." 328 J. or three pairs of ._ 5. Nothing couldbe simpler.it up and down in front of the -.organ situated in the middle ear. The boundarybetween in front and shows' the 'greatest variation. effector 'space is 'not merely a motor space built up thousand Intersecting directional steps. 9r ooseffector space. We measure all these by infinitesimal units. people indicate this plane near the ear opening.steps.. von Uexkiill: The worlds of animals and men 329 callelour-moiorsphere. .norrnal person carries around with. .:« word that explains' nothing. He will' find that the attempt usually and they missone 'another' by . and by some it is even placed in .

li>. n e zones of ebb and flood tid Th specimens Use their hard shells to sea b '. This is further :~by their-internal structure. ere are. since the: entrance must be · even if its aspect has changed. Without these.· Determination of the front door by visual visual space 'is insufficient in most cases.. antennaewerecut off immediately flew toward the hive in its new '. The ability to find their front door in a purely operational : also' be demonstrated in insects and molluscs. spawning grounq. WhjCh... I es.~.not as a compass that always points to the North. o on the rocky ground betw:mgth t e snail.. n so domg.shows Even more striking is the 'ho ' .emicircular canals of a fish." then: the animal must be back atits starting point whenev : reduced the 'nervous markings to zero as it moves about. Figure 6. the ele~entary structural unit is a stationary ~~ne' the no t a motor magmtude is di . . space. bees 'gather in the air. they return to their beds I . It is apparently destined to · a compass . . be it a nesting . . · whole body-are analyzed and-marked accordingto three . in visual space.flown out. animal. that'theorgan hasan added significance beyond projecting the. . This' . bees antennae must the :a~u~e~e :':~heg~~a~e~o:~t:~~{O~ the front.. In normal 1'£ th .. although these ". . I~ will then be found that the. As SOon as the the same route The . doo~..-: There is nodoubt that a compass for the front door . where the flight hole -.l. unlikely cen the existence of a com ~ue'h he ?nly alternative left is' to . If ill the movements. : canals. h Patella . f ha~ do visual ImpreSSIOns. Locus also owes its'existence jec ive receptor SIgn. · Figure'S shows the s.have no semicircular canals .t7"a Wi a space. it tubular system in which liquid . it was found that bees '.'''''uvw s hie. their front door .is a implement for ail animals with a fixed home.' bi ti '. "spot. . . ' a compass for thefish's own 'front door'. · The following is a very convincing experiment.pass rn t e animal's operatiomil we d 0 not know Its nature.: under nervouscontrol in thethreespatial directions. The semicircular canals of a fish' . It IS not a configuration inherentin surround- _' . e spend th ti fb op a ed out of the rock Here e nne a e b. Themotion : liquid faithfully reflects the movements of the whole body. to wander and graze Over the rocks around them e iey recedes. 7). ""L1Vl.'This mean's that their orientation is mainly operational only e:::e~~:.of these experiments.was previously : Notuntil five minuteslater do the bees turn and fly toward the In' a later stage . . they orient th . The worlds of animals and men . It is obvious that ~must be of paramount importance to the..' J.. A beehive is " a location of 2 meters from its originalsite while most of the . It IS equally.mselves ~y o~tJcallmpres~. von Uexkidl . ~s IS IrectlOn..' 330. . . 331 antennae. Parella (Fig. · planes into the animal's effector space. they do not always alone the snail ~ould are so primit~ve that with their t it could be guided by a s t ts home. At high tid' th . pressed hard onto the rock. .

The skin surfacesthat release the same local sign in us' 'touched.e smallest units and are therefore able to differentiate the number of places. .:. lattice. of the eye.' objects receding from the eye become .size.'. until they' finally . of places ("ith orientational units. Visual and tactile loci coincide in their' Only animals with eyes are visual andtactile space 'gated.they have their tactile hairs...projected into the outside world. . . ' animals. . ' : When fingering an object.': also have -receptor signs for location. cup grasped WIth the outstretched '. These are called local signs. " ~ .\ 1 333 ing substances. .'owhig tothe-global structure. . When we finger an object. . .'lie dose together. up grow... is the subject to the.!. The worlds oj animals and 'men . .h~l1 obtaI~ a progrC?ssively . . T?e delicacy of this .' tactile ~pace is of prime importance. The otace-mosau '~bjeo. i· :. And . . Each element of vision has a . and then re-enlae in i re. since there is local sign for each visual .' ~ tactile space enter into conflict A er.thIS IS where visual and guided to the lips gro . like the feeling hand . Each of them is in a place:' If they are now moved down the subject's 'back without the distance between them.evermore comprehensive parts of the environment are covered 'place. until they shrink to a single locus'. von Uexkiill .:::. the place-mosaics in th .'. the more details will be cruder than it does to the h e eyes of a fiy must appear consider-.'. the ':. ~ ". things in his Umwelt. and both serve to give it form. which represents a._". in addition to the receptor sign of tactile sensation.:" . which explores the oral cavity. . In consequence. to render " diminishing a picture more -and m~s 0 vanous ani~aI. I and the world as seen throUgh~h ce-mosaic. ua e ements WhICh grasp the same ' .number of visual elements va ' .: .:slt.. ' tac. Figures 9a to 9v~ een reproduced as water colors. have a photosensitive skin: . ff/. . 'place in the Umwelt. ' . between the pla~e mos~ 0 f ma~es it possible. the 'two are clearly distinguished. of the environment. . retinaof the eye. on the nape of a subject's neck at a distance of more i em. :. roving eye. furnishes a place or tactilespace.Hi. they come closer and closer together tactile space.' ' local sign'.oes not change its observer does not see the c pace predommates. AIJ nocturnal . the section of the "tha t 'reaches one 'visual element grows with" increasing distance..coincicie. photograp~mg It again with mosaic.nes ~eatly III the eyes' 'of \n:::sP<}fi<lm£>lv . ..es Over all the things in the sub" sp:eads a subtle mosaic of places : depends on the number of"vis ~~ct world.elements. our exploring fingers its surface with a 'delicate mosaic of sensory units. .ts in 'an animal's world. :. .the places or: loci are combined jiirectional units. because an '.smaller. "have identical skin 'regions to produce 10c~1 signs for both . both in tactile and visual space. the place or locus represents are no differences. '. .c. according to their importance in Next toothe 'tip of thetongue. like the tick. ' \ \ ·l~: :. The farthest plane The visual space of an insect inflight "~l . Since the lattice Ph~t:gr~ ~e . FigureS illustrates the visual space of a flying insect It is . Th e coarser the la err enVIronments mus t diff er . If the points of a are placed. vary. the. .all cave dwellers live primarily in tactile space.'. and does not exist in his. at which point they in a . . th ' ' e smallest spatial vessel within which ·:"'t :j . the contain th. In this case tactile s ' .and tactile stimuli. were made by the lattice method. objects do not grow small .1 ~ . ': . so-long as'. coarser mosaic-images ha b p e WIth the picture is disturbthe lattice.tile space. This has been proved by Weber.:.. ronment. • .'."Eyeless animals who. t '.exceedingly in. ~ : ~"j'. " In 'many animals. uman eye ' any Image can be transformed into a l' ' fine mesh or lattice on it this m th dP ace-mosaIC by superimpos. In the.332 J. ws III VIsual space but d III tactile space. Rats remain quite' unhampered in their motions' even if they have 'vision. It follows that. very small elementary sections. eyes.Visual space .

