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98 9 4 8y i h y l evie ll i h e e ve

INVERTEB ATE LEARNINGAND M MO Y:

From Behavior to Molecules

h a J Car w a d Chr t L Sah

ep me f ych y d y Y e ve y New 06520

N OD ON

ic eth i t h ve p vided d ich t dit f t d ye d mem y c p b i ie f v ie y f i ve eb te

Sch e 2 , v F i ch 50, Li d e 60). I ece t ye , hte e h e d h bee e ewed bec e t h bec me c e thve eb te m c p vide p we f m de y em w

exp e the ce d m ec mech i m f e d h v ie y f d cip i e ch e eth y p ych

m de e b y h ve me ed c ce ted e devve teb te p ep t which the e c mbi ed pp che c

f c ed the e e theme f the ce b i f beh vi phi ve e y h c me be k w imp e y em "

t he t dy f e d mem yA imp e y em " pp ch i t the e c ve d m f vee e ch he e e t te y f ch pp ch y e c c t v ved p ic f m f e i d the pecify the p ec e c d t e f the ce ch e th t e h pp ch h bee cce f y pp ed mbe f ve ty tem he e c de eyeb i k c di the c Kim e

W dy e 83, W dy 8 ) c di c c diti i he pi e 1 80 1 8 ) d c diti i f he ict t i memb e e p

bb t h mp et 1 83, 8 , e et 1 82 vey et 1we ce y tem ce ebe m It 82) the

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A n n u .R e v .N e u r o s ci .1986 .9 :435 -487 .D o w nl o ad edf r o m w

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6 CA & S LEY

ew rd y ch dry h here i c rre y pidde el pi g c lid i i er eb e d i er ebr e re e rch,

ech ic l d c cep l le el c ce r i g he ge er l e

he cell l r mech i m f le r i g d mem ryhe p rp e f hi re iew i r ey rece pr gre m de i he fle r i g d mem ry i i er ebr e im l . F r re f b h f

d c r i le g h, we re r c he c pe f he re iew i wir bec e f he rece r d ce i he re we prim

c ider ly i ce fassoc a e ea gi i er ebr e S c dbec e he ce r l heme f he re iew ce er e r l mech i mle we ly c ider prep r i i which me f rm fa mo ecu a a a ys sh bee c rried , r i le i e fe ible. We

di ide he re iew i hree p r . We r pr ide brief er iew f m j r p ych l gic l pri ciple d p r di ed i he m der dy le r i g d mem ry We he c ider h w he e pri ciple dp r digm h e bee pplied i rie y f i er ebr e im l . Wc cl de wi h emp y he ize me f he c mm hemec erge ide h h e e erged fr he dy fle r i g d e r

v b at s

L N NG: P N PL ND P D G

Le r i g i i fere ce m de he b i f b er ed rel i hbe wee im l beh i r d i p e perie ce Speci c lly, lei pr d c f im l experie ce wi h e r m re i l e e

f he rel i hip be wee he e e e Re c rlll d 1 6 S hleye l 1 8 ) I m ki g c mp ri f le r i g d mem ry mi er ebr e d er ebr e i i p r ic l rly imp r

imil ri y f e e a ss he ki d f im i h di erepecie c le r b d he ype f beh i r m di ed by le r i g

re ec i e e l i ry hi rie h m y be ery di ere precl di g me i gf l c mp ri f he i li re p ded r hbeh i r l re p e e hibi ed by di ere pecie i le r i g experimI ere e rel i hip , h we er, re pecie depe de hpermi direc d me i f l c mp ri f le r i g cr di ere

pecie f im l .Le r i g p ych l gi dyi g er ebr e h e ide i ed m y

i ere e rel i hip h i e ce le r ed ch ge i beh(Re c rl ll d 6 ci h 1 ). Ne r e h l gi

e r bil gi dyi g i er ebr e h e l ked he er ebrle r i g li er re d h e d p ed m y p r digm w pr d

Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org

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R BRA LE N NG 437

le r g i ver ebr es se e ly is w p ssible e ermw e er f c i l s m l ri y e s s be wee ver ebr e i verle r g pr cesses ef re we r ssessme f i ver ebr

le r i g we brie y isc s me key e i i p r gms se im er s y f le r

Nona o a ve Lea n no assoc at a refer ch ge i beh vi r re l f

e perie ce wi h ei he i le im l w s im i h emp r lly el e e w m s c mm e mples f ss ci

le r i g re h bi se s iz i .a tuat o refers ecreme resp i g which is e se s ry p i r m

f i e) s res l f repe e prese s f si le s mS s t zat orefers i cre se i resp i e s im l s e r m re prese i s f er e se r i s s m l s.

o a ve Lea n nsso at a gimplies e le r i f rel i s ip be ee r

m re eve s i e e vir me , c be ivi e i w is i c yc ass ca co t o st um ta co t o o o a t a

l ssic l c i g is bes ill s r e by vl v's 2 ) e rly eme s gs A rm l g's re c i me p w er e c is im l s r S s s iv i he c i i e resp se) F llsever l prese i f e he c i i e im l s r S p reme p w er vl v f h he e by i elf c me elicic i i e resp se R) ic w s simil r e rm l resp se me p w er liv i r ce r lly, i c s ic l c i i i he US re pre e e he m l wi h epe e ce he imre p se. he m l le r s s ci i be wee he e w s im l

is ss c i c m ify m y be v s D mjk r 82cki s ).

I s r me l le r i g r per c i i i g, i c r , is respc ge . h s me s h he prese f he S is epe e

e im l e ibi i speci c be vi r l resp se s, e imle r s s ci i be wee e resp se i emi s e c se e

h resp e D mj rkh r 82 cki h ) he rse mple f s r me l le r w s h r ike's rep r 8 8) f le r i esc pe r m p zzle b . W e i i i lly pl ce i e b h re i c l y freei emselves. ever, f er m y ri l errse e ces hey le r e whic speci c resp se c i e resp se

ll we hem esc pe re f rceme )


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438 CAREW & SAHL Y

mpo a Pa nLe r cr c y epe e p he me erv ep r

he US he er m e v r S ) Depe c be e r e e re p e me re

me erv f r e r v r e fr m m ece. . her bb cmembr e re p e; G rmez 9 2) h r e ver ; G r

1966 G rc K e 966 Rev kyG rc 97 )

F O e Con on nhe e mp e f c c r me c fr mh r ke re pec ve y e cr be b ve re r f

c F r r er c mpe h he e r e he rec p r f he CS US c c c

re p e re f rceme pe cSe on O e Con on n

he e fe re f ec r er c h mc c me pr ce c e re p e w h ever be p rew h he e c r 98 ) p r m, w re

S2) F r e mp e f er c r c re be w(S ) h ck (US) he e c be p re w h ew m ,

S2). F w h r , he m w h w ce h (S2) eve h he h h ever bee p re w h h

B o k nA c c e per meby K m r ce phe me c ek ng

h emph ze he mp r ce f pre c ve re h p f e r K m 969) K m f h m

m c e m re w US e S S2-Ue e y we c e re p e b S S2

m rece ve pr r r y e e eme ( ) f he cm , hey f e h w e r he ec e em (S2

repe e re e f he c mp p re w h he U (S *h , he e C (S2) f e c re c eve h

bee repe e y p re w h he US he pr r r S bc S2 h e h whe US we pre c epre e ce f C w h wh c be prev y p re , h erfere w h rb ck c her CS K m 969)

Con n en v Con u I er em r f pre c ve re h p , e c rv r e e p e c ve v e f he S by y em c y er


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wh ch US occ e ce wan ingen po S occ e ce. h wa do e yg v g d e e gro p o a mal he ame m e o S-US pa r

y g v g o e gro p e ra m er o US pre e a o . h all gr

had he ame amo o S-USn igui ypa r g ), d er g amoo S US n ingen yco ela o ). he po ha whe he US wa l kely o occ r he a e ce o he S a p e e ce, he S o

edi ed he US, eve ho gh may have had a ro g emporal rela ow h . Re corla o d ha a he p ed c ve val e o he S dec ead d a l y o evoke co d o ed e po d g.

Con one nh ono d o ed h o he e l o a pec al k d o pela o h p r de cr ed y avlov 19 7) a d mo e ece e ded y Re co la 1969) a mal ca ac ally lea ha a S p ehe a e ce o he US Whe h occ , h o y lea g ak.

oced ally co d o ed h o ca e o a ed y eve al me al ma p la o o e ample p or o S-US pa r g a m

e g ve e pl c ly pa ed S-US p e e a o h a ma a ega ve co ela o e wee S a d US e dow g he S w h o y p ope e eq e lea g a k

Ove ha o nhe al e ce o each eleme a compo d m l g ea ly e ceamo o lea g o ha m l Whe wo m l a e p e ecompo d he pa c la m l ha mo e o gly pe ce vea mal w ll e d ove had w he weaker m l , .e. le co d ow ll occ o he weake m l avlov 19 7). Re co laWag e

Re co la 1969, Re co laWag e 197 ) gge ha he epa a e mw h a compo d compe e or a ed amo o a oc a ve e g

he mo al e m l ge g he la ge amo . Mack o hha each m l compe e o a e o 1974)

Con one mo ona Re pon e C Rhe ER proced re wa rod ced a a d rec mea re o

co d o ed ea y E eSk e 1941) A emo o al e po e e ho gh o ecome a oc a ed w h m l ha a e pa ed w

h whe he co d o ed m l ay a l gh ) pa red w h heq e pre e a o o he l gh w ll make he a mal ecome

he co d o ed ea ca have d ama c e ec o o go g ehav o .e ample, p e e a o ohe l gh ca d p appe vely mo va e

ehav o ch a eed g o a -p e g o ood a d coac l a e aver vely mo va ed ehav or ch a e cape or avo d


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r spons s us r is mo iv ion lly onsis n r spons o li i ing ondi ion d s imulus or r vi s M in os 19 4).

pr ding ri ov rvi o l rning prin ipl s nd p r digms

illus r s nim ls r p l o orming v ri y o sso in v n s o urring on ir orld n s ions ollmin y s p r digms v n ppli d o s udy o l

nd m mory in num r o inv r r n m ls. n s ion d s ri vior l p rim n s nd n dis uss llul r nd, r l v n mol ul s udi s

A IS M LLIF RA

rn ng in on y s s n o s rv d sin Aris o ol , u sys minv s ig ion g n i l ssi s udi s o von ris 1921 1950dis in m ods v n us d o s udy l rning in s : rpro d r nd pro os is ns on p r digm.

