Morse 1986 Aquacultural-Engineering

7/28/2019 Morse 1986 Aquacultural-Engineering

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Aquacultural E ngineering 5 i 1 9 8 6 ) 3 4 7 - 3 5 5

B i o t e c h n o l o g y in M a r i n e A q u a c u l tu r e *D aniel E. M orse

D e p a r t m e n t o f B i o l o N c a l S c i e n c e s a n d t h e M a r i n e S c i e n c e I n st it ut e. U n i v e r si ty o fC a l i f o r n i a . S a n t a B a r b a r a , C a l i f o r n i a 9 3 1 0 6 , U S A

A B S T R A C Tbztensification o f production, modernization o f technique, an dimpro vem ent of cuhivars are needed to increase the econo mic eJfi-ciency of marine aquacultztre industries. To help achieve such improve-ments, mo dern m ethod s o f biochem ical and genetic engineering can beadapted to control biological processes which intrinsically limit theefficiency attd yield o f production. Suc h applications no w have beendev elop ed/ 'o r impro ved production o f molhzscs. Cri t ical l ife-cyclestages o f several species o f abalones an d o ther comm ercially valuablemolhzscs w hich thus fa r have pro ved am ena ble to improved control bythese mea ns inch zde reproduction, larval settlement, metamorphosis,an d acceleration o f early growth.Analysis o f the physiological a nd m olecu lar m ech an ism s ~4zichcontrol reprodu ction in rno lhtsc s reveals" a prostaglandin-depend entregulation o f spawning in abalones an d certain other species. Spawningo f gravid adults can be ind uce d b y the addition o f prostaglandins to thesurrounding seawater, or, more reliably and inexpensively, by activa-tion o f the endo genotzs en z)wzatic synthesis o f prostagland in-relatedspaw ning triggers" in response to ad de d hyd rogen peroxide. Peroxideactivation o f the prostagland in-depettdent spaw ning reaction has beenfo u n d widely usefid fo r obtaining synchronous an d copious release o ff idly com petent gam etes in a large nu m be r o f species of abalones',oysters, sca llop s, mussels, clams, an d other vahtable molluscs; anu m be r o f these species ha d not been successf idly spawned by othermethods.Set t lement and m etamo rpho sis o f m an y molluscan species can bebrought utzd er similar biochem ical control M ost e j]~ciently an d

* P o r t i o n s o f t h is r e p o r t h a v e b e e n p u b l i s h e d p r e v i o u s l y (M o r s e , 1 9 8 4 a , d ).3 4 7Aqzmctdtural Engineering 0 1 4 4 - 8 6 0 9 / 8 6 / S 0 3 . 5 0 - E l se vie r A p p l i e d S c i e n c e

P u b l i s h e r s L t d , E n g l a n d , 1 9 8 6 . P r i n t e d i n G r e a t B r i ta i n


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348 D. E. Morseinexpens ive ly , the na tura l b ioc tz er rz ica l r equ i r em en t fo r indu c t ion o ft hese c r it ic a l d e v e l o p m e n t a l p r o c e ss e s c a n b e m e t b y p ro v i d i n g c o m p e -t e n t la r v 'a e w i t h u n i q u e a m i n o a c i d c o n s t i t u e n t s a s s o c i a t e d w i t h t h en a t u r a ll y r e q u i re d in d u c i n g m o l e c u l e s . T h u s , y - a m i n o b u ~ ' r i c a c i d( G A B A ) c a n b e u s e d sir np l~ , s a f e l y a n d i n e x p e n s i v e l y t o i n d u c e c o r n -p l et e a n d r a p i d l ar va l s e t tl e m e n t a n d m e t a m o r p h o s i s - - w i th m i n i m a lm o r t a l i O ' - - i n m a n y c o m m e r c i a l l y i m p o r t a n t a b a l o n e sp ec ie s. T h i sa n d s im i l a r n et ~ ro t ra n s m it te r -r e la t ed , a m i n o - a c i d - d e r i v e d c o m p o u n d sa r e p r o v i n g c o r n p a r a b l y e f f e c ti v e f o r t h e r e l i a b le i n d u c t i o n o f s e t t le m e n ta n d m e t a m o r p h o s i s i n a n u m b e r o f ' o t h e r v a h t a b l e r n o lh ls c at z sp ec ie s.R e c e n t a n a l y s e s o f t h e r e q u i r e m e n t s J a r r a p i d e a r l y / z l v e n i l e g r o w t hi n a b a l o n e s r e v ea l s a m e c h a t t i sn z o f g r o w t h - c o n t r o l w h i c h i s S ll sc ep t-i b l e t o e x o g e n o u s h o r m o n a l a c c e l e r a t i o n . G e n e t i c c l o n i n g a n d a m p l i -f ic a t io n oJ" r e c o m b i n a m - D N A t e m p l a te s f o r t h e e f f ic i e n t p r o d u c t i o n o fe s se n ti a l g r o w t h - r e g u la t i n g h o r m o n e s , a n d s u b s t a n c e s r e q u i re d f o re n h a n c e d n u t r i t i o n a l e ff ic i e n Q , a n d r e s i s ta n c e o f c u l ti v a rs , a r e n o w i np ro g re ss . G e n e t i c e n g i n e e r i n g f o r t /z e d i r e c t i m p r o v e m e n t o f c u l t iv a t e dspec ie s soo n w i l l be w i th in r each .