lsua space' of snails.335 Fig~re 9b. . .!\)¥. Figure 9c corresponds - to the ix:hage~furnishedby the eyI. as in tactile space. Figure 9d c p spins a we~ that remains regIstered by a mollusc eye It h orresponds approxlmately to the . contams nothing beyond b .. the threads of a cobweb. ... :vonUexkilll The worlds oj animals and m~n .of an animal if we of visual elements in·its eye. photographed through-a "creel! The same village street. er of light and dark areas. connec~i~un.334 J. ~o~~letely. ns etween places are established !..'. It is easy to see a world :which contains so few details....) . seen by a mollusc the: number Th~y en·able.' m~lslble to its prey. sows that the vi I .. The sa~ villagestreet.us:to gain insight into the world . and we may say: the sider ' . of a' housefly. . ..

the adult's visual space" too. the cars that. whichis the counterpart of the photosensitive Iens of.of distance signs. which we call the horizon or the farthest plane.past me did not become more remote. . step by. "I The farthest . andnoticed some workmen on the gallery.relaxed. ten meters to infinity.fartherobjects.signs appear which signal 'backwards'. .' 1. ~ . onto the retina. but also the hand which guides the di ' needle.of: the farthest-plane is not rigidly fixed. When I took my . reports that as a little boy he was passing the ':garrison church. some .. find that not .first walk of doors after a serious case of typhoid." this orbit.operational and tactile space. .. whichsurounds visible things. . ~The difference' between the visual spaces of a child and a portrayed Figure which. the eye is set for a range of from. were no nearer and.of the little dolls begins. like a colorful on which alf visible things were depicted.. do we learn ~o push backt .it sets a limit to. the human eye is elastic and can be bent by special ciliary This curving has the same effect as focusing the lens in a camera 'Whe'n the .336 J: von_Uexkull steps.of ten meters. 'forward' appear. When preparing :an object under a reducing by directional which acts by condensing a large number of places onto a small area. in · He " ' io. The loca ' . at a ' Skm. w~ recognize the things. only larger and smaller ones. When the muscles ' . When -the' elastic lens distends the relaxing muscles.farthest plane that encompasses world-Only gradually. until. Beyond the twenty meters. .of the . .' environment as being-near or.drove.plane 'Unlike.distant plane with the aid .i)"'"'J'U' Helmholtz. he asked his mother to_ reach down. to focus the objects before it.1 . :. moon and stars wander: without any difference in' depth on same mostdistant plane. " . visual space is surrounded by impenetrable wall.Throughthe muscular movements.step. the farthest plane hung berore me at' a -distance of about twenty meters.lens muscles' are contracted.The visual space enos here with a. take's' much _smaller direction steps corresponding to the that have been 'drawrr close together.th~ h photographic camera-s+ namely.' uman eye has the same function as .-Sun. smaller .only the eye. and the" all . reproduces an experience Ut. pbjects at first become only larger or. only as soon as theyreached thefarthest plane: _ ~ :The lens. far within a radius .