F F yin oc duM z a s ag s a b s t m v a

p rim n l s ing s ion on i sp r l olors r pro dground gl ss dis s ndividu l s r pos d o o ompl m ns imulus olors, nd n ini i l s in pr r n s r ord d. s n r rd d on nonpr d olor S) y llo ing drin sug r r S) r s s l nd d on olor no sug r

s giv n n l nd d on o r olor). Wi is pro dM n l nd o or rs d mons r d s n l rn o ppronum r o olors v n p ir d i US Approviol r l rn d mos r dily nd o luis gr n l s r dily M196 196 M nz lr r 19 M nz l l 19 4). mpor l

ris i s r sp i lly impor n or in l rning olorr sso i ion M n l r por s is l o l rsso i ion i n in rs imulus in rv l o up o 2 3 s , i

p s n d o US Ho v n olo S is pr s nollo ing s l nding nd ding, olor is no sso i d i

US is rd ondi ionin is no iv ) Grossm nn 1M n l 1968, Op ng r 1931) is i onsis n i rli r ndings

l nding on on olor nd n ding on no r ill pr r l ndolor o ding olor Op ng r 1931).

n on r s o v r r s nd so o r inv r r s H ins1983 , rl yK rn 1984 S l y 1981 ), pr di iv v lu o

S do s no pp r o impor n or sso i iv l rning in x n is u s ion, ouvillon l 198 ) us d lo ing p r digm w g ups f s: O g up was s a d w h a s g d d


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R RA LE NING 441

s m l s S US, a d s s ly a d a ompo d sS S2-US (Blo g G o p) a s o d g o p as a d s

ompo d a d a s gl s m l s (Co ol G o p). T g ompa d fo ma o s o d o d

s m l s (S ) f d a l y of US mpo a os l o o o ld p l ss spo s o S2 Blo g G Co ol o p. T s as o o s v d. fa Co v llo a

o la v ly mo spo s o S2 Blo g G o p.W o o mo s m l a p s d og as a omp

s m l s a d a d a US, s o s o s asso a d o of o s m l s l m s. T s s m ds im s se e i n.Tsal of a l m a ompo d g a ly s l a

a l m ( . . o l m a ov s ado " a o s ps o ). s v al m s s s o ov s ado g Vo (192 ) fo d a a pos o o a odo a d a olo a om

s o ld s o d mo o a v ma d odo a o o ma olo . S m la ly s l a a d s m a o mo ly

odo s l va a olo s mpo a (Klos al19 ) ally odo also app a s o ov s ado olo s p s d odo - olo ompo ds f y g l a g p o(Co v llo a 9 3)

T o of l a g s app a s o a p as f(M l 9 9). T a s a a s gl a g al, o mm d a ly af l a g a d d l s 2 3 m

o aga as s 1 0 m follo g o d o g a d d ays ov 3 5 days. s gly, follo g o ly 3 Cpa gs app a s o o m mo y loss ov l f m of

). T a g als app a o p od ma mal l a g gs g v add o al a g s o o d s o

19 9). M l os la s s o a d lo g- m m mo y p as o afo p as a of o Co s s s m mo y s mo as ly f d d g a a s o asaf m mo y as om s a l B s p g l os o ( CS), ool g o+ C, o C o N a os s s 2 mfo\ o g o d on g s o a d am d as l a g om

s p g sam a m m follo g a96 , 9 6 M z l a 19 4)T a of fo ma o s d y s m m g o

as s v ly v s ga d (W 19 2, CollCa g 19 2Ca Coll 9 3, Go ld 9 5). om s s d s apa s a m m d mag s of la dma s o ampl( 9 5) fo d a s a l o l a o d s g s a vpa s a d d p ma ly o l s v ly spa al l


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the le e ts. e o ludes that bees a sto e owe patte s as a lowesolutio photog aph i thei b ai s.

Probosci Exten ionhe p obos is exte sio pa adig was o igi ally developed by Kuwaba a1957a d has e e tly bee used exte sively. I this p o edu e, o age

bees a e aught at the e t a e to t e hive a d ha essed i dividually is all tubes. A suga solutio the US p ese ted to the a te ae eli its a

e ex espo se i whi h the a te ae ove i a o wa d di e tio , the a dibles ope a d the p obos is is exte ded I a ol a to y sti ulus theCS su h as a owe odo is pai ed ith suga wate to the a t ae, beesqui kly o e t ial develop a o ditio ed espo se to the odo that was

pai ed with the suga wate US E be1980 198 ,Va es hi1971,MasuhMe el19 2Me el et al19 4). olo -suga wa e asso iatio s a elea ed o e slowly Kuwaba a19 7Masuh Me zel197

his pa adig has bee ve y use ul o studyi g highe o de lea i gphe o e a i the bee. Fo exa ple, it e a a d his ol eag es havede o st ated o ditio ed i hibitio itte a et al1983,Couvill

t 0 ha ees ha had e li i lu pai ed p es tatio s o the odo a d suga wate exte ded theip obos ises whe pla ed i the t ai i g situatio with o odo p ese t a d

the et a ted the whe the odo was p ese ted. his is i o t ast to beesthat had expe ie ed odo suga wate pai i gs A als t ai ed withpai ed CS US p ese tatio s exte d thei p obos ises whe the odo isp ese ted a d ot whe it is abse t Mo eove , bees show a d a ati

eta datio o lea i g to the o e l u pai ed CS whe it is subseque tlypai ed with suga wate . hese two di gs ta e togethe p ovid goodevide e o o ditio ed i hibitio i the bee additio , Me el

i e a( 98 ha e show ha he va ables tha p odu e se o d-o deo ditio i g i ve teb ates also esult i se o d o de o ditio i g i th

bee.I additio to p o os is e te sio bees espo d to suga wate sti ulip ese ted to the a te ae by ovi g he a te ae towa d the d op o sugawat . his espo s a also be o itio ed. A ve odo -suga watpai i gsthe be espo ds with a te a ove e ts di e ted to the odo

E be198 ikewise, pai i gs o a ovi g st ip d patte with suga tothe a te ae esult i o ditio ed te al ove e ts to the ovi gpatte E be S hildbe ge1980

eu a S udie of Bee Lea inghe bee b ai sup aesophageal ga glio is app oxi ately1 a do ai s about900 000eu o s i tho1967 he idb ai o ushoo bodies o the b ai have bee thought o a ato i al easo s to be


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NVERTEB TE E N NG

mport t or l r g th Mo s 198 ). Mor ov r ol gt l lob s d th mushroom od s shortly r l r g pp rs o m mory orm t o r r t 1980 M l 1984). Th mushr

bod s r v put rom bo h th v su l d ol tory t rs thd h v xtr m ly high d sity o b rs d sy pti o t o s Alt 1919 Mobbs 198 ) I tr llul r r ord g rom uro s i th

r s qu t d ult us d v du l uro s r v ry sm ll ll od s r l tr y r mot rom s t s o uro l t v ty g r ord g rom xo s or d dr t s Axo d m t rs r typ lly

th m Erb r 198 ).D spit th s di ult s o l ss o i t r uro s lo t d th

p rt o th b r i h s id ti d th t shows r spo s h g s duri

l r g h s uro s r h r t r d y th r mult mod l r sph y r spo d oth to odors d to su r w r pr s t d to t tErb r 1978 980 198 1) Dur g l r i g th sug r w t r r spo s

d h uro s o ly r spo d o h odor CS h odor r spo s s s l v how v r th s lls o ti u to r spo d o m y odors E1978 1981). I t r st gly th s uro s r also r spo s v to mop tt r s d ollow g t l st mul t o with sug r w t r r spo siv ss to g v mov g p tt r s s g tly lt r d 1981).

AX AX/M S

h g rd slugL max max mus s g r l d h r ivor whospr rr d d s m d up o oods su h s rrot pot to d mushroG lp r 1975) rst show d h t o sumpt o o o o th s oods

dr m i lly d r s d si gl tr l by p ri g di g to th with CO ox os s wo l ss s o st mul dom h d g b hL max ol tory d gust tory. Slugs r ttr t d y odor o lo omotow rd pot t l ood sour s h th pot t l ood sour is lo t

max v rt ts ps to t st th ood tt r t st s ou t r d ths t d. A s r s o p r m ts h s d mo str t d th sso

l r g rly od y s ug s p r or p rti u r odmpl S hl y t l 1981 ) ou d th t slugs qu ly l to

s l tiv ly odors p r d with th bitt r t st o qu di sul t wh g ho tw th odor p ir d w th th qu d th ir orm l

odor) omp r d both to v slugs d to slugs xp r g th odor t st pl tly u p r d r l t o sh p A m s o h o thgroups show d o d r s th r pr r or th odor h l r

v rs o w s r t d or pprox m t ly 3 d ys d th m x mum CSi t rv l w s pproxim t ly 60 s S hl y t 1981 ).

As dis uss d ov CS s to om sso t d w th US th



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ust be te po ally elated to t e S a d it is t oug t t at t e S ustalso p edi t US o u e e Ka i 1969, Res o a 1968 Wag e 19

xpe i e ts i s ugs ave assessed t e i po ta e o p edi tability by

usi g p o edu es ode ed a e Ka i s 1969) b o ki g des g Sah ea 19 1 b) Ka i obse ved that p o o d t o i g to o e e e e t So pou d sti ulus S S blo ked o p eve ted o ditio i g to t e

se o d e e e t S . slugs p io o dit o i g to a ot odo d a ati ally edu ed o blo ked lea i g to potato odo S ) w e a

o pou d p ese tat o o a ot d potato odo was pai ed w th aqui i e US. I o t ast s ugs w thout p io t ai g to a ot lea edavo d bot a ot odo a d potato do whe t e o pou d was pai edwit t e qui i e. additio eit e s ugs t at expe ie ed t e o pou d

S S -qui e pa g p e eded by ba kwa d pa i gs o qu idi e aa ot odo o slugs t at expe ie ed t e a ot odo -qui i e pai io lowed by potato odo -qui i e pai i gs showed a y evide e o blo i g

A l slugs showed o d t o g to t e potato odo . he te po a pai i g opotato odo a d qui e thus was ot a su e t o d t o fo asso ativ

ea i g Rat e t e p edi tive el tio s ip betwee t e odo a d tq s mp r a m as Kam a

