Garber s 1989

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Annu Rev Biocm. :-Cpyrght © by Anual Reews I A rghts rered

MOLECULAR BASIS OF

FERTZTON

David L. Garbers

Howard Hughes Mdical Institute, Dpartments of Pharacology and Molecuar

hysiology and Biophysics Vanderbilt University Medical Center Nashile Tennesee 37232-0295

CNTENT

PERSPETIVES AND SUMMARy............................... 719

SELECTIE TRANSORT OF SERMATOZOA................... 72ACTIATION OF MOTILIT............................................ 722

dntatn f tv Mu ...... ....................... ............ 723Identcation f Egg Peptide Genes. 724

Mhanm f tn. .. .. .. .. .. .. .. .... ......... .. .. .. .. .. .. .. .. .. . 76

HEMOARATlON................................................. 77dntatn f tv Mu .. .. .. .... .. .. .. .. .. ... .. .. ... .. . 7Mhnsm f An 78

RECEORS FOR EGG ETIDES ....................................... 78dntatn f ptr Mu.... .. ... .. .. .. .. .. .. .. .. .. .. .. .. .... .. .. .. .. .. . 728Cpri wt Sti Reeptr 730

guat f tr ctv....................................... 734

GAMETE ADHESION............................................. 734

dntcatn f ctv Mcu .. ... .. .. ... . .. .. .. .. .. . .. .... .. .. .. .. ... .. ... 734

INDTION O THE AROSOME REAON........................... 735dntatn f ctv Mcu .. .. .. . ... .. .. .. .. .. .. ... .. ... .. .. .. .. .. . 73Mhanm f tn.... ... ... ... . ... .... .... ........ ......... ...... .... ..... ..... .... 73

MEMBRANE REOGNITON AND EGG ATIATON.................... 73

SIS AND SUMMAY

he mecu eents tht ct in cdintin t dictte successfu fetiiztin ccu t mny diffeent ees incuding nim behir itse. Hee ewill concentrae on he molecular basis for he specificiy of ineracions

00-4154/89/070079$0200

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b y U n i v e r s i t y o f C o n n e c t i c u t o

n 0 8 / 0 1 / 1 3 . F o r p e r s o n a l u s e o n l

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20 GARBERS

btwn th gmts nd thir assciatd strcturs th spcificity tht inlrg msur prhibits friliati crss th spcis.

A mdl, in simplst f frms that summarzs knwn intrctins b

tn th amts is ivn in Fir 1. Hr spcific activatin f sprmmtility, ttrctin f sprmatza t th gg, adhsin f sprm clls t thgg inductin f an crsm ractin, mmbran fusin btwn th gmts, and subsqunt g ctivtin ar idntifid Evidnc xists t supprt ths spcific intrctins in a wid varity f nimals, nd xmpls willb discussd in dtail latr As dmnstratd in th mdl f Figur 1 ggsr gnrlly nvlpd in cllular matrics and/r dhrnt clls tht thprmatzn must frst ncuntr In much f th litratur th sits fspcific intrctin btwn th sprmatn nd th gg nd/r th cllulr

nd cllulr cmpnnts f th gg r nt clr fr givn spcisThrfr, whn w spk f mlculs assciatd ith th gg, th pssiblmananc f such lculs from clllr r acllular structurs surrundinth r from th , itslf, as wll as th dpsitin f ctiv mlculs tth ggssciatd strcturs by cls f th fml rprductiv tract ar llincludd

It is imprant t rli that friliatin may ccur in th bsnc f manyf th spcific vnts dscribd i Figur 1. Hwvr, frtilitin rtscrtainly wuld b xpctd t b lwr undr naturl cnditins spcilly

Fgur te o potena neracion tween speatooa an he egg (ncuingaear and euar atce ha uoundthe egg).

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. D o w n l o a d e d f r o m w w w . a n n u a l r e v i e w s . o r g

b y U n i v e r s i t y o f C o n n e c t i c u t o n 0 8 / 0 1 / 1 3 . F o r p e r s o n a l u s e o n l y .