which co ·visual space . Accordingly..fr'om which a sectionis . because they are built f ~s. hlCh III tersect each other space independent of subje~r °If subJe~tlve perceptual signs. P: p. Then we sh. be: they beetles.fly [scliemu! ic ): dissected on the rigln (according a 10 Hesse) · Cor: Chitincornea. . II ' I men In t eir Individual soap bubbl "ct a so see all. The worlds of animals and men 339 'half a meter frQID them. .all re ' '. our thly. which give its space a solid framework.xl<:~i' universal space. · ': Whether" the farthest 'plane encloses visual space in this or manner _·it is always there. or the cows" [2. the chandelier would vanish 'c~rtain distance. which encloses each of us Pll II we recogl1lze the soap . ASSUIIi.. bubbles. There' !. This behavior is like that of a yachtsman . and then 'close by or under it-again. is anxious to stay within sight of an island . "But other observations suggest that the most remote plane also 'in other ways in the housefly'S world . . which must t~e image projected by their lenses at" varying depths. which represent a lier photographed. flies. (b): K: nucleus. Nf: nerve fiber. their abruptly whenever they have flown half a meter or so away. butterflies.' mosquitoes or that people-a meadow'.lng that the optic apparatus made up of the fly's "elemeritsfunctions as' a portrait lens. ' :v •• tt • .the '. .Sz: visual cell TW9 omriiatidia in the meadows .338 1. von' Uexkiill '. rhabdom. Structure of the compound .eye of (a): Whok: ey~. compare· Figures 12 and' 13. :the squirrels leaping from branch to branch. hei as we . it would seem' justifiable to that their .around us. . enclosed within soap bubbles.' Now the-eye of 'a fly JFig.en this fact is clearly gras ed ~ha .farthest plane is at this distance. Kr: crystalline lens: Krz: lens cell. and in each there exist the' directional operational space. but interrupt. As an this phenomenon.ll.and contain all that is visible to them.. We know that flies do 'not circle around a hanging lamp' or chandelier. without.'. (: . The birds.and thus cause the fly to return. w~ do :e still cling to tJ~e fiction of an faCIlItates mutual com .erl: relinula. depending on distance of-the object· seen. 11) is built in 'such a way that its ' -elements (rhabdorns) are long nerve configurations.. the eye. which define the' mam permanently surrollnd~d by their hen j hi ir own space W . Exner has surmised that we might here 'dealing with a 'substitute for the muscular lens apparatus of. We may therefore picture all. Each soap harbors different loci. Chandelier as seen by man 13. 0 only because this conventional mUUlcatlOn.erlco'n. Chandelier as seen by fly Figure 11. and 'with a portrait lens.·Rh.

A third.time is the product subject. borrowed from carried by water.other hand. the blossoming of a ~ower~ can be brought within the range . moments per second As I In the snail's world OCCur m .' he studied . . '.c i 1· j. .340 1. An Instance of time contraction.infer from this th r times. The snail's movement: .ith the collaboration of another. who ~~edon fast. lI 1 . N: rod.n.time. Nor do its. slides und:rai~ :nall IS placed on a rubber ball . ' · The human ear does not discriminate eighteen air vibrations in· second. . .by which more eighteen pictures are taken per 'second. On the. there are animals whose. :von Uexkiill : Receptor ti~e . which wou'ld v:~ishe:~~ . · It has b~en found that eighteen taps applied to· the skin 'within second are felt as ·even··press\lfe . of-the fighting fish to its own mirror image. if the rot I... . . 14. Liter.1/18 of a. ovements. In this way. As already stated. u-. it acted as. III Its crawling its f?ot.I simple Umwetteu and time are of no imme di . WI urn away but if f are a dmlllistered per second. E: eccentric wheel. if ~th black-and white sectors was turned slowly. . a . 1/50 second.. food: After this training. The does riot recognize-its own' reflection if it is' shown him eighteen per second. -. to .~s become a tempo of three to four at the snail s receptor time moves nr"""PC~D~ . In onary. b ./ IOnspeed of the black and white tain at a certain speed and s'ooenathvOl reactions became more uncer. we . S: snail U". .LUJ""<U1L only when numerous receptor cues. "".sthen moved up with the stick each second it '11t snail is given one to three . . since all sensations are accomplishedu the same moment sign..~~e:~g~al had become gray . for It is the expression of an indivisible elementary the so-called moment sign. late use to the subject Th . A moment is the smallest time vessel.-. This is a technique .h f . all motor. Furthermore.'.H'UJ<. ase of slow-motion photog. 'bufhears them as one sound. . perception.' Ka~l Ernst ~on' Baer has made 'it clear that . . remains in the same 1 ' . if wewish to' observe motions too swift for the human eye. :Tlle sirigle pictures then follow each other in jerks of 1/18' second. If'a "process is . .lfou e~n to climb onto the stick. it is condensed into a short of.second.. and in 'whose motor 'processes are' consequently enacted more slowly or more than in ours. ' ereat ter they sh' ft d ' did not happen until the black se I e to the opposite. snar t an ours do to us..appear at reduced speed. ·.foll~wed each other within proves conclusively that in t:~ ~~~ld-:.. . ball.' .. perceptual consists of -shorter or longer moments thanours. the time speeds them up.is iven i .. student trained the fighting fish to snap toward food if a gray 'disc was ·rotated behind it. We may." Time as a succession of moments varies from' one' another.snail's world a rod that oscili~te.ft 'II besi : Our or more .. . according to the number of moments experienced by subjectswithin the same span oftime. A vine ~ n I.held in place by a bracket Th th It?Out fricnon.' The worlds of animals and men 3'41 . us e snail unhamp di .. The snail's shell . all motor processes _ as in the c sh. " for in this case the fish received a li h '.The question arises ·whether. l~g . ere. span of and processes too swift for our human time-tempo (of 18 per such as the wing-beat of birds and insects. ~er second h.' .'the tate of 1/18 second. Mot-or processes are thus extended oyer a longer.. " . t shock when they approached was gradually increased th . above-mentioned work.slow-motion photography.Flgure 14.: .. '.a resu t. processes too slow for our human tempo.photographed once an hour and presented at.tile reaction.As -~6tion photography slows' motor processes down. and then projected at a tempo.rlt mustbe presented to the fighting fish at least thirty p~r second. '. can be made visible'. the duration of a moment amounts to . motions seem slower to the il th aster than m ours. the identical forall sense modalities. : 'The first' experiments of this kind were made by a young :·scientist. !. the snail will climb upPo~~/i/t~:all ~tl~k i. · Kinematography projects environmental motions onto a screen at accustomed tempo. At this speed the bi k ~ors .