INTRODUCTIONM a n y c o m m e r c i a l a n d d e v e l o p i n g a q u a c u l t u r e i n d u s t r i e s f a c e s e r i o u se c o n o m i c i n e f f i c i e n c i e s a n d l o s s e s r e s u l t i n g f r o m i n s u f f i c i e n t c o n t r o lof r eproduc t ion , deve lopment , growth , and surv iva l o f cu l t ivars ;re l iance on ine f f ic ient , cap i ta l - in tens ive , energy- in tens ive and/orlabor - in tens ive technolog ie s : and re l iance on increas ing ly unavai lab lel a r g e b o d i e s o f u n p o l l u t e d w a t e r . T o i m p r o v e t h e e c o n o m i c e f f i c i e n c yand y ie ld o f mar ine aquacul ture indus tr ie s , and to adapt these indus -t r i e s t o p r e s e n t e c o n o m i c a n d e n v i r o n m e n t a l c o n s t r a i n t s , s i g n i f i c a n ti n t e n s i f i c a t i o n o f p r o d u c t i o n , m o d e r n i z a t i o n o f t e c h n o l o g y , a n di m p r o v e m e n t o f c u lt iv a rs ar e n e e d e d . R e c e n t l y d e v e l o p e d m e t h o d s o fb i o c h e m i c a l a n d g e n e t i c e n g i n e e r i n g c a n b e a d a p t e d f o r t h e i m p r o v e dcontro l o f those b io log ica l processe s which in tr ins ica l ly l imi t the e f f i -c i e n c y a n d y i e l d o f m a r i n e a q u a c u l t u r e p r o d u c t i o n , t o h e l p a c h i e v ethese required changes . As i l lus trated in recent re search and deve lop-m e n t w i t h a b a l o n e s a n d a n u m b e r o f o t h e r c o m m e r c i a l l y v a l u a b l em a r i n e m o l l u s c s , s u c h c r i t i c a l p r o c e s s e s i n c l u d i n g r e p r o d u c t i o n ,d e v e l o p m e n t , l a r v a l s e t t l e m e n t , m e t a m o r p h o s i s , a n d t h e a c c e l e r a t i o no f g r o w t h h a v e p r o v e d a m e n a b l e t o i m p r o v e d c o n t r o l b y t h e s e n e wm e t h o d s ( M o r s e , 1 9 8 4 a , b ; H o o k e r a n d M o r s e , 1 9 8 5 ) .


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B i o t e c h n o l o ~ " i n m a r i n e a q u a c u l t u r e 349I d e n t i f i c a t i o n o f t h e u n d e r l y i n g m o l e c u l a r m e c h a n i s m s c o n t r o l l i n g

