Carew Invertebrate and Learning

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<ul><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 1/53</p><p> 98 9 4 8y i h y l evie ll i h e e ve</p><p>INVERTEB ATE LEARNINGAND M MO Y:</p><p>From Behavior to Molecules</p><p>h a J Car w a d Chr t L Sah</p><p>ep me f ych y d y Y e ve y New 06520</p><p>N OD ON</p><p>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 </p><p>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</p><p>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</p><p>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</p><p>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 "</p><p>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</p><p>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</p><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 </p><p>k h 8 ) d hipp c mp cN ht et 8 L35</p><p>0 4 06X/86/0301 435$02.00</p><p>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</p><p> w w . a n n u al r e vi e w s . o r g</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1</p><p> . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 2/53</p><p>6 CA &amp; S LEY</p><p>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, </p><p>ech ic l d c cep l le el c ce r i g he ge er l e</p><p>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</p><p>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</p><p>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</p><p>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</p><p> v b at s</p><p>L N NG: P N PL ND P D G</p><p>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</p><p>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 </p><p>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 </p><p>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</p><p>pecie f im l .Le r i g p ych l gi dyi g er ebr e h e ide i ed m y </p><p>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 </p><p>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</p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1 . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 3/53</p><p>R BRA LE N NG 437</p><p>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</p><p>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</p><p>Nona o a ve Lea n no assoc at a refer ch ge i beh vi r re l f</p><p>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</p><p>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</p><p>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.</p><p>o a ve Lea n nsso at a gimplies e le r i f rel i s ip be ee r</p><p>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</p><p>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</p><p>is ss c i c m ify m y be v s D mjk r 82cki s ).</p><p>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</p><p>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</p><p>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</p><p>ll we hem esc pe re f rceme )</p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1</p><p> . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 4/53</p><p>438 CAREW &amp; SAHL Y</p><p>mpo a Pa nLe r cr c y epe e p he me erv ep r</p><p> he US he er m e v r S ) Depe c be e r e e re p e me re </p><p>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</p><p> 1966 G rc K e 966 Rev kyG rc 97 )</p><p> 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 </p><p>c F r r er c mpe h he e r e he rec p r f he CS US c c c</p><p>re p e re f rceme pe cSe on O e Con on n</p><p>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 </p><p> 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 , </p><p>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</p><p>B o k nA c c e per meby K m r ce phe me c ek ng</p><p>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</p><p>m c e m re w US e S S2-Ue e y we c e re p e b S S2 </p><p>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</p><p>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</p><p>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)</p><p>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 </p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1 . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 5/53</p><p>INVERTEBRATE EARN NG</p><p>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</p><p>y g v g o e gro p e ra m er o US pre e a o . h all gr</p><p>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 </p><p>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.</p><p>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.</p><p>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</p><p>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</p><p>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</p><p>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</p><p>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)</p><p>Con one mo ona Re pon e C Rhe ER proced re wa rod ced a a d rec mea re o</p><p>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</p><p>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 </p><p>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</p><p>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</p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1</p><p> . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 6/53</p><p>440 C W S LEY</p><p>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).</p><p> pr ding ri ov rvi o l rning prin ipl s nd p r digms</p><p>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</p><p>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</p><p>A IS M LLIF RA</p><p>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.</p><p> F F yin oc duM z a s ag s a b s t m v a</p><p>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 </p><p>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 </p><p> 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><p>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</p><p>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 </p><p> l nding on on olor nd n ding on no r ill pr r l ndolor o ding olor Op ng r 1931).</p><p>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</p><p>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</p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1 . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 7/53</p><p>R RA LE NING 441</p><p>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 </p><p>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</p><p>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</p><p>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</p><p>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</p><p>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</p><p>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</p><p>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)</p><p>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</p><p>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 </p><p>). 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</p><p>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</p><p>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</p><p>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</p><p>Annu .Rev .Neurosci .1986 .9:435-487 .Downloadedfrom www .annualreviews .org</p><p> b y C N R S - m u l</p><p> t i - s</p><p> i t e o n</p><p> 0 8 / 2 1 / 1 1</p><p> . F o r p e r s o n a l u s e o n</p><p> l y .</p></li><li><p>7/27/2019 Carew Invertebrate and Learning</p><p> 8/53</p><p>442 A W S LEY</p><p>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.</p><p>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</p><p>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</p><p>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</p><p>pai ed with the suga wate US E be1980 198 ,Va es hi1971,MasuhMe el19 2Me el et al19 4). olo -suga wa...</p></li></ul>