The small animal comes to rest only when it reaches its food. ' perceptual-cue. El1~iro~mellt'. Of all the different things m Its enviro a1ly on ItS longitu rna axis. which alone of all things in its world do not emit stimuli. the 'animal ' its legs. eight beil-shaped organs located on the periphery of the umbrella (represented symbolically in Fig. lashing drives "t' SW1. ed with dense. the 0 .' . medusae have a mobile mouth (manubrium) with a muscular system its own.''''Yy~. . '. " d however the Paramecium . driving the sea water in and out ' through fine pores. which draws 'the.. are also able to get along with a single fUnctional cycle: Here the. urchins. which its Umwelt takes in only the ever-ide!1~lcal:e~~r~~~a~~~ impeis it to i . and breathing are carried out by the same rhythmic contraction of the muscles on the edge of the umbrella. At the same time the stomach . . The ever-constant pulsation 'keeps the . entire organism consists of a swimming pump mechanism. there IS . Thus there are "«"'~u.. without central To iIlustrate the contrast between animals built in this way and animals. Sea urchins possess a large number of such reflexpe~sons.' a reflex animal. again elicits the same effector cue ad infinitum. for the same reflex runs all the time from each bell to muscular band at the umbrella's edge. .::~w:r~C of this kind.. as in Rhizostoma. say. von Uexkfi/T. we can speak. and ejects it again. filt~red: The sale manifestation of life consists in the rhythmic up-and-down ' swinging of the elastic umbrella.ari. The stimulus thus produced elicits the next umbrella-beat. d~'l . carry a large number of spines -which. distends and Contracts alternately. To ensure continuity of this motion. and dominates the rhythm life. . the bacteria of putrefaction.'. . long as these 'reflex arcs remain mutually independent. when a sea urchin runs. The liquid content of the stomach is propelled through labYrinthine digestive canals. I have coined the phrase: when a dog runs. the legs move the anima]. the term may even. By defIec~~~~lw?ereuPov~edt ~en~ay. with a ' single functional cycle.~stb~~ . In very siQ1P17umwe~~nforh~~. These facts show how nature is able to fashion life according to her plan. 'In the medusa's the same bell signal rings all the time.fi' Umwelt of tire Paramecium of which performs its Own reflex function by itself. I The worlds of animals. whose clappers strike a nerve end at each beat.''''''' which have tentacles with self-contained reflex arcs. . !: A few multicellular animals. which is Connected to the receptors on the umbrella's edge.. and this releases the same receptor cue. arcs operate quite independently and are not directed by An external organ that contains a Complete reflex arc is aptiy termed reflex person.~ .that in this case the same receptom ~ IS remo. Moreover. rows of h ide by sid. ~ riated. through the water '. ~he~ever'f~~cape The same obstacle-cue always elicits the same ' motr?n 0 . ' .no ne nt andthe Umwelt of the Paramecium ' 'In Figure 1$. such as the marine medusa Rhizostoma. ~ . . whose walls absorb the nourishment and '. Figure 15. and men 343 : CUeis always extinguished by the same effector cue. . I " . SWimming.the accompanying oxygen. feeding. In this way the medusa gives her own effector cue. All other stimuli are cut off. plankton-filled sea water into itself Unfiltered.animal floating on the surface of the ocean. extended to animals with several reflex arcs.' " b ckward motion.eParameclum IS cover sown.. followed by a Of reactl~n. like porcupines. such as other: medusae" . I I' . while it revolves conunus iftly hose S1 ~ .342 J. the envlronme. In the case of a single functional cycle. Moreover.. ~ U"'J. these reflex central organ. . ~ut t~e temporal and spatial framework of the ~mhw~~~:::. ThiS' consists h a'~:l again begins to swim forward. 16).

and poison claws or pedicellariae) scattered over their surface.:ne :~ edusa with periphe~a/ organs . We think in terms of the sea urchin projecting the receptor cue of outward into space.long.. avoided only by the appearance of the. For the sea urchin. a substance secreted by the skin. it remains ineffective. Furt ermore. . snapping claws. sea urchins respond to any darkening of the horizon by a .. if it consists of numerous reflex persons. absolute domestic .rnAn< of spines which.isince each """. the spines part.. have acornreceptor organ. " The tick's life manifestations. instead of using these isolated reflex arcs... which normally seize every approaching object. the tender tube-feet are never attacked by the sharp snapping fangs. stimulus and heat stimulus. For pain inhibits the pain· eliciting action. chemical stimulus emitted by the sea urchin's enemy. namely. ' . a ship. _ . which has no superior: : center. as we have seen.A . = of claws (cleaning Claws... a fish. · receptor signal of pain in the central organ. since it has no visual space."·.. Even our representation of its Umwelt is not sufficiently simplified. despite . 'de enden t reflex persons. . 16. This peace is not dictated · by a central organization. ri "'* The worlds of animals andmen 345 344 j. where our sharp teeth are a constant danger to the tongue. are composed of graduated pressure stimuli and chemical stimuli. Although some of these reflex persons act in unison. A reflex republic such as a sea urchin may well have numerous receptor in its Umwelt. is displayed toward a cloud. : Fi~re : ~rr. : .'Wi'jIlMf'WttYwwmWflitt:"ri . ho~ever. :. Besides . and the real enemy. tain sea urchins have four tu¥-fe~ for climbing._. Thus in response to one and the same.s:e!~: a~~e to turn a forest of spears against any be~nng a. ' . the starfish.It is achieved by the ·presence of autodermin. even though consisting only of butyric acid stimulus."" cycle operates in utter seclusion. cer .the utter independence of each reflex person.. . For p . clapping claws. they work quite independently of each other. this possibility does not exist.": 'throughout the skin in such great dilution that. domestic peace must be secured differently . As soon as two skin surfaces however.emitting object that approaches e . b th skin there are delicate.~~s :ta~ed to the lime shell by means potn~ed·. Undiluted. the poison fangs spring forth 'in their stead and bury themselves in the enemy's suction feet. of snapping by the pedicellariae. it becomes effectual and prevents release of the normal. as in our case. What actually .. Its receptor cues. eace reigns. But these trecentor cues must remain completely cut off from eachother. autodermin lames the receptors of the reflex persons. totally isolated magnitudes. : . '. iqn Uexkull ~-- --. In the tick's world the prey may therefore possibly as an entity.I. upon contact of the skin 'with a foreign body. as shown in Figures 17a and b.! . '.' . . We may therefore refer to a reflex republic in which. It is diffused. Even this represents a superior type of organism.LV'. consist substantially of reflexes."''''vu''' cycles.. In a sea-urchin's reflex republic. since the ~"'1J. . deve~~p. .