Expe i e ts wit slugs a so i di ate that t e asso iative p o esses oma a be i ue ed by se o o de o ditio i g S ugs st p

se ted wit a ot-qui i e hase 1) d the g ve potato a ot paihase ) d a ati ally edu ed t i p e e e e o potato odo evethou h potato had eve d e t y bee pa ed w th the qu e Sahley e1981b) o dete i e whet e t is di g was t uly se o d-o de oditio g RizleyRes o la 19 t e pe o a e o t e slugs e eivi gse o d-o de t ai i g was o pa d to slugs i o t ol g oups otexpe ie g both pai i gs Slugs i he o t ol g oups showed o a

the p efe e e fo potato odoMa y a a s a e apable o usi g post-i gestive o seque es o eati g

a spe i ood to odi y t e exte t t w i h t e ood will be subseque t yg t P g t vues t at a e ost t i lude ut itio al de ie es Rozi 1969 ahleHa pe 197 ) a d gast o testi aldist ess Ga aKoelli g 966) W e a i als expe e e e the o t ese

o seque es t ey show a p ofou d ve s o to the spe ood p e edithe o seque e Ga iaKoel i g 1966 Rozi Kalat 197 ). It is

o o ly t ought t at the ood a e sio is a exa ple o asso iativelea g a o k 1977 Do au k a d 1982) Re e t expe i e tss ow that slugs use post- gest o al o seque es to odi y thei eed g

e a ey Gelpe 985) ou d t t slug wi sele tively avoid eat gdiet a e a s gle ea whe a esse tial a i o a id et io i e) ise oved f o the diet o t ast he e ova of a o esse t al a

a id ala i e) does ot p odu e a a ge i eedi g


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VE TEB ATE EA NG 5

F al y add t o to ood av da e lea gmax as bee s ow tobe apable o appetitive lea g. Slugs a be t a ed to app oa a odot at as bee pa ed w t e t e t e taste o a u tose solut o o aoppo tu ty to eed o a ow ext a t Sa ley et a 1 8 .

o study t e el ula eve ts u de ly g asso at ve lea g imaxGelpe i a d olleagues developed a se i ta t p epa at o o s st g

e eb al ga gl o bu al ga gl o a d l ps oGe pe 77Gelpe i et a 78 . Food ext a t applied to t e lips w ile e o di g obu al ga gl o e ve oots el ably el ted a oo d ated ytpatte o oto eu o a tivity eed g moto p og a t s as bede o st ated to be t e eu al o elate o eed g Gelpe et a 78 a dto o espo d to be av o i t e ta t a al Re goldGe pe 977

ai i g p o edu es si la to t ose o ig ally used by Gelpe i 1975t e i ta t slug ave bee su ess ully used to de o st ate asso at velea g t e sem a t p epa atio a gGelpe 1 80 a edp ese a o s o a oo s mu sC a a e as eo e ps o ep epa at o esul a apid a d sele t e supp ess o o t e eed gotop og am to t e taste pa ed w t t e US e eu o s ed at g tlea g appea to be e t ally at e t a pe p e ally lo ated slea g a be obta ed w e t ai g s do e o o e l p a d estdo e o t e ot e ulligaGelpe 8 .

Seve al e ve ells ave bee de ed t at appea to ave a ole tseque e o a t o s asso ated w t t e gestio o ood e ee oto p og am " ese i lude seve al o o eu o s w t t e buga glio Gelpe i Fo syt e 1 97 Gelpe 981 a d sal va y eu ow t i t e e eb al ga glio oGelpe 77 opela dGelpe i198 A se oto e gi te eu o t e meta e eb al gia t ell modulato y ue e o t e eed g moto p og am Gel e i 198Ret og ade ll g o t e o e t ves om t e bu al ga glia to t e e ebga gl o eveals a lus e o about 1 eu o s ese eu o s a e o sp te est be ause si ila eu o s ave bee ide ti ed P eu an aeasee below d ave a o a d-l ke e e t o eed g G llette e a 198

iela d Gelpe 8 suggest t at o a d like eu o s imaxmay be dopam e g s e bat appl ed dopam e t gge s t e eemoto p og am a dose-depe de a e .

Alt oug asso at ve lea g as bee demo st a ed L max bott e w ole a al a d i t e solated e vous syste t e e a e seve al

mpo a t d e e es betwee t e two p epa at o s I t e w ole a aodo taste asso at o od es a appet t ve be av o lo o otio

o t ast e sem - ta t epa at o a tasteaste asso at o mod eso su mato y be av o eed g). esults om seve al expe e ts sugest t at bot t e w ole-a mal a d t e solated-b a lea g volvs m la eu al eve ts GelpeCulliga 198 a d subseque tly



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6 W S LEY

u ige e pe i 98 ) ep th f wi g f t e-quip i i g i the wh e i whi h e u i - v i e e

upp e i f he fee i g m p g m i evi e i he i e b iti ppetitive iti i g e u e p u e b th i e pp h e p e the pe i p i e wi h ug i u e fee i g e p e t the e S h ey e 1 8 . hi u h t - e i i ify the f y gui e ef we the e-gui e u m y e p e.

L UROBRANCHA A CALIFORNICA

he m i e m uP e an ha ae p f with h

e i i p b i e e i f we by bi e- ike e p he im fu y ex e i p b t i e the f with ti Mpit D vi 1 7 ) we he e ibe i ti hi beh vi by i ive e i g. hey f u h imexpe ie e p i e f p e e ti he S) g e e t i

the US) w u qui k y e e e p i g the f S A h t h expe ie e th the S he S ut i exp i it y u p i

e i hip i h w qui v i e t he f MpiD vi 1 7 I e expe i e the i i g p e u e we e

m e i i v i e e i g p igm. A im we e give e iti i g t i i i g f p e e i f thei u u p i e with e e i h k t he vei i we e

by i te i i e v f h If the i e v i e he f he h k w mi e e i g w i i e by v i e f he f Mpi i 1 75 Mpit e 78

i ive f we e e e wi h exp i i y u p i e g up A i i e e p i i y u p i e g up we

y e f v i e h im i the p i e g up. Si e

g up we e i e i ex ept f he p p i i g f he S Uf v i e e p e w ue e e i i be wee the h k. e e i g y the p i e e i e e ge i e iAv i e e p e f im exp ie i g p i e S-US p e e

i t ex ee h e expe ie i g S US p e e i u1 h f wi g i i g i i Gi et e e 1 84 h

h w th t the pe f e f hi e i g t k i epe e m iv i te f he i

he i u u pe i i y f he e e v i e h bee e

D vi e gue D vi e 74 1 80 . hey f u hi i e v i qui h m ge e h we ge e iz i v i e beh vi e e e vi e 74 ) M e ve

e eivi g i e e i i i i g {tw i e e S u te b


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13/53

R BRA L AR G 4a ross groups of a i a s wi h o o e s i u us paired wi h he US)avoided o he spe i food ha was paired wi h he US a d showed o ha ge respo s ve ess o he u pa red s u us If o asso a ve f

o ribu ed o he ear ed ha ge behavior, o e wou d expe hefa ors o a e ea h odor s ar Dav s e a 1980).he eura ir ui u der i g feedi g has bee ex e sive hara er

he ora vei w h fused a era e a es a d he rh ophores are he he ose sor orga s i vo ved i food de e io Ma eraDavis 198Dav s Ma era 198 ) Se sor re ep ors are o a ed w h he se sep he a a d are hough o be respo sib e for de e o of food s

B ker e a 198 a b Axo s fro hese re ep ors pro e o he ea d rhi ophore ga g ia whi h are o a ed per phera a he base of

e a es a d rhi ophores respe ive Bi ker 198 a) A pop aabou 100 euro s has bee ide i ed wi hi ea h periphera ga g io hapro e s o he e ra erebrop eura ga g io or brai hese periphe

euro s appear o have he ose sory, e ha ose sor , or b odarespo se hara er s s B ker 198 a)

he e ra o ro of feed g has bee a a ed o siderab e deDavis e a 19 19 4a b 1984 Gi e eDavis 19 Gi e e e a 1980

198 a Kova e a 198 1983a b S eg er 19 S eg er e a 1 e ra ga g a appear o be vo ved feed g he erebrop eura abu a ga g ia he per phera ga g ia are d re o e ed o erebrop eu a ga g a a d he bu a ga g o s a so d re a

he erebrop eura ga g ia v a he erebropeda o e ives Ma euvo ved eedi g have bee hara er ed P e a aeaWe

ou dis ussio o euro s ha appear o have a ro e i ear i g A group ofeedi g o a d euro s " he para erebra euro s C s G e

a 19 8 198 ) have bee s udied i aive a d o di io ed a i aor a he C s are ex i ed b food o he ps hu gr a a s

o ras ra ed a i a s food s i u a o produ es a e sive baof i hibi or pos s ap i po e i s SPs) i he PC s his sa ha ge reased i hibi o ) o urs i sa a ed a i a s as we DaGi e e 19 8 Davise a 198 A us er of i er euro s heI sre e ve dire he ose sor ex a o a d ur a e ex as apses o he re ra or o o euro s a d h bi or s apses o hpro e or o o euro s he I - s appear o be he ajor sour e ofI S s o he para erebra euro s Kova e a 198 a b) As a resuei her o di io i g or food sa ia io , he I - s respo d o food s i uwi h a pro o ged barrage of a io po e ia s, resu i g i suppressio feedi g o do Gi e e 98 ) Cha ges i he I- s appear o re e age era de rease feed g o iva o ra her ha a spe e ha sasso a ve ear g DavisG e e 19 8 Dav s e a 198 Kova e1985) I eres i g whe he ervous s s e is re oved fro we - ra


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4 4 8 C W S L

a imals the euro s sti l showbarrage o S s i respo se to oodstimu i deli ered to the atta hed ora ei howe er u like i the semii ta t a ima s rom brai s re o ed rom ai e o trol or satiated

a ima s showed i disti guishab eex a yrespo ses to ood stimu i.resumab y surgi a iso atio o the NS e o ed the i hibito y sig aarisi g rom satiatio but e t i ta t the i hibitory sig als arisi g rom

o ditio i g This preparatio th s o ers o siderable promise orstudyi g ellu ar me ha isms o ea i g that a be disti guished rommo e ge era ha ges due to a ter d moti atio a state.