r e p r o d u c t io n , d e v e l o p m e n t , m e t a m o r p h o s i s a n d g r o w t h i n a b a l o n e s a n do ther spec ies has revea led severa l e ssen t ia l p rocesses which can bec o n t r o l l e d e a s i ly a n d e f fi c ie n t ly b y p r o v i s i o n o f th e r e q u i r e d b i o c h e m i -c a l r e g u l a to r y s u b s t a n c e s o r t h e i r i n e x p e n s i v e n a t u r a l a n a l o g s ( M o r s ee t a l . , 1 9 7 7 a ; 1 9 7 9 a ) . T h e s e b i o c h e m i c a l m e t h o d s a r e re l ia b l e, s a fea n d c o n v e n i e n t , m a k i n g p o s s i b l e t h e l o w - c o s t i m p r o v e m e n t o f m a s s -p r o d u c t i o n a n d e f f ic i e n c y ( M o r s e e t a l . , 1 9 7 7 b , 1 9 7 8 . 1 9 7 9 b ; M o r s e ,1 9 8 4 a ) . S u c h m e t h o d s h a v e p r o v e d w i d e l y a p p l i c a b l e to t h e im p r o v e dproduc t ion o f a l a rge number o f spec ies o f aba lones , oys te r s , sca l lops ,musse l s , c lams and o ther va luab le mol luscs under cu l t iva t ion fo r fooda n d o t h e r p r o d u c t s w o r l d w i d e ( M o r s e , 1 9 8 4 a ) .O t h e r m e t h o d s f o r th e a c c e l e ra t i o n o f g r o w t h , i m p r o v e m e n t o ff o o d - c o n v e r s i o n e f fi c ie n c y , a n d t h e e n h a n c e m e n t o f t h e r e s is t a n c e o fc u l t i v a r s w h i c h a r e p r e s e n t l y u n d e r d e v e l o p m e n t i n v o l v e t h e u s e o fg e n e t i c e n g i n e e r i n g f o r t h e a m p l i f i e d p r o d u c t i o n o f h o r m o n e s a n dr e la t e d g r o w t h - r e g u l a t in g m o l e c u l e s, p r o d u c t i o n o f va c c in e s , m a n i p u -la t ion o f gene t ic con ten t , and c lona l p roduc t ion . These and s imi la rm e t h o d s , i n c l u d i n g t e c h n i q u e s f o r c o n t r o l l e d g e n e a m p l i f i c a t i o n a n dt ra n s f er , a r e u n d e r i n v e s ti g a ti o n f o r th e e n h a n c e m e n t o f p r o d u c t i o n o fa n u m b e r o f i m p o r t a n t m a r i n e p l a n t a n d a n i m a l sp e c ie s .

C O N T R O L O F R E P R O D U C T I O NA b s o l u t e c o n t r o l o f r e p r o d u c t i o n is o n e o f th e m o s t c e n t r a l a n d b a s i cprocesses requ i red fo r e f f ic ien t aquacu l tu re . Ana lyses con duc ted ino u r l a b o r a t o r y r e v e a le d t h a t r e p r o d u c t i o n in a b a l o n e s a n d a la r g en u m b e r o f o t h e r c o m m e r c i a l l y v a l u a b l e m o l l u s c s i s c o n t r o l l e d b yh o r m o n e s k n o w n a s p r o s t a g l a n d i n s ( M o r s e e t a l . , 1977 a, b ; 1978;Morse , 1984a) . In a number o f these spec ies , p ros tag land ins re leasedin the spawn of one an imal wi l l t r igger the spawning o f o the r g rav ida n i m a l s n e a r b y . T a k i n g a d v a n t a g e o f t h i s o b s e r v a t i o n , w e f o u n d t h a tw e c o u l d i n d u c e s p a w n i n g i n g r a v i d m a l e o r f e m a l e a b a l o n e s s i m p l yb y a d d i n g v e r y lo w c o n c e n t r a t i o n s o f p r o s t a g l a n d i n s to t h e s u r r o u n d -i n g s e a w a t e r ( M o r s e e t a l . , 1 9 7 7 a ). H o w e v e r , t h is p r o c e d u r e w a sd e p e n d e n t u p o n t h e u s e o f r a t h e r e x p e n s i v e h o r m o n e s , a n d p r o v e dless than per fect ly re l iable .In sea rch ing fo r a more re l i ab le and less expens ive method tha tcou ld be used fo r the con t ro l o f reproduc t ion in a l a rge number o f