it would still be impossible to join these loci.346 J. In the light of further observation. ..1:he ~ed'urr:hbi's envi~onmenr :: .a urchin's Umwelt . we generally assume jhat the '·shape of an object is the receptor cue originally given. of a wad of cotton 347 passing Form and motion as perceptual cues . Even supposing that each receptor Cue of the various reflex persons is ' provided with a local sign and that thus each is in a separate'place in the sea urchin's world. is adapted to the moving form only. ~.. Jackdaw and grasshopper Figure ~7a. 'However. Now. but that owing to the intersecting blades of grass it cannot recognize the grasshQPper as an ' tity. the form only separates from the Interfering subsidiary images at the time of the jump. form and moving form two mutually quite independent receptor cues. quently this Umwelt must of necessity lack the receptor cues of form and .. . but motion may even appear as an independent receptor cue. " Form and motion appear only in higher perceptual worlds. without form. Figure 18 shows the jackdaw on the hunt for grasshoppers. a secondary receptor cue. The jackdaw does not snap' after 'it it hops. Couse. . however. Not only are static . we may assume that the w does not know the form of the stationary grasshopper at 'all.' 'I i' . and that motion ' · is added only as an attendant phenomenon. '. The se. motion. as we have difficulty in finding a familiar form in a puzzle picture. -. ing to this hypothesis. It simply . This would explain. One is at first inclined to suppose that the jackdaw is familiar with the form of the grasshopper in repose. ' ' ' . the 'death- ----(b) i -. Figure 17b. von Uexkiill The worlds of animals and men happens is better conveyed by the analogy lightly Over its photosensitive skin. for many animal worlds this is not true. 'thanks to experiences in Our own world. !.cannor see a sitting grasshopper. which presuppose a combination of several places.

however scallop responds by pushing.!." cue of the female in flight. scallop '~y flight from the vicinity of its enemy.. Pine on 'the other. which plays a decisive I~ Its effector world. a slight motion of the stick the pea is swung back and forth in . .' . It was possible..". On the other they roll up easily . affects the 'scallop not at The starfish's characterisiic form is! sensory cue for the scallop.-. Most leaves spread out if tries to pullthem into a narrow tube petiole foremost.._ ... However. . .. . many will invariably throw themselves on the pea. which act as .are sitting.·to show ~hat small sticks dipped in gelatine were pulled into the earthworms" i :'J'~ ~ . I '.. The whole process represents a uu. Asterias. Darwin early pointed out that . not sef for either form or color but solely for a certain . In repose it is' never taken for a female.:. .out its long tentacles. The eyes are. .. ' must intervene.. 'The pea is covered with a sticky gum.~handle both leaves and pine needles according to their shapes (Fig. serve it both for protection and for food. Experiments have shown that shape and color of a' moving ooiecua wholly indifferent. I have built a. The Fliescircling around a chandelier. The worlds of animals and men' 349 all. not-their tip. The swinging pea deceptively imitates the ~P.'fly rod' consisting "Ofa small stick from which a suspended on a thin thread. hand'.and offer no resistance if seized at the tip. If .. Environment and Umwelt of the scallop It was inferred from the earthworm's spontaneously c~r~~ct 'handling . which' corresponds exactly to that of its enemy. this does not characterize the enemy precisely enough. .. ! may wei! conclude from this that static female and female in two different receptor cues. They approach the starfish and receive the new stimulus. motion'is as slow as that of the starfish. are males at females flitting by. .' organs. So long as the starfish is at .. environment. must exist as a receptor cue in Us perceptual Th!s assumpti?n ha~ been proven false. Figure is provesthat motion without form can feature as a nercentu cue.. ~ r:"» ~ --. .. within range of its hundred eyes.r' ----'\ '. too. .' . nuptial flight. The existence of a receptor cue for form was long surmised in Umwel(of'the earthworm. First. '. tempo.'. The earthworm 'drags leaves and pine needles into its narrow cave.. . These are always males...tbe starfish.a sunny window sill on which a number of flies. As soon as the starfish moves.'" 348· J.. arid some will remain to it.jo elicit the' second functional cycle. must be grasped at base.~. is the' scallop's dreadedenemy. and needles that the form of these objects. then by 'pretending dead' they ·disappear entirely from their enemy's sensory world cannot even be found if sought. .t!i. If their static form does not the perceptual world of the pursuing enemy. conclusively extinguishes the enemy's receptor cues. and by this effector' . 19. a scent. which always fall in pairs. ~on Uexkall feigning' reaction of many insects. .: II 'I' :..' if they are to be dragged into a narrow hole with' J . upon the scallop rises and swims away. It presents the scallop in its environment and in its Umwelt. which removes' .The object appears as a perceptual cue in the world only-if its.