LYMNA A STAGNALIS

Re e tly the po d s aiym aea st g a shas bee show to be apableo appetiti e lassi a o ditio g Audesirk et a 198 ou d tLymnaea are ab e to ear a asso iatio betwee a eutra hemostimu us

S amy a etate a d a appetiti e US a mixture o su rose a d aseidigest . A ter 1 trai i g tria s s ai s hat experie ed paired prese tatioo the S a d US showed sig i a tly more eedi g mo eme ts to the S

a s a s a xp r c m r xp cu pa re presetatio s o the S a d US or pre e tatio s o the S or US a o eMoreo er s ai s that ha e experie ed spa ed trai i g e trials per day

or three o se uti e days T!= 90 mi showed greater rete tio o thelear ed asso iatio tha s ai s that ad experie ed massed trai i g t als15 rials o a si g e day T45 mi . teresti gly i o trast to other

mollus s Sahley et al 1981 Haw i s et a 198ym aea showedba kward as we l as orward o d ti i g ba kward o ditio i g e e tswere shorter li ed( days ompare to orward o ditio i g e e ts6days A exa der et a 198 exte ded hese studies i a exami atio o e e ts o agea d moti atio o the lear i g. T ey reported that you gs ai s aged 8 wk ear ed the tas i both ood depri ed a d sated

o ditio s whi e older s ai s aged w ear ed t e tas o y u der odepri ed o ditio sThere ha e bee o reports o e lular hages t at might u derly the

lear ed ha ge i beha ior Howe er the euro a ir uitry mediati g t eeedi g respo se has bee des ribed i o siderab e detail e jami

Rose 1979 1980 M roha e jami 1980 Rose e ami 19791981a husym aea holds ear promise or uture europhysiologi astudies o appetiti e ear i g

AC A INA FULICA

The la d s ailA a na ashows a dramati ha ge i pre ere e oood odors as a result o e perie e ro lhase 1977 1980) ou d hat


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15/53

NV RT RAT ARN N 49as a s l f s c d f d g a p c c f d b vs a ls s bs ly p f d d f f d y ad f dc pa s a v l f d d . p f c was d ca

p f al a wa d d d c as d la yl a d ap d l wa d p d wa d a v l d . a ls a w p s d d b f d d d s w a p f c f d . v c as

f d d p va a c d l a g f b v l s a d adl a g was ly l g-l v d a 48- f d g p sl a dp f c was bs v d p 3 w . add ag f

c d ac s f l a g. a g ad l s aa l g f d g p c w f d a d d a g

T s as ym aea( xa d a 8 ) b va al v l aca d la ac s a d f s f f appl a g.

HIRUD MEDICINALIS

d c al l ci ud as ly c ly b f c s f l a gs d s. T b av c v g s p al as g w c ca b gg d by l g

s l s. T s v d c f ass c a v l a g l c s was by H d s g (19 ) w f d a a als p cpa d p s a s f a p s l s ( ) a d a l ss w d a a d s g sp s l g (w d

v a g als) pa d l s v pa d p s a s f S a d S p s a s f

al c ly Sa l yR ady (198 ) f d a s g l d by a ac l s l s l c c ld als b d

ass a v l a g s g sp s was a d fp c g pa d p s a s f a ac l C a d a av s v c as l c s p c g al pl a d a d p s a s s w d l l c as

s g sp s a d l c s p c g CS al p ss w d a s g ca d c as sp d g CS.

add ass c a v c a g s s g sp sd la d y ass a v fa s ab a f p

s g (L c yK s a 1985 c y al 985) a d c l c d s g E. B d p s al c ca

Sa y 1 8 ) av b bs v d s v a lab a s S ll a l y 198 ) s w d a c v s f abaf a 0 s p d (sp a s c v y) a d s fac l a d s c s a pl d a l f a al (d s ab a ) a



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0 A W & S

pp d p o o h b o g p od s d d h b

dd o o h sho g , h b o h s b obmod mo omp b h v o s s h s h sho k d

spo s D b sk s 198 ) d h w dspo s R 1 9 )o v s g v s d y g spo s d

f o h h, og sys m h s b d vf s - o d g sys m S) s h of y o p d

o h g g o , h m d s v o d sho g B g oR p d s m o of h s p od s h b o

spo s d o o s s m o p od s d sh b o Bs o o o dop m so d sh s h S spo s f o s b o k d bo h by p b o w h m hys g

o go s d by p b o w h m d o , phosphod s s v o d 198 ) o s sph m o og s s h sy h s s o A P hg g of h h h s b obs v fo ow g s s z o 198 ) h s d gs og h w h h k o h g s h

o s ho gh o m d h s m o s gg s s s z o A ys as b ow) h obs v d f o y

h g s w h h m y b d o s o o -m d d A P B o d 198 )ss s k ow bo h m h sms m d g h b

s s z o of h sho g sp s of h mp m s d o hos mp oy d h b v o p

y b s d s g y d d body w p pod d by B y oN ho s 19 9) App y o h g o

h o ss of p m y m h os so y s P s) m dds o g spo s R h h g s pp o o

mo o o L m d g h po S oS h y1985 .SCHIS OCERCA GREGARIA

19 , Ho dg 19 b) od d c v s mp opg p d gm s s : H d ss s sS h s e a)o o k o h s

w s sp d d bov d sh of s so o d v d h o gh od h d o h g) wh h y ow d

m d o w h h s . A m s q k y d o ho d

h d b sho d w y f om h s . h s b s p d gm h smod d d d ov h y s mb o w ys bo h o mp p o m b o s d s d o p o h


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NVE TEB ATE EA N NG 5r rc h r c v pr d c g h r g r

r y 1 sh w d h c s s c d r h g p ssk b h r rc c d r H y19 0s b

s y d h c s s r gr ssh pp rs c d r h sk r v rs v s s ( d s d/v br ) r br w rd (s g r w r r gr ss). g pr b s w h h rH rr dg p r d g r 1 r h r r d h pr c d r L c s sw r r r d p r c r r - b

v rs v h g h h d. y g g sc y gr ph cp r s d r g h r g r 19 w r b d y

hr d s c r s r g s (r p d - s sc s d c s w c r r r c v y)

s c v y p y d d r g h r g d p d g p h rg r r d A sc g v r h s r r g hb r p r d by H y d1 83 wh d h pr v N wZ d s c ( h W ") c d h b g p s r

bs c c r c c v y h g sc s g r g s gg s g h h s s p r ph r c ch k chg r h sc s c ss ry r h pr ss hr sp s .

rs s ps w rd r ys s w r d by E s sh1 65wh h w d h h pr h r c c g g (wh ch c r s h

b h c ss ry d s c r h p r r g h sH y 1 65r c rd d h c v y h c r r g

v v d h r g ( h dd c r h c ) d d hd r c y c d h r g r h r Wh d v r d h c g p d c r g r w d cr s s r s y s r s h r g rr s s w r b d s d g g - rv - sc pr(H y 1966 A p r bs rv h s s d s w s h

r g d d d p d db ck r g p s s c h g w s b d r h d r h g- sc

bs y s yH y 1 d v p d dr gr g c r d by c p r d d h h y c d c d

cr s s ( p r g") r d cr s s ( d w r g") r g r .

p r h ch s f h c d d ch g r g frW c H y 1 rs b ck d sy p c r s ss (w h h g

g w C + s ) ss ss h r s c p c k r rr s h c d d hc s r s T s

p c k r sh s pr d c d by h r g h y c g bsy p c r s ss r r g d d h h r s



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45 W & S

e e s was c eased w g p- a g a d dew g d w - a g Th s, c d g a e ed he pace a

he ce s H weve , a e a s sy ap c p a s we e

s ce he eq e cy sh s w h a s ss c ed we e s g s e s es h a e he ch ge e s we e b a ed y W acH y e ( 9 ; see a s H y e

8 a, ) wh da h he p es s a ce he ec eases w h p a g a d dec eases w h d w a g, a) ha

he hype p a g a e p e s we e a ge a p de weq e c es (d w - a g) a d d ced w h h gh eq e c e

g) hese es s a e c s s e w h he hyp hes s hap d ced a ed c a K c d c a ce he e s Ba

hese a d e a ed se va s, a de has bee p p sed (W acH y e 19 , H y e 19 9, 198 a, ) p a g ha a p ged a v age-depe de K c d c ce he e s sh

s c pace a e eq e cy hese ce s p d ce he c edec e se g es se ved d g - ea g a d d w - e

espec ve y H y e ( 98 b) he p p sed ha a e e e cem a m s v v r v g a k a "s r a ra gK + c d c a ce va es ha a e y p d ces he app p a e ca ce ach eve g a e ed eq e cy d a e a y

de s spec a ve p v des a c ea ve a e e esea ch.

H RM SS ND CR SS C RN S

he d a ch sc,e m en ash ws a e a e p s veph ac c ehav B h a pe e d a d e c sed e pe

es s ead c e wa d g s e (A 9A 19 8, w 985a) h s p s ve ph a s ca be s pp essea a ece ves p e p s e gh c c w h a(19 4) s desc ed h s as c phe e a d C wA ( 9 8)p v ded he s de s a a ass c a ve d ca

eh v . s g a a a ed a g a d es g p cedu e, hey sha h ee days g, c s s g 50 s pe day

p ed w h a ( he ) ed a s g c c ease esp se a e cy e ea s gh . ps ece v g h s pa ed a

sh wed s g ca y g ea e s pp ess ph a s ha a c g ps, c d g h se ha ece ved a d pa edp ese a s, a d C a e he ea g was e aseve days a d, p e a g, he pa ed g p sh wed sav gs

eacq s pe a ce seq e h s s desc p


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19/53

NV RT BRAT ARN NG 5v l a g ermissen a h o s a o al s

rolo g r o a sav gs o h ro r s of l ar g havb a . o a l s l s s has b r o

a l Alko ( 80 8 a ho a h s of or ah l ar r s o s a r s o s s c has b o s raro bach ( 9 9 98 ho fo ha cr as r s o

c s follo g l ar g r o o a d c s g ral b s g c c s l cy c d d lo o o o o a l gh . Mo l , o of h o(R ha s a 8a o g s ( a lK 85 hav alsob sc b n a o L rh l r al ( 98 hrov

v ha fol o g a r g of gh a ro a o gh

o ra o of h foo ha s s m lar o h o ra o s roro a o alo all s g a r b hav oral as rsha o r s o s ( h ch a als ll avo ark ss b g a l

a bo o (198 ) av gg o og ha g se m ssenda' ab l o l gh ark s

A ll lar a al s of assoc a v l a g e missendaha b aoss bl b h r l v s l c a a ss b l of h s so

volv h v s al ( a h a a v s b lar (U a h a sh hav b a o rabl a l (fo v s

98 984 . Th s of e missenda o a o l v ho o ors ach( o T A a hr T B all of h ch r s o o l gh ola g g ra o o al ha g v s s o ro aga

o als (D 967 Alko s 97 ar c lar t hab foc s o T ho or c ors, h ch sho a lo g-las

ola a o (L D ha o las a s s a o ofl gs co s a s a o a b a r as b a s s a (A

Gross a 978 Th o c co c a c s ho o c o ( ha k hav b sho o l h o a K c r s [ ar

la (10 a a - d] a ar a co c a a (r s o s o gh a arl ar + co c a c a a o a + o a (Alko 84 . Th s ag

s s a r h ho o ors h o c ga gl o h h haso v s al o s. Th v s b la a a a s ( h s a ocys

ha lls h h sho as ls a v s o s h sy p c c s g h s l s h b

r z a ca b s ar z as follo s T A a B ho oc orsh b a h o h r al ha lls a T c lls h b ll

g g o ( a s T B s a hc lls ( l o 9 8 984 W a 82 Mo lb a f g so o o ro s oss bl volv



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c c b h v or row 8 , r ss& Al o 8 dAl o 82,Goh Al o 84).