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350 D . E . M o r s e

spec ie s , we fou nd t ha t t he enzym at i c syn thes i s o f p ros t ag l and ins in t hereproduct ive t i s sues of mol luscs (and cer ta in o ther spec ies ) wasna tu ra l l y dep end en t up on - - and l im i t ed by - - t he ra t e o f syn thes is o fm i n u t e a m o u n t s o f h y d r o g e n p e r o ~ d e ( M o rs e e t a l . , 1977 a , b). T hishyd rogen pe rox ide is mad e na tu ra ll y , i n the r ep roduc t i ve t is sues, bythe b iosynthe t ic enzyme which ca ta lyzes the ra te - l imi t ing reac t ion inthe b iosynthes i s of the pros taglandins ; the peroxide then i s used byth i s enzyme as an ob l i ga to ry o r r equ i r ed co - subs t r a t e i n t he p ros t a -g l and in b iosyn the t i c r eac ti on .Based up on t h is fi nd ing , we deve loped a co nven i en t and i nexpen-s ive me thod fo r r e l iab ly i nduc ing spawning , simply by add ing a sma l lam oun t o f hyd roge n pe rox ide t o t he s eawa te r su r roun d ing t he an ima l st o b e r e p r o d u c e d ( M o r s e e t a l . , 1977a , b ; 1978). Th i s m e tho d conve -nient ly and re l i ab ly ac t iva tes the synthes i s of the endogenous pros ta -g tandin spawning t r igger , and thence induces the animals to spawn.Cop ious spawning o f bo th ma les and f ema les i s i nduced synchro -nous ly . T he eggs and spe rm ob t a ine d by t h is me thod p rove fu ll ycompe ten t fo r no rma l f e r t i l i za t i on and deve lopment (Morse e t a l . ,1977b; 1978) .

T h i s m e t h o d f o r t h e d i re c t m o l e c u l a r c o n t r o l o v e r th e p r o d u c t i o nof the na tura l spa wn ing tr igger has pro ved to be rapid , re l iab le , inex-pens ive and wide ly appl icable to many spec ies ( inc luding a growingnum ber w hich cou ld no t r e l iab ly be spawn ed by o the r m e thods) . As aresul t, th is meth od has bee n wide ly ad op ted in research an d indus t r i a lapp l i ca ti ons , f o r imp roved con t ro l o f r ep rodu c t i on o f a la rge num berof spec ies of aba lones , oys ters , sca l lops , musse l s , c lams , and o thercom merc i a l l y va luab l e mol luscs unde r cu l ti va t ion in t he US andabroad (Morse , 1984a ) .C O N T R O L O F L A R V A L S E T T L E M E N T , M E T A M O R P H O S ISA N D E A R L Y P O S T L A R V A L S U R V IV A L

Ser ious econ om ic an d ope ra t i ona l ine f fi c ienc i e s and h igh los sestyp ica l ly a r e found i n con ven t i ona l aquacu l t u r e i ndus t ri e s r e su lt ingf r o m p o o r c o n t r o l o f p r o c e ss e s d e p e n d e n t u p o n l ar v al m e t a m o r p h o s i s(Morse , 1984a , b ; H oo ke r an d M orse , 1985). By ana lyz ing t he na tu ra lr equ i r emen t s fo r t he i ndu c t i on o f l a rva l s e t tl emen t and m e tam orph o-s i s in a number of aba lone spec ies , we found tha t these processes a res t r i c t l y dependen t upon spec i f i c b iochemica l subs t ances wh ich t he


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Biotechnolog3' in ma rine aquaculture 3 51l a rvae normal ly contac t only a t the sur faces of cer ta in a lgae in the i rn a t u ra l e n v i r o n m e n t ( M o r se e t a l . , 1979a ; Morse and Morse , 1984) .Th i s p roces s ensu re s t he subs t r a t e - spec i f i c r ec ru i tmen t o f aba lonelarvae to sui table h abi ta ts in coa stal waters . In cul t ivat ion , how ever , i ft he na tu ra l l y r equ i r ed i nduc ing molecu l e s a r e no t adequa t e lyprov ided, the l a rvae show very . po or success a t metam orph os i s , wi thsubse quen t ly resul ting very low surviva l and po or grow th as juveni lestyp ica ll y obse rved i n t r ad i t i ona l l ow- t echno logy aquacu l t u r e op e ra -t ions (Morse e t a l . , 1979a , b ; Morse , 1984a , b ; Hoo ke r and Morse ,1985).