If putourselvesinthe bee's place and 100k at the field from the o 21.'evidently been adopted for the reason that the receptor organs" earthworms are built too simply -to fashion sensory cues of shape. in which blossoming flowers alternate with buds. ' . it became even more urgent to answer the question: are. their form and the b d ge to stars or crosses according 11OlogJ'lcal '. whereas' they avoid compact forms. the other with from its base.eto overcome difficulties which to seem utterly Insurmountable. earthworms had to' abandoned. the blossoms are chan d ' '. they. . the earthworms differentiated between. anmg or them.350 J. Taste discrimination in earthworms holes indiscriminately by either end. example shows how nature is all.I.. they' do between the tip and base of the leaf' Although earthworms handle leaves in keeping with their form. the two ends of stick exactly' as. It has been possible to show bees alight by 'preference on figures that exhibit broken forms.' Figure 20. But as soon as one end was with powder from the tip of a dried cherry leaf. von Uexkiill The worlds of animals and men 351 " . . This has. ~Figure 21. buds do not.' which' was designed on this basis. This question has since been solved. stars and crosses. Thus the hypothesis of form' perception in. guided not by the shape but by the taste of theleaves. . u ~ assume the unbroken shape of circles 'The bloss~~~~~:~e~. such as circles squares. the lowest animals in whose Umwelt we may expect to find form a perceptual 'cue? . ~~!en:: di~covfiered quality in bees is evi·dent. The bee is seen in 'its envrron blooming field. Environment and Umwelt of the honey bee o of its Umweit. contrasts a environment with its Umwelt. such.

no one will attribute goals to a sea urchin or an earthworm.scents. this assertion._~" i Now.same sensory cue. It is enough to assume that-the receptor cells for local signs in receptor organ are segregated into two groups. '. petty everyday worries into the life of the tick. does not matter from this These studies have reduced the 'problem of form to the simplest mula. pace."t. "receptor images' of a very general nature result. Excellent mvestigauonsreveat that in the case of bees these are filled with and . i. it makes difference whether the ... our. or the tick have no such schemata. ave .ofthese experiments .can accomplished by' coordinating the life manifestations of animals' the' viewpoint of a plan.. as we... .opposite kinds of action are governed by . has the opposite effect in their is striking how the two .: Goa_! and plan ... . By this expression we have already. Since we humans are . This is a fallacy which has led research astray again and True. '. swarms together on the white a er e ymg. ~~'''VjJU'UlJl':.sound to 'which the animals are adjusted . goal to the next.. owing to. quently there are' no true perceptual images in their worlds. sitting rn. In a neighborin room on. fiddling in lively f hi an d!ust~a~lOn .concerning the sound perception of night moths. by a pure plan of nature.m t rough the window and s~t next to it under the ~l!s S~nglemale heeded the female kind of physical or chemical ti mul a re was unable to ascertain Experiments made with h s mu us emanated from the paper. relations of meaning are only true signposts in our exploration of Umwelten. e plan artful microscopic structure of the ~r~: nura. 0 a eise. or butterfly has ever seen th Ion or . VUIl. lways the same. IS sitting m a room before a and pay not th~ sli ht~~~sex p~rtners gather ~nfront of a a glass bell. It' must therefore be our first objective to extinguish the will-a' wisp of a goal: in our comtemplation of Umwelten. a light coloring. those in one according ... The earthworm. since no . which preclude all doubt. which is . these moths are totally A fine observation by Fabre revealed h . According to information I· received .the effectis . ' . even in describing the life of the' tick. waylaying' prey. are easily visible.. von Uexkiill The worlds of animals and men 353 . . the reader must be given insight into' Umwelten."' outward. flyaway upon perceiving a while species which have protective coloration alight in resoorrsenc same tone. ¥. The .accustomed to carry our existence laboriously one.. be teleological actions.. There are no further differentiations. He then put males of this very rare species cam fl ~ape~.saw in the case of the tick. He placed an eyed hawk t ~hc~ntr~st between goal and . Night moths which. ve m this respect.: can be no question of discriminan . the schema 'broken'. which in turn are tailed into the over-all plan of nature. Actions directed 'toward a goal do not occur in any other all. .purposlveness. Figure 22 is' . we spoke of it as.een more specimen. SInce t e sounds she makes cannot or b ~~t.352 J. the scallop.' long ago.bIe wh~n we learn that for this one high tone of the bat 't g II~oth s hearing organ exists . I Grasshoppers be/ore a microphone . albeit unintentionally. hi e co or of Its own ski Th in t 1S mstance appears even m drni in."' . who fiddles in v!. those in the' other according to the schema . Whether the forms are physiologically more effective.. To prove. . This . we are convinced that animals live in' the fashion. ~unng the night. attenhbon to a specimen. sound manifestation of a bat or one produced by rubbing a glass' per :. '. grass oppers and crickets h b . where she moved her abdomen a mo ema e o~ a sheet white under a glass bell next to the sheet ofbout for so~e tune. If the schemata are . Perhaps later certain actions of the mammals may prove to. I.