Ev d c mpl c pr m ry h h yp B pho or c p ors s r sul of h l r w s rs ob d by CrowAlko 80), who

x m dB c ll v y s v r l ho rs r d o d spo o s v y o d r d p B pho o p ors w s s

r s d m ls h r v d pr v o s p r d r omr domly d co rols Mor ov r, h s p r d B pho or p omor d pol r z d r s po ls d h h r pu r s s c s, v

xo om z d d c h h s c s w r r s c o F r h r s d s by F l yAlko 82 ) d W s l 82) h vco m d d d d h s obs o s 2 d ys o low

yp B c lls from p r d m ls w r o lo r d pol r z d, b d d shcr s d l h r spo s s d s d l v d pu r s s

Ch s p v mo o ro s corr l d w h r h v r por d Crow 8 , k dAl o 8 Goh 85). For x mpl

row 8 ) show d h h p v p d l mo o ro 5) xhs c h h r h b ory sy p c p r spo s o l

omp r d o r domly r d m ls Mor r ly Goh l r por d h o h r pu v mo o ro MN ) show d sm

cr s mpuls fr u cy r spo so l h s p m ls r c v

p r d r comp r d o r dom y r d co ols, up o 54 hrr d h MN s r spo s o l h w s pos v ly corr lh m ls b h v or l r o l y d v rs ly corr l

pho or c p or pu r s s c . F lly, dd o o rh s l d o cr s d xc b l y B pho or p ov d c s s s h r prod c s co com d rxc b l y of yp A pho or c p ors R ch rdsrl y 84). oll c v ly, h s d h v l d o h hypo h s s h h ssh pho o b h v or s d o r -prod c den n emeno

h yp B pho o c p or s l h d d r or po l v y l r v m h sm volv de ease h B ll sr spo s h s b propos d by row 8 b s b low) h s r r B

c vy s ho h o r s B-c ll h b o o o yp A c lls, whhypo h s z d o b r spo s bl for m d pho o x s hrou h

o c o s o r u o s d mo o uro s lko 8 84)A h x l v l o lys s, l o d coll s x m d

s o d r o of h o d o d ch s hpho o spo s

cqui i ioh c u s o proc ss h s b pos ul d o volv p r -s

m l v pol r o hB pho or c p ors h occurs ross


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R BRA 55

tr g r s (A o 1979, 980 ,b) h s mo s p rt y probs rv t o th h po r g of th g r or pot

m s r v g p r r g ( rowA o 1980) Mor ov r th sh m t b m m by p r g gh ro t o s

pr p r t o (A o 1980 ,b) or ght u h r s muso rvous sys m ( r yA o 1982b).th s so b su st

th t m mbr h g s th B s b us y r t o ssh g s ( r y 198 ) h umu v po r o s hy

to b r v from two sour sa)r bou x o h B s t hrm o o ro t o u o r s from h b o rom u

x t tory pu from op o s wh h thr bou ( t th o s of p r h rot t o ) from h b t o from h B s h r s (A o 19 9, 1980 ,b)

h sy pt x t t o fo ow st mu us p r s thought h by m s of pos t v f y y pt pu p

m mbr po r o , w h us s bo h r s of vo t gt urr r s t v o , o o by pr

v t o of wo Kurr ts A Ie , wh h fur h r r s sth put r s st of th B (A o t 198 b A o 9

r s pu r s st w tur h subs u syg v g r s to gr t r po r t o , so o r g s mu

ty r thus hough o r su t from h umu t v pot o th t r s s ou of pos t v f b y b w sy p u po r t o A ss f ur of h s p r g mo

S US show s mu ousset. hus, th s mo , pr p , woupr t th U o s t fo ow som t m t r by o s ( . . b

o t o g), wou st pro u sso v h g s f ht rm t s mu ous y

h t r su t of umu v o r t o s hou htob r s umu o th yp B photor ptors (A o 19 9) R

xp r m ts by o orA o ( 98 ) r o s st t w th th s hypoth s sUs g th + tor y , Ars oh y fou th t p r g gh ro t o r s tr u r th B photor ptors ov r th t

pro u by ght o ow v r, h ss ry ro of r u K As w s th ro of h s r t y b brough to u st

Grov r r y (198 ), who show th B s st xh b t psp umu v po r o whA c r b o whth s o th b h so u o

o to r s f tors su h s umu v po r

umu t o , sy p mo u tory s h v so r tmp t e missenda or x mp , Wu r y ( 98 ) r port ths m r toA ysias sory uro s (K K 1980, K t 982),s roto r s s pu r s st r s Kurr ts h s


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56 W S L

a2+ currents n photoreceptors. M re ve x ac f p c a l aha b sh w c as B c ll pu s s a c a d D McEl

Fa l y 1 83 . h p c a l a c a u ha xh bu c c w h a Fal - lla p a hu su h p

ca ch lam N p ph a d cl d(O2 ad c c pag n s s have recently been shown reduce both I a d n cells

a a ba a a 1 8 h ha p p d h u ha a 02 c p a s f m c lls h p c a may fac l

duc f a ddu ac u a a ba a al 1 84 Al1 84

Re en onR f h l as h u h o nvolvea a + duc d l - m

duc f a l as w K cu a d al l s supp h hyp h s s f w Al 1 8 F xamplal 1 82a b f u d ha was s ca ly duc d h B c lls 1a pa d a a d h y 1 82b h w d ha d c of

ntracellularCa2+ reduces l , but does not a ect another K current IBwh ch s a a a cu c s s a as h a K

a d h calc um d p d K + . c as h s a lm c xp m s als mp ca h Ca + -d p d K + c

Fa l y Al 1 8 F ma a 8 Fa l y a 1 8 h m p s ula ha h p d f day ha h la d h m mb a p al f h c ll u p a

but a d rema n reduced The magn tudef th s reduction 0 40%c mpa d c ls s h u h b s c acc u f

ha c d a p al bs d c ls days a Al 1 84 .On a b ochem cal level, an mportant clue concern ng poss ble molecular

mechan sms nvolved in the retent on o the learning came rom a study by

Neary et al 1 81 wh h w d hacond t oning nduceda cha hl l f c p a 2p a 20 000-m l cula w h ph ph p h y s ferm endasu s ha p ph sph yla m

c bu M , N a yA 1 83 b a dd c ha h s p ph ph yla wa la d acf '

and eary et al 1 84 h w d ha h ph ph yla wa ulab hCa a d cAM f w N a y 1 8 . F hstudy ofp oteinph sph yla was ca d u by Al al 1 83a wh ca aly c subu f h cAMP-d p d p as h

a d f u d ha duc d h la K + cu s m ha hlAoS c duc I was h u h b m l d h l aAl a 1 82a cAMP-d p d ph ph yla wanot considered



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NVER BRA ARNING

to b m y m c sm ( l o t 1983 ) t s o tct o oduc s su ss o o bot ( l o t 1982b) d

t 1983b). s l d o ocus o os o yl o by m s o C

c lmodu m c sm R c y, cos U u d ( 8 )o d t c oo s c s o c modul d d os c duc botA d B B o o c o s y o os

c llul C ccumul s du , v s c modu m d d os o y o ofIA d c s, d sust d os o y o of s K + c s cou d oduc o

t o o t ssoc v

An A na iv opo a

R c o e m en aby o s s d o m o ssu sco c g b v o l sul s d c llul mod l o ol o d co o s : s ssu dd ss s t u s o o t

c t v d o ssoc t v com o ts o t co d t o , d s s t s o o t com l x c cu t t ct o s b

d oto c o s c ss y o x l co d o d b s s o b v o l x s C o (1983 ) x m

co t but o o ssoc t v d o ssoc t v cto s to t co dsu s o o oto ct c b v o o e mi endaby v y g t umbo co d o g t ls d t t m b t g d t st g. F( v o ) s l s ss o oduc d o c s b v o

d t ly d s o d o y o ssoc v d o m c com bl to dom co t ols) t st d 1 m

lo g . Co s s t t v ous obs v o s (C ol o 1978, C o 1979), mul s ss o s (3 d ys, 0 t ls c ) s

o ssoc v c s st o s v d, ollo d 30 m by s c ssoc v c s o (1 83 ) m

s c t g t o ly t ls oduc d o ssoc v bc s c l ul s ud s us v y ls ( .g. l o 1980b

l o 1982b) m y o y ld u l co l s c b d ctly b v o co d o c s udy o (1 8 b) dd ss d do

sul s of t b v o l d c llul s ud s o Al o d o- oe mi en a os t v ly oto ct c d co d t o g sults

su ss o o t s o s , y t t s b su st d t t oto s o s B oto c to s s c c l o oduc

(s l o 83 1 8 ). o c en an e o o s o s oduc d byco d t o gde ea e otot c c b v o ? As d sc b d bov l o

d coll gu s o os com l x c cu ct o s mo g y oto c to s d t l t d t u o s d moto u o s o



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24/53

458 EW & S E

f r e ecrea e e a r ( lk n 983). r w ( 985a, ) pr p e al erna e m el. n a p c op ca u , row ( 98 a) foun

e m en a ow a gra e ncrea e n po e p o o a n re pon

an ncrea ng er e of e l g n en e . on on ng regn can uppre n f p o o ac c e a or a a l g e am ne nclu n n en e n w c e LLD n p rw ul e e pec e e e er a en r n a ec e c n( arle Alkon 98 a) ow ugge a ome o er c ange n e m g accoun for e con on ng

n a ec n paper row ( 985 ) a re e que n re lrea ne a e appr pr a e c n n n w c mea ure c an e

p o recep r ul c rre pon e n w c c ange n p

ac c e a r fe missenda are ac ua l e pre e ur ng e a rale ng Spec cally ur ng e a ral e ng an mal are u jeca ap a on an o no eg n o em a p o o ac c re pon e for em nu e . erefore row e am ne e -ce l p o orecep or re pon e pre u ly c n ne an mal ) mme a ely a er l g na e 5 m n f c n nuou l g . W en e am ne n co nu u l ga er g n e e -ce l re p n e wa gn can l re uce n a

a rece e pa re ra n ng compare o ran om con r l n f -cell ac an ( n a m ze cell ) n e ampl u e f e

a e genera or po en al u w en ac of p o orecep oe am ne un er c n on f llum na n a re em le c n nw c c n ne e a r e pre e , c n n n pr uce r duc

on n e cell re p n e n ng e oppo e of a o er e Alkon an co wor

w foun in eased cell ac af er con n ng ( l n 98 ) npre u ly pu l e rep r lk n an c w rker ( ee lk n 9