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AI OF TILIZATION 721

whe stress is ivolved d shoud re etwee zero d the orl vlue.urret evidece uets specificity t the species evel for ech of theitectio depicted lthough speciiciy is oe ot bsoute but etive i

e of te cocetio of effecor oecue required to ect respoeHowever i soe cse specifict i soute d eve the hihest cocetrtios of eecto olecule will o iterct with receptor oecue ofother specie Epe of retive d bsolute specificity c e obseedwith sll peptides foud i the eggcoditioed edi of vious iltht itrct with cell sufce eceptos of spemtoo. I closely reltedpecies the specifici e sed o reltie potecies but i oredisttly relted species o crossrectio is visie eve t the hihestpeptide cocetrtio

The biocheicl d biooicl eposes of perm cels to specificefector olecue re eerl sil or idetcl Therefore it is resobeto ssue ht the ched ehvior of the petozoo i portt deerts pesre for the coevoluto of efector olcule d receptor idisites. he poteti coevolutio of effetor d receptor oecues ivovedwith fetilito rise the posiilty tht uttos t the er cell levelcould i soe cses edite specito itself or this to occur, uttioi efecto olecue egg) d i ecepto oecue spemtoo wouldoccur such tht fetiiztio etwee the utt e cells d spertozo

o eggs of te geel popuio would e redly reduced or ero butfertiliztio rte would be hh betwee te two utt The offpr ofthi ti the coud coceivly lso ot effectivel fetilize ecept withech othe hreby resutig i th se pheoeo s eogphic isotio

sed on sech o bcteil heoeceptors 1 d other receptorfilies 7 worki hotheis would e tht cel ufce recetor ofpetoo re hhl cosee th ctolsc dos but showcosidere ritio i the receptorbidi dois. uch odes couldprovide resoly siple eptios for wht ow ppers to be cosiderle diverit i the structure o effector oecue ut etetio of the bicbiooic esposes deostted i igure coss the species

Periet eviews o the oleculr sis of fetiiztio iclude hpio(8 Tier Vcquier 9 Vcquer 10 W 1 12 dGrers

SELECTIVE TRANSPORT OF SPERMATOZOA

Eculted spertoo ust oe e first tspored to the site of fertiiztio withi the fele. eective rspor withi the fele does ot ecessriy ivove specific olecules ssocited with the e ut y ivolve




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722 GARBERS

mcs asd in fic i a h im f ain as as caffc mcs asd by h s idc. In ais mammas, i is dcmnd ha smaa may sid in sicd gins f h

edci ac na ans h si f fiizain (8and ha sm miiy may b dcd ihin thes gins (18 Seas sggs ha h ans f smaa h si f fiizainccs cinciden ih he ime f ain, and ahgh h mchanism fans aciain is n fy ndsd h simain f idcamiiy ciiay miiy by facs ihin he fica id, and/ eas f sm chmaacans sm miiy simans cincidnwih he as f h egg, ae iky eanains T enia f sciesscific gain f ans has any n y bn sdid, ahgh

i sns n nia mechanism caab f ning diminishingcssscis fiizain

CTITION O MOTILITY

Miiy ae canes and/ incasd sm ciy ha bn din bh intbas and eebae in esnse eggassciacd meces19 20 Ofn, h, i has n bn ca whh n h siffecs a d a scific mce ind wih signaing annscifc ct g. a meabic sbsa. Rcny, a sbstancesen in smina asma f is hat stimats adenyae cycas aciyas ied and identied as bicbnate in (21 22); it as simaedsm miiy The ahs ha sggsd ha h bicabnae ffc nadenyae cycas is secific f h enzyme fnd in semaza. Thsimain cs f bicabna in d nmay b aibd is as a gna mabic sbsa ssiby is abiiy eeae inaca , in is cas a as, a mc ha cass gnaeffects in many differen ells �lso appars e secic eects in esazn n hf mst be cais ih inains f hmechanisms actin f meces tha mih be cnsided nnsifcLmuu poyphmus sematza eesen an am f cs ha ajacad and main in an immi sta ni ggcndiined mdia aeadded t te ces (23, 24) eimia ets hae sggested ha the actiemc in h egg media is a tide since ase tamen a aiayed ncia destys te actiity (24). n Orthops ee seaza aa t bcm immie ihin h aca mati y ca f hegg ni "aci mecs a eeasd cinciden ih fain f h

scnd a bdy sbsqeny, miiy smes and fiizain ccs25) An ame f miiy nhancemn, sciay nd cndiins fsighy acidic etace , i eesened by sea chin seaza,h gg-cndiind mdia can inceas sm siain as and miiy (26 27)