But no who has ever observed a. is not produced. she had also 'hatche egg of. She hurried in the of the-black chick's peeping. which was her and blood. The auditory and the visual cues of the object 'elicited 'two contradictory functional cycles in her. she is not in the disturbed by the sight of him. No goal is pursued in either case.it is a necessary link in the chain of cycles which lead to mating.354 ' J. even if the chick is invisible. All right. as. we must give. This distress call mother hen run immediately in 'the direction of the sound with plumage. The hen shown in Figure 24 behaved even more oddly and Together with a clutch of eggs of a white stock. the mother hen is in no position to loosen a noose. I . Normally. Toward this chick.If a is tied to a peg by one leg. The optical image is ineffectual. the question of instinct hi h d~ s goal.should intervene at this point and activate the next functional Th~ nature of this second receptor cue must be investigated more in both cases: In any event. she begins to peck furiously at an imaginary antagonist. in the' world. T4e perceptual cue of peeping normally comes ' from an eneiny'whois attacking the chick. But the seemingly strange behavior of the males is easily explained if examined under the aspect of a governing plan. Very experiments have fully clarified this particular case. which determines:' their perceptual cues. " ' Both experiments show the same thing." Thus the partners make no advances whatever. the reader will say.. the proper effector cue. Does the acorn need an insti IC t oes no~ really get anyone anyme to grow mto an oak tree.by the effector cue of beak thrusts. In both cases a specifi:c receptor cue initiates a functional cycle. which chase • foe away. . but when she perceived it among the .she pecked away at it. insects. von Uexkid! The worlds of animals and 'men 355 be heard. as we have seen in the case of the tick. '. the existence of true goal actions. ones. They: are ruled directly by the plan of nature. Here' again. According to plan. this cue is extiQguished . since the normal object~is eliminated. but not-peeping chick is not a sensory cue would release a specific activity.her own black breed.mother hen hastening to the aid of the in a poultry 'yard can doubt. " image and effector image contrast nature's plan with the sub iect' . The struggling. up purposive action . if the fettered chick 'is set before the mother hen's eyes under a glass so that she can 'see ~t but not hear its distress call. It would be quite incongruous if it too. . goal action. figure 23 illustrates the resultsobtained in these experimen~&. it peeps loudly. another receptor cue. but. we will not have to . we have an interrupted chain 'of functional cycles.she behaved quite inconsistently.. " _I. or does . . chick's two sensory cues had not been fused into an entity. As soon as she catches sight the chick. which would be necessary to extinguish the firstperceptual cue.

since it """lrPLth neither matter nor force. the sea anemone rts mear~mg for the crab. Without plans. The processesillustrated in Figure 25 give a series of findings ' from studies of the hermit crab. To trace them is one of the most enthralling . the sea anemone's'. . This is enough to put the crab into dlf!erent n. S""Shell. they exemplify the plans of inanimate nature which they. which is expressed in the crao's. IS expressed m the acuon ?f the crab. and when physicists present Bohr's beautiful. IS robbed of its shell. Instin~t is merely a product of perplexity. . von Uexkiill The worlds of animals and men 357 .all.n Its she~L In the second case even its shell had been taken It. Image o~ the sea anemone assumes a 'defense tone'. These plans are because it is hard to conceive the nature of a plan. continue our stroll through the Umwelten. since no subjective ~oal is in .~ . Figure 24."feeding and the crab begins to devour it. even' level of arthropod WOrlds.tor Image of the sea anemone assumes a. ·:'f ~.. albert a futile one. But the finest hammer is not enough either. s prevailing mood.or. in either 'case. nature's 'designs will also be recognized as swaying the 'spmnmg cobweb or the building of a bird's nest. of a plan of nature. without the sovereign ordinances of there wouldbe no order in nature.case the crab had 'been robbed of the actinians which it c~rned o.356 J. models. F = Food ' As the figures show. T~e same crab and the same sea anemone are before us all the time in t~e first . the cuttle~he receptor.". the perceptual image furnished by the '.. We shall therefore not let ourselves be turned aside. a concept which IJlL". but will . In the first case.and blackchick J '\ 1 a host of b~ne-iorming cells work instinctively to form a bone? If ' deny this and substitute a plan of nature as a regulating ~act. that of the " crab. which plants the sea anemone Its sh~ll. According to these moods. if one bears a concrete example in mind.. that is. if we gainsay the superindividual plans of nature.' observations are of especial value because they show that. . If you no hammer.. object of a certain order of magnitude with a cylindrical to conical can assume meaning for it. " The sovereignty of nature's living plans is expressed most clearly • study of Umwelten. where the crab's shell the protectlv: mantle of actinians. It has been found that (he requires' an extremely simple spatial schema as a receptor . Every crystal is the. the recep. Then you hit your fingers. . '_ Hen . In the third case. If th~ same crab. Changes in significance of Actinia in'the Umwelt of the hermit crab P"" Protection. Ima~e gam~ a 'dwelling tone'. Yet it is not so difficult to gain an idea plan. to era wl into it. . . which repel its enemy.' The mostbeautiful plan will not drive a nail into the wall. only chaos. and ill the third case a crab that wore both shell and actinia bee~ left to starve for some time. one and the same cylindrical object _ in'this case" an~mone ~ changes its meaning in the crab's world aCcording t~ crab. if you ha plan and rely on chance.t0ods.