984) cellular c rrela e a e ee a e up n analy f a ery earp a e of e -cell p re p n e. r w argue a n

e a rally rele an p a e f e p recep r re p n e e amnce e e a genera e only a er e an mal a e een l

a ap e for man m nu e . A an al erna e o e ne work ypon l ng -cell n n f ce l ( lk n 98 984) r w ugge a c n n n pr uced m n shed-cell ac y a

rec ly ran lae n o m n e p o o a . a po le mec ane l ng- erm re uc f e cel p o re p n e, r w rep r e ru e a w a c n on ng pro uce w c ange n

a n pr per e of p ecepa a gn can nc ea e n l ge oke p o orecep or e en za on a canno e m m cke

l ck n ana gn can elay n rec ery f en y f l w



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25/53

R BRA AR G 5

p g l gh ( ow 82 83 . S c + l v l homo ul e c g ( lko 8 k ow pl

mpo ole ecep r p o gul of cellulml c r u e mp r l co o e l er o f he

p ope e of pe pheceph propo l l o colle gue ow e o ec

mu u ll xclu v o h m ch m coul co u e o lHermissenda. ur her work w ll u u e l cl r f h ue o

P S ORN

A ys a a f n a m mollu c h h e u e x v

u c ve well c ve f rm f le r g. lp m focu of h ch p o oc v le r g oc ve le g p c c ll h u o e

por ckgrou or e oc ve work ever more rev ew h ve ece l ee pu l he (K elSchw r 98 , K el e

l 83 K l 84 M po m A ys a xh h uo z o . he clu he g ll ph w

e ex ( ke e l 0 he l w h w l e ex (W l er eee elow k g ( rewK el 9 e c pe locom (W l er e

9 8 he ee em (Kup e mP ke 9 8 . We e cou cu o u e of he g ll pho w h w l e ex mo x v l l ze . W v u cu o

r co e g eh v r l he cellul u e .

B hav ora S d of Hab a on and S n za onec u e mpl c el vel w ll l ur l eh v r h h ece v h mo expe me l e o A ys ahee he efe v w h w l ex f he m le o g ( he g ll

m l helf h x c e el c e l gh c le ph , wh ch ev ke w h r w l of h org o h m l

he me mulu repe e l el vere he re ex e l h(P ke e l 9 0 em f r ever l m u e e

oweve e ex mpl u e c e mme el e o e el veo g e z g mulu o o h h uch l wh ch p uc h u . D h u o mpl

c e more ge er l f c l pr ce e o ( rew o h h u o e c e me o

rm g week g v g repe e g e v ( w , k e 3 .



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460 AR W & SAH

C u a S ud f Ha ua a d S ae nve g on of u on nd en z on of e g l

re e nA /ysia een c rr ed ou on ree evel : n p c, opnd molecul r.

S AP A A S S e neur l c rcu for e g l nd p on w dr wre e een e en ve ud ed nd re vel we l undercon of m l popul on of ou 24 den ed pr m r neuron , p on mec norecep or , w c m ke mono n p c e cconnec on on o den ed g l nd p on mo oneuron we l

n erneuron ( rne e l 74, ellucc e l 70, Kupferm nn e74 erlm n 7 wk n e l 8 , . Sever l ne of

nd c e u on nd en z on re due n l rge omo n p c depre on nd e ero n p c f c on re pece n p e from e p on en or neuron on o e r r

( ellucc e l 70, rne e 1 78 . Moreover, qu n l nen or neuron n p e dur ng omo n p c depre on nd em

n p c f c li on owed o of e e proce e re preior g n Cas & Kan 74, 76) In a on o s or - rmc nge , e e me n p c connec on ve een own o e cc l depre ed or en nced n n m rece v ng long erm u

en z on r n ng ( ellucc e l 1 78, rew e 1 7K ndel 1 84 . An mpor n n om c correl e of e e o erv oeen de cr ed e en ( 83 w o found e num er, ze,nd ve c e complemen of c ve zone ( n p c rele e e n

neuron erm n l re gn c n reduced n long erm u ed nd gn c n ncre ed n long- erm en zed n m l . Moreov

recen ud , e en (1 84 found ong erm u edn m l ve gn c n fewer v r co e (pre n p c ermen or neuron n con rol . ence e e mp e form of e rn

produce profound u r ruc ur l wel func on l er on en or neuron n p e of e g ll w dr w l re e .B O H SI A A A SIS o n l ze e on c curren modul ed n e

en or neuron dur ng u on nd en z on Kle nK ndel(1 78 1 80 e m ned e dur on of e en or neuron c on po

nd found w reduced n dur on dur ng repe ed c v odr m c ll prolonged dur ng e ero n p c f c on. Accomp

e e er on n p ke dur on, n p c r n m on w

or en nced. Vo ge c mp ud e owed u on wcomp n ed progre ve decre e n vo ge en ve nw rd2 curren (Kle n e l 1 80 Recen qu n ve mul on u


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R BRA L R G 6G n r chB rne (1 ) e however ha Ca nac va on ma no

acco n or hab a on an ha ep e on o ne ro ran ma o p a a gn can ro e K e nKan e (1 0) r her o n ha

he ero nap c ac a on wa pro ce b a ecrea e n an o warc rren ha pro onge he epo ar za on o he p ke an h enhahe vo age en ve Ca c rren , wh ch n rn enhance nap c

re ea e. In a on o mo a on o Ca e o change n ra oac on po en a a componen o Ca acc m a on n epen en o

hape ha recen been en e . Bo e e a (1 ) njec e ne ron ce bo e w h he Ca en ve e Ar enazo III an

ha ero on n enhance ree a acc m a on n re pon e o conra on vo a e c amp epo ar a on n ca ng ha he Ca change

were me a e b ome n r n c change n he han n o Ca ben or ne ron Th n r n c enhancemen o Ca acc m a oho gh o ac nerg ca w h he re c on n K c rren ran m er re ea e. Con en w h h ob e va on G ngr cB rne

(1 5) ge ha n a on o p ke broa en n , ran m era on ma a o con r b e o he ero nap c ac a on.

evera ne o ev ence e ha ero on n one o he aran m er nA ly a(Br ne e a 1 76 Ba e e a 1 1 K

1 3 1 5). Imm noc ochem ca e n ca e ha ero on ncon ac he en or ne ron (K er e a 1 ) ero on n m m c

nap c ac a on b broa en n he ac on po en a ecreacon c ance an enhanc n nap c re ea e rom he en or ne r(K e n Kan e 1 1 0) A vo age c amp ana howe hac rren mo a e b ero on n er rom prev o e cr bec rren ( he e a e K a K Ca epen en K an m

en e K (K e n e a 1 ) A pa ch c amp ana b ege baco eag e ( ) r her charac er ze he ero on n en ve K ( he " channe wh ch ve r e o a n q e K c rren I ) n hene ron I ac ve a he re n po en a mo era e

epen en an n epen en o n race ar Ca . C o re channe can acco n or ncrea e n p ke ra on Ca n xenhance ran m er re ea e re pon b e or en za on ( ege b1 ).MO U AR A A SIS In nve ga on a a mo ec ar eve mo progrha been ma e n he ana o hor erm en za on. Ba e o

n epen en ne o ev ence ( ee Kan echwar z 1 or rev ew),K e n Kan e (1 0) propo e a mo ec ar mo e or en zapo a n ha ero on n ncrea e a en a e c c a e ac v ne ron erm na hereb ncrea n n race ar c c c a eno n



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6 C EW & S LEY

hos hate (cAMP). cAMP n t rn stim lates the act v ty of enzymes ( rote n hos hok nases) res ons b e for hos hory at n s bstrate rote nsone of which is the channe or a rotein c osely associate with it

Phos horylat on of the channe closes t ivin rise to increaserat on of the s ke more a n x an th s more transm tter releaseonsistent with this s estion ie elba m et a (1 ) fo n that cAMP

closes the channe n cel attache membrane atches an more recet yh ster et al (1 ) have fo n that the catalyt c s b n t of the cAMPe en ent rotein kinase also c oses the channe in cell free membraneatchess estin that the kinase ac s on the internal membrane s rface to

os hory ate e ther the channe tse f or an assoc ate rote n thatre lates the channel.

ort for the molec lar mo e was obta ne by aste cc et al (1 0whofo n that nject on of the cata yt c s b n t of the cAMP- e en entrote n kinase into the sensory ne ron broa ene its s ike an increase its

transm tter re ease Moreover fac tat on set n mot on by seroton n can blocke by injection of a s eci c rote n inhibitor of the kinase ( aste cciet a 1 ) s estin that the time co rse of sensitization is e to the

ersistent activity of the cAMP- e en ent kinase (an not for xam e tothe stable hos horylation of the s bstrate rotein). onsistent with thishy othesis Bernier et a (1 ) have shown that cAMP is elevate three tofo r-fo in sin e sensory ne rons in res onse to serotonin or stim lationof fac tatory ne ra athways an that the t me co rse of ts elevat on

arallels that of the resyna tic faci itation. Ex eriments involvin thein ect on of a b ocker of a enyate cyc ase (G P - ) in icate that thee evat on of cAMP r n sen t a on a ears to be e to ers steactivity n the a enylate cyclase com ex (which has at east threecom onents a rece tor a re ator G rote n an a catalytic s b nit)th s s tho ht to be the t me kee n ste n short t m sens t at( aste l cc et a 1 chwartz et a 1 )

Recent ev ence s ests that m lt le fac tatory transmitters mayconver e on the a enylate cyc ase com ex n the sensory ne rons. Abramset al (1 b) have sho n that n a ition to se otonin t o en o eno sne ro e ti es inA ly iathe sma car ioactive e ti es PAan Pfaci itate syna tic transmiss on from the sensory ne rons by a cAMP

e en ent c os re ofthe channe . Moreover recent i m nocytochemicalst ies by Ono Mc aman (1 ) an istler et al (1 3 1 5) haveshown that one class of i enti e interne rons inA ly ia,the ce s

hich ro ce resyna tic faci itation of the sensory ne rons an whichwere ori ina y tho ht to be serotoner ic (Hawkins et a 1 1b), o not seserotonin as their transmitter Th s i a ears that there are at least threea ents that mimic heterosyna tic faci tat on as well as the nat ral


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NVERTE RATE EARN N 6

ran m er o ha may conver e v a eren rece or on a commonmo ec ar ca ca e n he hon en ory ne ron (Abram e a 1 8 b

In recen year a oc a ve earn n ha been cr be n a n mberbe av ora y em nA ly ia, nc n(a he e en ve aro a y em)

he w h rawa re ex o he man e or an) he a w h rawa re exdhe ee n y em ane) he ea -wav n y em We c each o hen rn.