Al thou gh t he na tu ra l l y r equ i r ed a lgae p roved t o be unsu i tab l e fo rd i r ec t u se i n aquacu l t u r e ope ra t i ons ( spec imens co l l ec t ed f rom thef ie ld f requent ly car ry many smal l preda tors , and the a lgae grow toos lowly for l a rge-sca le grow th o f preda tor - f ree sur faces), charac te r iza-t ion of these a lgae has made d i rec t b iochemica l cont ro l feas ib le(Morse e t a l . , 1979b) . From ana lyses of the na tura l ly requi red induc-ing molecules ava i lab le a t the a lga l sur face , a number of s imple andinexpens ive b iochemica l inducers capable of sa t i s fy ing the l a rva lrequ i rem ent was fou nd. M ost e f f i c ient ly and inexpens ive ly , the na tu ra lb iochemica l r equ i r emen t fo r i nduc t i on o f s e t t l emen t and me tamor -phos i s in aba lones can be met by provid ing the l a rvae in cul ture wi th7-am inobu ty r i c ac id (GA BA ) , o r s imi la r r ead il y ava i lab l e am ino ac idcons t i tuen t s a s soc i a t ed w i th t he na tu ra l l y r equ ir ed i nduc ing molecu l e s(Morse e t a l . , 1979a , b). Th us , G A B A can b e used s imply , safely , andinexpens ive ly t o i nduce com ple t e and r ap id l a rva l s e t tl emen t andm e t a m o r p h o s i s - w ith m i ni m al m o r t a l i t y - in m a n y c o m m e r cia llyimp or t an t aba lone spec ie s (Morse , 1984a ) .Th e con t r a s t i n re su lt s ob t a in ed by t he use o f G A B A and by cu lt i-va t ion in the absence of th i s na tura l ly requi red inducer i s dramat ic .W h e n G A B A is a d d e d in lo w c o n c e n t r a t io n t o c o m p e t e n t a b a l o n elarvae, vi r tual ly a l l of the larva e set t le , a t tac h to natu ral o r ar t i f ic ia lsubs t r a t e s p rov ided , and r ap id ly commence t he i r gene t i ca l l yp rogrammed me tamorphos i s t o t he j uven i l e fo rm. Fo l l owing t h i ssynch ronou s i nduc t i on o f me tamo rphos i s , t he r e su lt ing hea l thy j uve -ni les grow re la tive ly rapid ly wi th ve ry h igh surviva l . In co nt ras t , l a rvaec u l t i v a t e d i n t h e a b s e n c e o f G A B A o r o t h e r a d d e d i n d u c e r s h o w ave ry low p e rcen t age o f s e t tl emen t , r e t a rda t i on o r com ple t e f ai lu r e o fm e t a m o r p h o s i s , a n d s lo w a n d a s y n c h r o n o u s j u v e n il e d e v e l o p m e n tand g rowth , w i th ve ry h igh m or t a l i ty (M orse , 1984a , b ; Ho ok e r and


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352 D . E . M o rse

M orse , 1985) . Typ ica l ly , su rv iva l o f juven i le s one m on th a f te r me ta -m o r p h o s i s i n d u c e d w i t h G A B A i s o b s e r v e d t o b e i n t h e r a n g e o f9 0 - 1 0 0 %, w h e r e a s s u r v i v a l o f s i b l i n g s c u l t i v a t e d i n p a r a l l e l w i t h o u tG A B A is typ ica l ly ~< 1% (Morse , 1984 a) .I t s h o u l d b e n o t e d h e r e t h a t p r o v i s i o n o f G A B A o r s i m i l a r n e u r o -t r a n s m i t t e r - r e l a t e d , a m i n o - a c i d - d e r i v e d c o m p o u n d s ( s u c h a s D O P A )a lso is p rov ing use fu l fo r im prov ing the e f f i c iency o f induc t ion o fs e t t l e m e n t a n d m e t a m o r p h o s i s o f a n u m b e r o f o t h e r v a l u a b l e m o l l u s -c a n s p e c ie s ( in c l u d in g v a ri o u s o y s t e r s a n d c l a m s ) ( M o r s e e t a l ., 1 9 7 9 a ;M o r s e , 1 9 8 4 a ) .