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dea Aive Mees

BASIS OF ERTILIZATION 23

Smll pepdes he ben sled frm eggs h cn smule spermmetblsm nd mtly unde pprprite cndtns n se uchns 2-32)

he t peptides studied in getest detil e spect (lyPheAspLeuAsnly-lylyVlly) slted fm Strongylocentrotus purpuratus Hemicentrotus pulcherrmus, nd esct (ysVl-hly-AlPly-ysVl-lyly-lyAg-euH btied fm Abaca unctulata eggcnditned med. Resct nd spect d n crssect deecbly hspemt f the speces cntnng the ppste peptd een hen eryhgh cncentns e used. he specifcty ppes t be dctted by thepimy stuctue f the cbxyl til f the peptide. nsdeble substitutns pssible n the Hteminl prtin f spect ith etentn f bilgcl

ctty but deletn f the Otermnl ly Vlly esults n lge ttl lsses f esptn-stmultng cty 33) Ms chemclly syntheszed ngues he hd n el ecesed ptency ele t spect,but ne nlgue (lyPhe-AspLeuSelylylyVlP ppes t be00 times me ptent .

mu 3) ls hs studied the stuctuectity eltinships f lyPhe-Aspeu-Selyly-lyVl-ly n egg peptde frm A. rassspina.

Elmintn f the cbxyl-temnl ly educed cy t /3000 f thentie peptde hees deletin f the minteinl ly educed ctityby nly 0 y culd eplce Phe thut lss f ctity. he eplcementf Phe by Leu nleucine Al, ly, P, in cntst cnsdebly educedctiity . Vline ls ppeed mpnt f ctity nd t subseuently ssuggested tht hydphbc esdues t pstns 2 4 nd 9 ee mprntf ptiml ctity.

he mst p, stuctuectiity studies he cncented n theeltinship f stuctue t espitin stimultin; f numbe f spectnlgues, espitnstimulting ctity hs cncided th cyclc

nucletideeleting ctiity30)

Shimmu

bes36)

peped us nlgues f esc hee nd fund th elte ptences ieddependent n the physlgcl pmete mesued. Mdfctn f theOteminl leucneH f esct did nt lte blgcl ciity, butsubstitutin f the t cystenyl esidues by Se y methyltin f thecystenyl edues esulted in diegent eltie ptences dependent nhethe espitin tes cyclic nucletide cncentins ee mesued.

he stuctues f ius the spemstmultng peptdes e n nn339) se f these esemble the suctue f spect esct nd hes

d n (ble )Suzui et l 40) he ttempted t study the txnmicl disbutin thepeptdes n he se uchn by testin the cude eggcnditined medi f pticul speces n the spetz f us the species hei esultssuggest tht eggcnditined med f speces ihin the sme de ill




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24 GARBER

Stuctue se the peptides that ae kw t stiulate pe eais ad tlity ud i the ecdtied eda vaus se uchs

eptde ae

Speac

Resac

eptide stctue

Gy-Phe-Asp-LeuAsGly-Gy-Gy-Val-GyGy-Phe-Asp-LeuSeGy-Gy-Gy-Va-GlyGly-Phe-Asp-LeuhGly-Gy-Gly-Va-GlyGy-Phe-Asp-Leu-h-GyGy-Gy-ValGGly-he-Ala-LeuGlyGly-Gly-Gly-Val-GyGy-PheSe-Leu-As-Gly-Gy-Gly-Va-SSePheAlaLuGlyGyGlyGlyValGlyGy-Phe-Se-Leu-Se-GlySe-GlyVal-AspGly-Phe-Se-Leu-SeGlySe -Gy-Va-GlyAsp-Se-Asp-Se-Ala-G-As-Leu-Ie-GyAsp-Se-Asp-Se-AaHLeu-IeGyys-Valh -GlyAa-P-Glyys-Val-Gy-GyGly-Ag-LeuH2

simulae he spermaozoa across species ies; his oes o imply ha hesrcures of he pepies are he same bu ha he pepie sucures aresimilar eough o cross-reac wih he same recepors L ptus a S

purpurtu fall wihi he same orer for eample a hey boh coaiserac or speraclike molecules bu he srucures of he peies are

iere (37.

deia Egg Peide Gees

Oe of he ieresig observaios me by Suuki a coworkers (29 3238 ) was ha varous pepies wi similar srucures coul be isoaefrom he eggcoiioe meia of e same species Sice coiioe meiawere isolae fom he es of may iviuals i as o o heherhe iere pepie sucures reece cosiues from a sigle sea urchi

or wheher iivial sea urchis posssse oe pepie a he vaiabiliy isrcures reflce iiviual variaio i he populaio. S Ramarao D Burks a D L Garbers (persol commuicaio) have ow isolaecDNA cloes from a ovaria cDN libray ha ecoes for sperac asperaclie sucures. Two iffere cles wih ise sizes of a bhave bee isolae; he ope reaig rame of he loger iser is show iigure 2 Eigh sperac or speraclike oeial ecapeies are ecoe fori oe regio each separae by a sile lysie resiue our of he eiesare serac a four coai srucures ha ossess a Gly-Gly-Gly-Val-Glycarboylemius suggesive ha hey herefore have moiliy-simulaigproperies (see above). Aoher ecapepie Gly-hr-MePro-Thr-Gly-AlaGly-Val-As wihi he same regio of he mRNA as well as a ecaeie