Condi ion d D f n iv A ou aIn one o he ear e e o c a ca con on n nA y ia,Wa er ea (1 7 ) em oye a R ara m o exam ne w e erA ly iaco

earn o a oc a e a chemo en ory C (a r m ex rac ) w h an averUS (hea hock). he behav ora n ex o con on n wa mo ae ca e ocomo on by he chemo en ory S. on ro rece ve n a rCS-US re en a on he CS a one or he US a one. ra n n a e

ee a e ay o owe y e e ex ay an ma howe com ara e e ca e oco o on n he ab ence o e

owever, he re ence o he C a re an ma howe n cmore e ca e an a con ro . In ere n y he S by e nobv o ocomo or con one re on e a er ra n n ea n Waa ( 7 ) o ro o e ha he CS a come o e c a cen ra e en

(ana o o o con one ear") ha mo a e e ca e ocomor her e no on Wa er e a ( 8 ) exam ne a var e ybehav ora re on e n he re ence o he hr m C o ow n

ra n n an o n he e re on e mo a e by he CS n a mo vy on en a h on o r e en ve re on e (he n

w h rawa nk n an e ca e ocomo on) were n can y ache re ence o he S w e a a e ve re on e ( ee n

n can y re e . The e be av ora re h ocon one - ear hy o he . r her or wa ob a ne by arew

( 8 a) w o recor e n race ar y rom mo one ron n he eocomo on hon w h rawa an nk n y em rom rera ne an con ro an a . In he re ence o he C , an ma

rece ve a re ra n n owe n can ac a on o ynaa he e mo or y em wherea con ro howe no ac a o

ro ce no chan e n re n o en a or n re ance o mo onn ra ne an ma e n ha he cr ca chan e ro ce

were re yna c o he mo one ron he e re o e her wbehav ora a a e he e a hor o ro o e ha he conexh b e byA ly ia n h ER ara m re ec an a oc a on be weehe S an a cen ra e en ve aro a a e re emb n con onemamma


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6 W & SA

Condi ion d Wi hd awa of h G land SiphonC a ca con on ng of g an phon w h rawa ha been

wo prepara on rg ca y re ce n v ro prepara on an an ma prepara onI VI RO O I IO I L kow ak Sah ey (1 1) pa re a pho c

m o he phon a he CS w h a rong ac e m g a he US. orma y he pho c m oe no pr ce g

rawa b af er repea e pa r ng w h he US he CS came o ew h rawa n even o of en p epara on wherea never ew h rawa n con ro prepara on ha rece ve ran om CS-US pre

a on . S b e en y L kow ak (19 ) e cr be a new para hamore amenab e o ce ar nve ga on a weak ac e m

phon e ve a he CS wh e he US rema ne a rong ac e he g . Af er repea e CS-US pa r ng he weak phon CS evoke

enhance phon an g w h rawa re pon e wh e a n mber of con(ran om CS-US pa r ng CS a one an US a one) howe no enhancemen

r hermore L kow ak (1 3) recor e n race ar y from g mne ron r ng pa re ra n ng an fo n ha he phon S evoke ar

ynap c re pon e n he e ce a ra n ng progre e . F r her emen n ca e ha ncrea e ynap c ran m on from phon ne ron con r b e o he fac a e np n he g mo one ronv ro prepara on prov e he oppor n y o ana yze ce ar evenC S r ng he acq on of a earne re pon e.I TA T A IMALS Carew e a (1 1b) exam ne whe her he g an phonre ex n a on o en za on m gh a o how a oc aThey pa re a weak ac e m o he phon a he CS w h ae ec r c hock o he a a he US. A er repea e pa r ng r a h

o e c gn can y enhance g an phon w h rawa comp

con ro wh ch rece ve an om o npa re CS-US pre en a on oCS or US a one Con on ng wa ac re w h n 1 r a anre a ne or evera ay . More recen y ng a re a e para gm Caa (1 3) have emon ra e eren a c a ca con on ng of w h rawa componen of h re ex In a eren a con on ncon one m are e n he ame an ma one (CS+ ) pa re w h

he US he o her (CS ) pec ca y npa re . Th each an maerve a own con ro . Weak ac e m app e o he

man e he erve a cr m na ve m (CS+ an CS ) an ahock erve a he US. eren a con on ng co be ac reng e r a wa re a ne for more han hr an ncrea e n rengncrea e n mber of r a . Moreover con on ng co be pro ce


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VE TEB ATE EA G 65

wo e a a e e on he hon k n a c m na ve m l ,cell la ana y o e en al con on n n en o y ne on

hon ( ee elow). nally, Hawk n e al (1983 ) o n ha

n e m l n e val nc on (wh ch e c e he C U no ce e ec ve con on n ) wa q e ee el a le lea no ce only w h a C U n e val o 0 ec lea n n wa le a

n e val . o con on n occ e w h o wa n e val o ec, o w h ackwa (U ) a n . o eove , hey o n ha

con n n e en e o he con n ency e ween he C an U, on he e ee o wh ch he C e c e he U .o nve a e he cell la mechan m o cla cal con

Hawk n e al (1 a) e a e en al a n n oce e n h

c c o hon w h awal. hey eco e om wo hon ne on ha o h ma e exc a o y mono yna c connec on onhon mo one on a l hock o ce n can ly ea e ac

o yna c an m on om a en o y ne on he hock wme a ely ece e y ke ac v y ( o ce y n acell la c

he en o y ne on) han he hock an ke ac v y we e ecn a e o he hock occ e alone he e e l e e em o al ec c y o he con on n m h e e o an ace en en enhancemen o ac l a on. he ev ence e eh enhancemen e yna c n o n : Hawk n e al (198 a) exahe a on o he ac on o en al n he en o y ne on ( n e ae hylammon m) an o n ha he e en al a n n o

n can ly ea e enhancemen o ke d a on n a ed comnpa e en o y ne on . Ex en n he e n n , ecen vol a e

ex e men y Hawk nA am (198 ) e ha e ac ve en en ac l a on nvolve mo la on o he ame on c channe

n o y ne on ( he channel) ha nvolve n en a onThe e ex e men e ha he mechan m o cla cal con

nA ly iamay nvolve a em o ally pec c, ac v y e en en enhancmen o he ame cell la oce ( e yna c ac l a on) ha

n en za on. he ev ence ha h mechan m can acco nehav o al con on n ha een o a ne y Cla k ( 19 ), who o n

j a ackwa (U C ) pa n oe no o ce con on n (He al 1 3 ), ne he oe o ce ac v y e en en ac l

yna c an m on o wa pa n o ce n canac l a on han ackwa a n .Ano he mo el o con on n em ha z n he m o ance o ac

ha o o e y D He (19 9) who o la e a c cal o yna c ac v y n yna c ac l a on. Ca ew e al (1e e h hy o he nA ly iaan o n ha o yna c ac v y (



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466 AR W & SA

ip o moto u o t at m diat t o ditio d po ) a ita y o uf fo a o ativ y a ti a d a

l a i t i i ta o ditio a ot du to a bb- y m a

but at o a ivity d d t a m t of y a ti fa ilitat a t v y-d d t mod l di u d abov m a i ip b t iti atio a d it ampli d fo m la i al o di ioi mod l t u ai t o impo a t u io o do a i i

mol ula ma i y i volv d i i atio ( p i ally a ad ) W a a p of a ivi y i im o a fo am li a

lim a y vid add t t u t o u t t t ta ad i di l modula d b pi a tivi S o u o

b i f appli atio of oto i (a t US) imm diat ly aft a t ai of pi

o a fou -fold at l v l of AM a ll x o d o 5- al( a d l t a 9 Ab am t a 9 4a) W t a d o t o d qo li of v d u t a a b i f l va io of a oduik a tiv y t o y u o m v a al f

p i i by i a i ll o mal AM po o (Ab am t al 19 3)1 A tio ot tial ai d it oto o mally a f

ato y ( pi b oad i ) po o o o i i o y u

t i a tiv t -d p d t a m t do ot o u i ta sence

ofa i ux (Ka d l t a 983).. S imula io of ad la y la f omAply ia vou ti u by

o o i a d ub a iall by a / almodul . abio o i o a iva y la i ma k dly du d follo i d

of almoduli o follo t additio of a a moduli ib to ( . WAb am t a i p pa atio ).

Alt ou t ult a p limi a u t at la iditio i a d iti atio i ys a a at l a t pa t of a ommoml ula a ad a d t at a i ux du i a tivit ma b a iti

m fa o fo a m t of t at a ad du i o ditio i

C a i a Condi ionin in h ai Wi hdrawa R ail i d a al x iAp ys ai m dia d by a id ti d lu t of

m a oa t o u o t at y ap mo o y apti ally omoto u o alt t al 983a). S iti atio o t x a

oto i odu a i a i y a ti t a mi io f om t (Wal t a 9 b) additio o o i p odu a i a

i ta a d a o omita t l vatio of AM ( yWalt 198O o t a 1983 olla k a 9 ) Mo ly Waly (1984)

av o a t la a tivato fo oli mimi 5-HT poi t o u o a d lo u u t 5 HT po u


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NVE TE ATE E N NG 67

a s a s a ou a ommo sa u a l m a ism. umo O o y 985 av s o a i addi io o 5- S

u o yp a o am d l va s l v ls u o s a d a 5- a d S p odu a a d u asso

i a i as i i u sis a . ak oll iv ly s da a sa s si iza io i ail i d a al x los ly s m l s all a d s o d a al x o may volv a s o o sM m d a d d as K o du a .