A C C E L E R A T I O N O F G R O W T H , I M P R O V E M E N T O FN U T R I T I O N , A N D R E S I S T A N C E T H R O U G H G E N E T I CE N G I N E E R I N G

A m a j o r r e m a i n i n g l i m i t a ti o n t o t h e e f f i ci e n c y o f a q u a c u l t u r e f o r m a n yan imal spec ies i s a complex o f p rob lems re la ted to the i r r e la t ive ly lowf o o d - c o n v e r s i o n e f f i c i e n c y , w i t h c o n s e q u e n t h i g h c o s t s o f f e e d i n g ,re la t ive ly s low grow th an d sens i t iv i ty to t r aum a, s t re ss and d i sease( M o r s e , 1 9 8 4 a , b ; H o o k e r a n d M o r s e , 1 9 8 5 ). T h r o u g h t h e a b i li ty t oi n d u c e t h e s y n c h r o n o u s s t a r t o f j u v e n i l e g r o w t h u n d e r d e f i n e d c o n d i -t ions (as desc r ibe d above) , it has bec om e poss ib le to d i s sec t th i s p rob-l em b y a n a l y s is o f th e h o r m o n a l r e q u i r e m e n t s f o r o p t i m a l g r o w t h ,nu t r i t iona l e f fi c iency , and res i s tance o f aba lone s an d o the r mo l luscs incu l t iva t ion . P re l iminary s tud ies revea l tha t g rowth can be s ign i f i can t lya c c e l e r a t e d b y p r o v i s io n o f e x o g e n o u s i n s u l i n a n d r e l a te d p e p t i d eg r o w t h h o r m o n e s ( M o r se , 1 9 8 4 a , b ; H o o k e r a n d M o r s e , 1 9 8 5 ).

T h e s e s t u d ie s , a n d t h o s e o f o u r c o l le a g u e s i n E u r o p e , h a v e e st a-b l i s h e d t h a t a b a l o n e s a n d o t h e r m o l l u s c s p r o d u c e t h e i r o w n i n s u l i na n d g r o w t h h o r m o n e s . S u c h h o r m o n e s p r o v e t o b e e s s e n t i a l t o t h emol luscs fo r : ( a ) the e f f i c ien t up take and u t i l i za t ion o f nu t r i en t s f romfood , (b ) the rap id convers ion o f these nu t r i en t s in to essen t ia le n z y m e s , o t h e r p r o t e i n s , a n d g l y c o g e n (m e a t c o n s t i t u e n t s o f p r i n c i p a lnu t r i t ive , f l avor , and commerc ia l va lue ) ; ( c ) r ap id g rowth ; and (d )res i s tance to t r auma, d i sease , and s t ress under cu l t iva t ion cond i t ions .I t shou ld be poss ib le , then , to use these hormones to inc rease ther a te o f g ro w t h , t h e e c o n o m i c e f f ic i e n c y o f fe e d i n g a n d t h e n u t r i t i o n a land commerc ia l va lue o f the mea t o f she l l f i sh under cu l t iva t ion . I t i s


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Bio techno logy in ma rine aquacu l ture 3 5 3impor t an t t o no t e t ha t because t hese hormones a r e pep t i des andpro t e ins , t hey a r e na tu ra l l y and r ap id ly degrade d by enzymes , and n o ts tored in animal t i s sues as s te ro ids a re . Thus , in cont ras t to thes t e ro ids , pep t i de horm on es can be used sa fe ly to acce l e r a t e g rowth .However , indus t r i a l appl ica t ions of these f indings wi l l requi reinexpens ive supp li e s o f t hese h orm on es f rom hom ologo us t is sues (i.e .f rom the mol luscs t hemse lves ) . Because mol luscs make on ly minu t equan t i ti e s o f t hese h orm ones , t he neces sa W supp li e s can be p roduc edmo st e f f i c ient ly and inexpe ns ive ly by app l ica t ions of recent ly deve-loped gene t i c eng inee r ing t echno logy , us ing t he c loned mol luscanho rm on e genes as temp la tes . For th i s purp ose , ou r l abora tory ' hasbegun ex pe r im en ta l work on t he c lon ing and amp l i fi ca t ion o f t hemol luscan ( aba lone ) genes cod ing fo r g rowth-acce l e r a t i ng hormones(Morse , 1984a ) . S imi l a r ho rmones now a re be ing p roduced f rom thec loned genes o f hum ans and t e r re s t ri a l an ima l s , fo r use i n med ic ine .pha rm aceu t i ca l p rodu c t i on , and ag r icu l tu r e . T he re is eve ry reason toexpec t t ha t such t echn iques shou ld p rove equa l ly usefu l f o r improve -m ents in aqua cul ture as well.