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DDUCE AMINO A SE�NC OF TH PERAC PCUR

et

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Gly L -ProVal-lieS GyGnAp-GlLt

TyTh-e-Ja-pSeSeAp-euGl-Ge-AlaHis t

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e-Se-Po-eisleSe-eu-SSeeuGI uSer-Ja rp-n nu-enle

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SeGl-Gl-py-Se-e-Po-Gl-eu-Se-leo L -pV- LSe-

e-Se-C-PoL�-y-SePo- Li-yPaaeGy-eSps-

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yVa-G-Va-G n-ps-e-an-AaL -Gu-eu-r-G-Gy L eV-

Se-Gl-e-le-Met-y-:- 1-ep-HpeU-Aa-Ser-VSer-L!-e-eu

*Se-s-T -e-p-Ty-l-S--e-Se-See Se VlVal-y- Gn-p

G n-T- AlPo-Se-His-Po-Met-p-GluSer-TyMe tph-ro LeuSr-et-l- e-

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Figure 2 Te dedce amino' acd seece of te peract p ecrsr. T he pedicted amno acds ae ed on the DA sequence otaned f m an olaed

2 .3 cDNA ne fo a ea cn ovary DA rary Two p ossible ntaion ses ( meonne) e maked wih n oele. e ine edue are

sared and the nne potenial decaepides tat are contious wit ec te ae b Oed. ne ot oea decap eptide exists .

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e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2 . D o w n l o a d e d f r o m w w w

. a n n u a l r e v i e w s . o r g

b y U n i v

e r s i t y o f C o n n e c t i c u t o n 0 8 / 0 1 / 1 3 . F o r p e r s o n a l

u s e o n l y .

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2 GARBERS

ream Ile-ApHAp-Tr LeAlaSer -Vl-Ser al eaaed by ngle lyne rede a ye be eed fr blgal ay Te erlaed lne naned w addnal eralke rre: Ser-e-A

LeAn-GlyGly-GlyValGly and GlyeSer-Le-Tr-Gly-Gly-GlyVal Gly Hybrdzan f e DA lne w mRA from oe eeled wmajr RA ee of and . kb

Te ee for pert re o be olted, ll llo tedetemto of ete o ot tee e oeved eo of te ee ttet o pee Te tutue of te RA, oee, e teetqueto Se multple pet petke tutue o t teme RA t te peue t et fdelt mutple ope of teme boo te peptde? le pert moleule eoded t

oe mRA moleue ot uffet? ee e ote potet peptde eoded b te RA. Do te e futo? e proe of te pedtedrerr al wll be f nere. t be elaed a rynlkeenzyme fllwed by arbxypedae Blke ay wld rel n arrae reng Fnally were re ee ede yneed? relmnarydt of D Burk d D. L rbe perol ommuto) uet tta rerr mlele fr rea fnd wn e egg, ggeng yne wn e egg

Meas A

Te peptde ue ube of el boem epoe tt eembletoe epoted omt ell tt repod to ot fto ormoe, dote erometl Tet eleto of l AP, lP, d of tellul e bee eported 0 4 4 A potoex r and e nraelllar p nreae 41 4 Alg etempol emet of te boeml eet et to be etbed, teeleto of tellul p ppe to be te pm ue of te eedmty a meabsm. Hasbrugh Garbers a estraett Moe A ould ue te me eet o pem motlt te epeptde d Repke be 43 lte oed tt ek d d ekbe lteed motlt me otet t te pote tt telul p epeeted te pm eulto of motlt. Te ddto of8broo P but ot of l P yl AP, r var ernlede al an mlae ea rn erm mly ; e mlanb 8-brmyl P m repreet propert of t eotde n

elted to truturl mrt to l P, e Smomur Garber(36) demntraed a elean of perm reprto rte oud be oberedn e abene f deeable e nraelllar l P.