I addi io o o asso ia iv s d s i d a ov , su o s of ail i d a al x also s o l a asso ia iv las

Wal s y 98 a) a li d a llula a alo u of a di io di io i o du o di simul a ously f o s

u o s a d a ail mo o u o . O s so y u o S ) as a ivayi a llula u ai d i ail s o k S), a oas a iva d u ai d i S a d i d s si iza io oas o a u i a i . ol o v als ai d s so y u o s S ) as si i a ly fa ili a d om ao S a d s si iza io o ols Wal sy 983 fu

o s v d a S u o s s o a slo d ola iza io a S ilS o s si iza io o ol u o s s o a y ola iza io .

su s a , as a fu io of is d ola iza io a vol a -s si iv o du a a iva d a s i o ial may modula dasso ia iv ai i as d o s sul s, Wal sy 98 a

opos a l ssi al o di io i is odu d y a ivi y-du omodu a io of sy a i a smissio , d a a ould a m

i a llula ss volv d i is o ss a s s ofxp im s Wal sy 198 985) av also su s d a

ivi y-d p d u omodula io may a m a sm of losy a i a m simila o lo - m po ia io ommo ly s v d ma mal a C S.

R s ud s y O o al 983 985 av x d d s amol ula l v l. y us d ila al lus s of ail s so y i a io mi al a alo u of lassi al o di io i pa adi mD ola iza io of a lus y m a s of a 5 s xposu o is a a s v d as CS, a d a 15 s xposu o 5- s v d as O lus iv d pai d o sp i ally u pa d

s a io s ollo i o ly o ial a d lus s osi i a ly l va d A o ompa d o u pa d lusalo a d S alo p odu d o si i a a s. limi a y s udip osp od s as DE a v y y O oy 1984 su s a

as s du a l as pa ly o s mula o of y las a a i i io of DE. ak o s sul s l d O

985 o su s a a a i -sp a m of



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468 A S

perhaps by means of a a -ca modu in dependen s ep, may be abiochemica mec anism for associa ive learning. T is hypo hesis can bedi ec ly es ed in view of e recen e or by Ing amWal e s (1984) ha

he ail wi hdrawa e ex is capab e of di eren ia classica condi ioninghe simi a i ies of he behavio al, el a , and mo e la es l ss udies of associa ive condi ioning fro o sys ems inA ly a ha havebeen independen ly analyzed ( he e ex wi hd awal sys ems of he man leand ail) are s riking Resul s f om bo h sys ems e p asize he impo ancof ne onal a ivi y, a2 +en y y lase a va on, an ed ion of a +conduc ance for a pai ing mechanism.

Condi ioned Ce a ion of Feedin Re pon e

A ly aare herbivores a norma ly feed on a va ie y of sea eeds.Suss ein Sc arz (1983) have described an in eres ing form of associaive ea n ng in h s feed ng behavio hey esen ed an ma s wi h seaww apped in a p as ic ne , allowing e anima s o as e he food and a em

o ea i b no swa low s he ne -enc osed food be ame ohe b cca cavi y Anima s receiving is rea men rapidly earned o ce

res on n o s ne e foo an e or as a n a ne or a e2hr. n essen ial componen of e ea ning is ha he food become s uck

i in e buccal ac ivi y ; his e a ho s s gges gives rise o s imuli

become paired wi lip s imuli from he as e of he food. The learning alsodisplays s im us speci ci y lip s imu i occ ing seq en ia y ( a hepaired) i h food s uck in he b ccal cavi y do no prod ce he condi ionedcessa ion of feeding Recen y M Schwa S Ma kovich and ASuss ein (in prepara ion) have fur her speci ed he parame ric fea ures of

he learning and s ow e o e c ea ly dissociab e fro adap a ion andsa ia ion. Fina y, SchwarzSusswein (1984) have found ha anima swhose esophageal ne ves a e c ake wice as ong o ea n as s amcon ols, sugges ing ha ese ne ves ca y info ma ion fo lea ning food is inedib e. hey sugges a gu inpu s in hese nerves can ac asposi ive o nega ive einfo ce s of feeding SussweinSchwa z ( 983)s gges a e earning n eir para ig resem les operan con i ioninsin e e ange in be avio is on ingen on he onseq en es of hanima s esponse Since he ne al circui offeeding has been examined inde ai ( ohen e a 978 Rosen e a 1979 98 ) his p e a a ionconsiderab e promise for a ce ular ana ysis of an in eres ing form ofassocia ive earning

Op an C nd n n f h H ad- Wav n R p nA ly a eadi y show side- o side head waving behavior in a numbe ofbe avio al con ex s incl ding sea hi g fo a ho d fas , egg laying, and i


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R RA I 6

sp n an us y" p r ng h r nv r nm n . ar w (1985a,hav r c n y sh wn ha h s r sp ns can p ran y c ndSp c ca yAplys acan r ad y p ran y ra n d (w h n 10 m n

m d y h r h ad wav ng r sp ns , ncr as ng s r qu ncy nh dy n rd r rm na h pr s n a n a s r ng ghan ma s nd av rs v . Y d c n r s d n acqu r h p ran r sp

w n s v d nc sugg s ha h p ran r sp nd ng s undc n r r n rc m n c n ng nc s :

. Wh n h c n ng nc s ar r v rs d ( h r g na p s v spun sh d and v c v rsa , c n ng n ra n dAplys a( u n y dc n r s) sh w a s gn can r duc n n h r r sp nd ng and wh

r g na c n ng nc s ar r ns a d, c n ng n an ma s ncsh w s gn can ns rum n a r sp nd ng h r g na s d .

. Y d c n r s d n arn h p ran as , y h s s m areadily arn when re nforcement s made cont ngent upon the rr sp nd ng.

na y, c n ng n ra n d an ma d n app ar us rn(such as v sua s mu d rm n h r dy p s n dur ng ac

h p ran r sp ns . Ra h r, n rna y d r v d cu s ( .g. pr prr r a r n s gna s app ar pr v d mp r an d ac du

acqu s n h p ran as .S nc s m h c u ar m n s nv v d n h h p ran

and h r n rc m n pa hways ar n wn ( n ng a 979, Jaarwar r dman s nac ), s rm p ran

c nd n n nApl s a ay p v qu us u n s udy ng h cm chan sms ns rum n a arn ng r v r, s nc m chan smc ass ca c nd n ng hav n uc da d n w h r r sp ns

n l , may a s p ss addr ss h h r ca qu sr a nsh p w n c ass ca and p ran c nd n ng n m c

rms.

DROSO HI A E ANO AS ER

caus s r a v y sma g n m and x r m y w dg n c m h d gy, h ru y,Drosoph la mela ogaste has n h

cus a n v and cr a v s ra gy n h s udy arn ng S ns ns ar us d d srup h c mp n n s arn ng n y n T

s nd g n s ha pr duc sp c c s ns n h arn ng r mappara us, and h n d n y h g n pr duc a c d y h sp np n a sp c c m cu ar c mp n n h arn ng r m m ry p(Qu nn 1984, Duda 1985). h va u h s appr ach s ha s us s a



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70 W & S EY

i r n m h l gy r m h hysi l gi al s u i s s rib in rs ns b ss s mm n b s s ns

h s m ns n ss v l ning n h y w s Ph rm a reg na h bl w y) by N ls n 197 ) li s n rm lly n hr b s is in s ns s g r a li h ir l gs. In N ls n s ria h m s im l s h CS) li h l g was air wi h sugar han r sul in i n r b s is nsi n h CS Sin

rigin l r , l arning has b n bs r wi h a n mb r ul n l vD aq ly l n v s

has b n ir wi h sh Q inn a 97 , D i 977, v s iQuinn 979) r a r a h an r ha h s r vi sly b n air wis g r T m l 1983). n a i n, i s l n av i a l ha w s

air wi h m hani al sha ing M nnS a 977) li s an ls l rn h n h l gs v sh Q nn 981)M s h g n n bi h m l analys s ss l

r i s n rs n h s ab li y l rn n r m mb r n ass ia i n wh h l a s s n a a h h

Qu nn l 97 ), r an r-s r s ass ia i n whi h l s signi n n s n a a r a r r m

983) Th r ur s ar s aigh rw r n a s ra gy ha v n g h s n m l h is, i s ll w

l iglass b s illumin by r s n lam . Sm ll rin rar us as r s l r sh a 90 V ) r h rsi ly mrim n s, an h sam ri s ar r wi h su r s a li ir

h gri ) in h a i i ly iv im n s. Th gri s ar wi h n s n h n la in h ub , s i s h h

b s r s m l an sly s i h n n n l rn n n s s . In s bs n s wi h n m

n ni n lyn wi h a T ma n rm l i s n a i h rar vi usly a r wi h sh an a r a h h ran r vi uair wi h su r s . Th a a ar usually r s n as h r i n v i ngr r hing h sh r sugar-ass ia r ing

i l, m nus h i n v i ing hing h n l h s ri

h r-sh l rn ng r g h s b n s is l m anh ar i n in h l arnin as . Mu a i ns in si X g n sr philan w a ar us l arning r m m ry i s. Th s in luun e (dn ) ruta aga (rut) a age ( a ) turnip (tur), rad sh (rad),an

amne a amnD 976 A v s- nQ nn 1979 Q nn l979, m l 1983). E h m n h s h ris i i inn / m m ry r m l b hn n i s sh w r iabl

l rnin wh n s wi hin 30 s ll win r inin bu m m y in


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I V R R AR I 7i s d cays rapidlyAmnals sh ws n rmal l arning ut a rrant m m ry.

An additi nal mutant is lat d y Wri ht ( 77) thDopa eca oxy a eD cmutat n has a d ci ncy in n ur transm tt r synth sis Li ingst n

T mp l 983 and als fails t l arn th d r sh ck ass ciat ns as w las d r suc s ass ciati ns n this cas n can us mild mutati ns andsh w that t s ac u s ti n that s l ck d rat s f f rg tt ng f thsu s qu nt m m ry ar n rmal mp l t al ). n th f ll wins ct ns w r y d scuss s m f th sali nt f atur s f ach f thmutants.

DuTh m m ry mutantDunce was rst is lat d y B n r and c ll agu s n

9 6. h s mutant has su s qu ntly n th f cus f int ns study.Duncewas in tiall th ught t ha practically n sh rt-t rm m m ry ut lat rp rim nts sh w d d nc f an n tial m m ry that d cays ry rap d

da 7 T mp l t al ). t app ars that th l c s f thng n is in chr m r 3 n th X chr m s m and it is th u ht that th sg n c d s f r an is ym f th cAMP-ph sph d st ras PdE ntw n ym s inD o oph lathat hydr ly cyclic AMP By rs t al Kau ar 98 Kig r 7 Kig rG lanty 77 Sh tw ll 983 . A 35 khigh-cAMP a nity w-cGMP a n ty f rm f th n ym s d f ct inth ncmutant M r r c rr lat ns am ng thnc n d s r duc dact ity f th n ym and r duc d m m ry ha n r p rt d Sh tw

983 .

R bRuta aga ut als a m m

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