F U T U R E P R O SP E C TSF r o m t h e c l o n e d g e n e - b a n k s n o w u n d e r c o n s t r u c t i o n f r o m m a r i n efood resource spec ies , s t i l l o ther usefu l genes and gene products canbe co nve n i en t l y s e l ec ted fo r t he ana lys is and improve d co n t ro l o fnu t r i t i on , g rowth , r e s i s t ance , r ep roduc t i on , and deve lopment . Moreadvanced appl ica t ions of genet ic engineer ing , inc luding the se lec t ivemo di f i ca ti on , amp l i fi ca t ion , and r e in t rodu c t i on i n to t he genom e o fspec i f i c genes for the d i rec t improvement of cu l t iva ted spec ies ofm ar ine p l an t s an d an ima ls , and t he p rodu c t i on o f vacc ines , soon w i llbe w i th in reach . In addi t ion , var ious s t ra tegies for the c lonal propa ga-t ion of ma r ine p lan t s and anima ls presently ' a re u nd er inves t iga t ion .T he newly deve loped b iochem ica l and gene t ic eng inee r ing t echno-log i e s desc r i bed he re fo r improved food p roduc t i on a l so a r e be ingapp l i ed t o t he ana lys i s and con t ro l o f mar ine fou l ing (Morse , 198 4


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354 D. E. MorseA C K N O W L E D G E M E N T S

T h e r e s e a r c h r e p o r t e d h e r e w a s s u p p o r t e d b y t he U S D e p a r t m e n t o fC o m m e r c e ( N O A A ) - U n i v e rs i ty o f C a l if o r n ia S ea G r a n t C o ll eg eP r o g r a m ( G r a n t s R / A - 4 3 a n d R / A - 5 1 ) , t h e U S N av y, O f f ic e o f N a v a lR e s e a r c h ( C o n t r a c t N o . N 0 0 0 1 4 - 8 0 - C - 0 3 1 0 ) , a n d c o n t r i b u t i o n s f r o mC h e v r o n U S A , In c ., a n d t h e A t l a n t ic R i c h f i el d F o u n d a t i o n .

R E F E R E N C E SH o o k e r , N . & M o r s e , D . E . (1 9 8 5 ). A b a l o n e : t h e e m e r g i n g d e v e l o p m e n t o fco m m er c ia l cu l t i v a t io n in t h e Un i t ed S ta t e s . I n : Crus tacean and Mol luscAqu acu l ture in the Uni ted S ta tes , e d s J . V . H u n e r & E . E . B r o w n , A V IP u b l is h e r s, W e s t p o r t, 3 6 5 - 4 1 3 .M o r s e , A . N . C . & M o r s e , D . E . ( 1 98 4 ). R e c r u i t m e n t a n d m e t a m o r p h o s i s o fHaliotis l a rv ae in d u ced b y m o lecu le s u n iq u e ly av a il ab le a t t h e su r f aces o fcrus tose red a lgae . J. Ex p. M ar. Biol. Eco l., 7 5 , 1 9 1 - 2 1 5 .M o r s e , D . E . ( 19 8 4 a ) . B i o c h e m i c a l a n d g e n e ti c e n ~ n e e r i n g f o r i m p r o v e dp r o d u c t i o n o f a b a l o n e s a n d o t h e r v a l u a b l e m o l l u s c s . A q u a cu l t u r e , 3 9 ,