I. . too. . such as crates. of running around of borrng m. will consist few objects . such an Umwelt would leave nothing . "n. ' But the real problem. images. along with few r ''''''<1U U1i. !'. a tone depended entirely on the actions which it performed with them. Thus new' percepImages with new functional tones are created.to see sitting in a chair..' . ' . tones I~ OUr con~emplation of it. If an object is used in different ways. but all the more For orrentatIO~ is much easier among few objects than among If the ParameclU. I had taken a young. Elsewhere... be su~e. " If a dragonfly flits ~oward. the su determines which functional image will lend its tone to· the < . and jumped up on them. experiments were'carnefl on dogs. must be excluded. erWn . Then the chair was removed and command repeated. and he recognized them ladder. assumed a 'sitting tone'.'.. . none of which are given perceptually? In ' objects that we have learned to use.functIOnal.. The problems was stated very simply and the answers' of dogs were unequivocal.» sense organs can be supplemented and changed by a 'functional (effector image) dependent . the selective activity of the receptors these gates. For many of these 'dog chairs' at all suitable as human seating accommodations. To irpnT"V'" !hese Vital things. A dog had been trained to jump up on a . could do it easily. cert~mty IS the functIOnal tone connected with the stimuli.aning to perceptual images only by their 'effector ~one.which then lend different tones to the same perceptual' chair may occasionally be used as a weapon. before it at the command. VItal things l~ an animal's world furnish a perceptual image. we see the function which we with them as surely as we see their shape or color. be desired as far as certitude IS concerned..d me. 'j'\< r' I'" . The only thing which' he was a knowledge of European tools. It then a~sumes. its world. an animal's performances grows. .n. and to grasp the full significance of their Images. held a climbing' tone for him.' welt only a~quires its admirable surety for animals if we include The . '. If. which manifests itself as a 'thrashing tone'J strictly human case. result its world is indeed poorer. ~. i.. entails a readjustment to new impressions.. To clarify this remarkable state of affairs. This experience with the Negro indicates that we have effector Image for each of the' functions which we perform with the.. drinking in a...ere there are as yet no spatially articulated receptor as... it may possess several. we shall supply these images with a functional tone. the objects in the Umwelt of a foreign subject more tangible and gain new meaning. very intelligent and agile Negro with me the heart of Africa to Dar-es-Salaam.? had a functional image of its performance. this was a 'dog si and not a human sitting tone. For each new' .evel wh. a ~ranch to perch on it. ~]ays the lead. ' in our specific Umwelt. that III the the objects acquire a new quality. Let us . i" As the number Of.. through this it had meaning. This effector image we inevitably f~se so with the receptor image furnished by our sense organs.on the action which it elicits. can only' be analyzed! man.. we may say that the meaning of the only three ~n".. but is also distinguished by' a tone. As a..".. such as the sea urchin. number of functions it can carry out. in all its implications. functional image. at a I. Un: 1 1 '/1 .. It was' found that the dog now treated as chairs the objects on which it could sit. the perceptually given 'rods and .' ' I L i . von Uexkull The worlds of animals and men 359 image. F 358 J. climbing in a ladder.. but what lends this a~tivity 11n"r.~. We may say that the ber of objects which an animal can distinguish in its OWn world the.. To. we must constantly bear in mind that are the performances of animals..Oi . which marks it above all other branches. When I bid him climb a short he asked me: 'How am I to do that. '. may be observed especially in dogs. In t~e tick. junctional or effector tone.! . its world would consist of homogeneous objects. How do we manage . shelves.obstacle tone.that is able to gather experiences.. The receptor image of rods and holes had been supptementec the effector image of his own action. the branch not only as a receptor Image In Us world. ' If we wish to use the functional images to paint the Umwelten of o further removed from us. ! j. ''''''1>. It grows within the individual span of eve~y animal. All animals that operate on a reflex plan. The new meaning manifested itself as a new attribute.that populate Its Umwelt increases. the number or'.". Effector images can be surmised only in animals whose actions controlled by central effector organs. I see nothing but rods and As soon as another Negro had shown him how to climb the . To be sure.... Moreover.lt possesses few functional images. the mfluence of effector images reaches far down into the animal as shown by the hermit crab. .> stimuli that reach her from her prey stems from the functional (con~ect~d with the stimuli) of dropping down. which conveys their meaning and which we shall briefly term the junctional tone. stools.. It could also be that 'table' and' 'basket' assumed a special tone for the dog. ' Since functIOnal ~mages can be deduced from the easily recognizable ornllanlces of a~lmals. From then on. of t~e stimuli. projected into their Umweli and t~ey len.' this way: a series of other objects. who learn to handle certain "it. 'chair'. as in that of the hermit crab. . all of them bearing same .

the number of dog objects remains considerably smaller than that of objects. running. only feeding. fe~t are equipped wi~h taste organs.a stimulus to them. 27). whose stimulation releases a . The wall has an obstacle and the lamp a light tone. o~ the proboscis. 28). and light tones are Everything else displays an obstacle tone. . To illustrate' this fact. let us imagine a room in terms of functional tones connected with the objects in it.360 J. they around it. Finally. while all th o~~ctSu. Flies orient themselves with great ease in the environment of our As soon as a pot of hot coffee is set down on the table. which for them has a running tone. the functional tones of the objects in a room be represented by a sitting tone for a chair.rnducecontinued wandering. . and· crockery on the table. first by man (Fig. for the lamp whose significance has already been pointed out. They wanderthe surface of the table. von Uexkan human implements by turning them into canine implements. Here it is particularly easy to e y s mwelt out of its environment. and thirdly by a housefly (Fig. The room in terms of the junctional tones connected with its objects by" .\~ t· . terms of the junctionallOnes conn~cled with its objects by a dog ) II' J The ronm t . And The worlds oj animals and men 36] Th e room In . sitting. a revolving piano stool does not have a sitting tone for a dog. Owing to its smoothness. because its heat acts as. secondly by a dog (Fig. for the fly. everything assumes a single running tone. The floor has a walking tone while the bookcase a reading tone and the desk a writing tone. a meal tone for the and by further adequate effector tones for plates and glasses (eating drinking tone). they are held fast by their food. In the world of man. Ifwe represent the recurrent similar functional tones by identical in the dog's world.r e room In terms 0 thejuncti01lallones connected with its objects by a housefly Figure 26.