2 6 3 - 8 2 .M o r se , D . E . ( 1 9 8 4 b ) . P r o sp ec t s f o r t h e Ca l i f o r n ia ab a lo n e r e so u r ce : r ecen td ev e lo p m en t s o f n ew t ech n o lo g ie s f o r aq u acu l tu r e an d co s t -e f f ect iv e seed -i n g f o r r e s t o r a t i o n a n d e n h a n c e m e n t o f c o m m e r c i a l a n d r e c r e a t i o n a lfisheries. In: Ocean S tud ies, ed . S . Han sc h , Ca l i f o r n ia Co as t a l C o m m iss io n ,San Fr an c i sco , 1 6 5 - 8 9 .Mo r se , D . E . ( 1 9 8 4 c ) . B io ch em ica l co n t r o l o f m ar in e f o u l in g . I n : MarineBiodeter iorat ion, ed s R . T ip p e r & J . Co s t lo w, Nav a l I n s t i t u t e P r e ss ,A n n a p o l i s , 1 3 4 - 4 0 .M o r se , D . E . ( 1 9 8 4 d ) . B io ch em ica l an d g en e t i c en g in ee r in g in m a r in e aq u a -c u lt ur e: t h e r o le o f m o d e r n b i o t e c h n o l o g y in t h e p r o d u c t i o n o f f o o d f r o mth e o cean s . O cea n s ' 8 3 , Mar. Tech. Soc. , San Francisco, Vol . 2 , pp.1 0 0 9 - 1 1 .M o r se , D . E . (1 9 8 6 ) . Ex te r n a l m o lecu la r s ig n a ls co n t r o l l in g r ep r o d u c t io n ,se t t l em en t , an d m e tam o r p h o s i s o f b en th i c m ar in e in v e r t eb r a t e s . In : Biologyo f Ben thic M arine Organisrns, ed s M. - F . Th o m p so n , R . Sa r o j in i & R.N a g a b h u s h a n a m , O x f o r d P r e ss , N e w D e lh i , 3 7 9 - 8 6 .Mo r se , D . E . . Du n can , H . , Ho o k e r , N . & Mo r se , A . ( 1 9 7 7 a ) . Hy d r o g enp e r o x id e in d u ces sp awn in g in m o l lu sc s , w i th ac t iv a t io n o f p r o s t ag lan d in -e n d o p e r o x i d e s y n t h e ta s e . Science, 1 9 6 , 2 9 8 - 3 0 0 .M o r se , D . E., Du n c an , H . , Ho o k e r , N . & Mo r se , A . ( 1 9 7 7 b ) . A n in ex p en s iv ec h e m i c a l m e t h o d f o r t h e c o n t r o l a n d s y n c h r o n o u s i n d u c t i o n o f s p a w n i n gan d r ep r o d u c t io n in m o l lu scan sp ec ie s im p o r t an t a s p r o t e in - r i ch f o o d


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Biotechnolo~, in marine aquaculture 355r e s o u rces . United Nat ions Food and Agricul tural Organizat ion FisheriesBulletin, 2 0 0 , 2 9 1 - 3 0 0 .

M o r s e , D . E ., H o o k e r , N . & M o r s e , A . ( 19 7 8 ). C h e m i c a l c o n t r o l o f r e p r o d u c -t i o n i n b i v a l v e an d g as t ro p o d mo l l u s c s , I i i : A n i n ex p en s i v e t ech n i q u e fo rm a r i c u l t u r e o f m a n y s p e ci e s. Proc. World Maricult. So c., 9 , 5 4 3 - 7 .M o r s e , D . E ., H o o k e r , N ., D u n c a n , H . & J e n s e n , L . , 19 7 9 a i. y - A m i n o b u t y r i cac i d , a n eu ro t r an s mi t t e r , i n d u ces p l an k t o n i c ab a l o n e l a rv ae t o s e t t l e an db e g in m e t a m o r p h o s i s . Science, 2 0 4 , 4 0 7 - ! 0 .Mo rs e , D . E . , H o o k e r , N . , J en s en , L . & D u n can , H . (1 9 7 9 b ) . I n d u c t i o n o fl arv al ab a l o n e s e t t li n g an d m e t am o rp h o s i s b y ; / - ami n o b u t y r i c ac i d an d itsco n g en e r s f ro m c ru s t o s e r ed a l g ae , I I : ap p l i ca t i o n s t o cu l t i v a t i o n , s eed -p ro d u c t i o n an d b i o as say s ; p r in c i p a l c au s es o f mo r t a l it y an d i n t e r f e r en ce .Proc. W orld Ma ricult. S oc., 1O , 8 1 -9 1 .

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Morse 1986 Aquacultural-Engineering