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CHEMOATTRACTION

deiai Aive Mees

BAI OF FRTILIZATION

Sperm chemotas has been reported n varous phla and has been demonstrated n man derent speces (45 across both the anmal and plantkndom. he frst and stll onl dentfed substance from anmal esestablshed as a chemoattractant s the peptde resact descrbed n the prevoussecton. he peptde was solated based on ts ablt to stmulate spermrespraon rates and was subsequentl shown to be a potent attractant (46).Mller (45) summares the chemcal characterstcs of attractants a numberof whch stronl resemble peptdes from varous anmals. Further researchon a starfsh attractant has suested t to be a peptde althouh ts prmarstructure s not known (47) Resact as been tested on the spermatooa of S purpuratus and L pctus as well as sarfsh spermatooa and t fals to attractthese speatooa there s also no detectable specfc bndn of radolabeledresact to spematooa from these anmals Consstent wth these observatonsMller (45) has compled etensve data demonstratn that speces specfctof sperm attracton s a common phenomenon he eperments to provdesuch nformaton have nvolved the use of rue or partall purifed etractsfrom es tested on the spermatooa of the same or of derent speces

Althouh the structures of the actve molecules reman to be determned, theepermena desn clearl establshed speces specfct to the attractantresponse

Reports aso est that spermatooa are attracted to peptdes of the fMeteuhe faml (48 49) In one case ths has been repoted for bovne and nthe other case for human spermatooa Snce pror to these reports chemotashad not been successfull demonstrated n mammalan spermatooa theresults were at frst ectn and suested that a molecule related to thspeptde faml mht be secreted b mammalan es However Mller (50)

subsequent demonstrated that the work of bal et al (49) contaned methodoloal errors and the rpo of Gness et al (48) has e to be confrmedFollown the apparent postve chemoattractant effects wh human spemcells Gness et al (5 ) reported the specfc bndn of Seteuhe tohuman spermatooa Appromatel 60000 receptors/cell were estmatedAttempts to measure specfc bndn of SetLeuhe to bovne orhuman spermaooa n a leas one oe laborato however have no beensuccessful (S. E Domno D. L Garbers personal ommunaon nemaor conce wth the prevous studes s the possble contamnaton of

spemaoo preparaos b neurophls or neurophl membranes t nowsees puden to reu o e eras eodoned meda folularud ec o determne wheher or not spem chemotas n he mammal




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728 ARBER

ccus. Sudies e penilly me cmpliced in he mmml ecuse he pcess cpciin nd ecuse he nml sge spem cells incein egins he femle epducive c pi nsp he se f

felzn; emns pssle h chemtcn espnses f mmmlspemz ll n e seved unless he cell pceeds hugh hesenml physilgicl evens

Meais Ai

The mechnsm y hch gdens esc e deeced y he spem cellemns e deemned. Heve s knn h C n he ecellulmedum is equied n de cells espnd he pepde gden (46)Since C is n equied he indng he pepides he simul

in espiin es i uld ppe h C mvemen m excellul incellul cmpmens epesens n impn signl spem cell enin hs een demnsed n c h incellulC cncenins ncese in nsien mnne in espnse spec es nd h such elevins equie he pesence excellul C+ 42A he h C+ onentaton may led hange n he aymmety flgell ending in espnse esc hs een ppsed 2 nd culdexplin he mechnism y hch he cell mves n pepide gdien. hesignl C milizin emns e clied lhugh cyclc AMnd/ cyclc GM e penl ddes n h cses hee s pevdence n he cells h hese nucledes cn egule n chnnels5358)

Resech f ee Ges 5) demnsed h he memne penil spemz culd e eguled y spec esc, nd Lee 60) lesuggesed invlvemen GTinding eguly pein in ppsedspe eep/K hnnel cupled nein The mdel Lee 60 iscive snce gunne nuclede eguly pens) e knn egu

ae K+ channe in oher e 61, 62 Th model wod sggest theegulin he +JH nipe y pimy eec n K chnnels,since ee 63) hs peviusly demnsed he exisence memne penildependen /H exchnge.

RECEPTORS OR EGG PEPTDES

Identcation of Receptor Molecules

Alhugh n -leled Bln-Hune dduc f spec s used in el

sudies f he spec ecep 33) he lck f ee min gup negedpenil css-linking sudes ideny he ecep Theee n nlgue(Gy-Gy-GyGy-y-ApLeuAn-Gy-Gy-Gy-Va-Gy) wa ynthezed

h ened epaton-tmuatng cv euvlen sec (64 he

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BAI OF FTILIZATION 79

aalu a cp Ial BlHur prac fr rcpr quval pc prac T GGG[]pac aralal a uqul crlk appar rcpr

ucl ura (). T appar rcpr a t a aglcp a a cuar g f 000 (u clufa ruc cn). T pcfc f aca f pp p a r cp pr vau prac aalu a falur f ralal pp cvall praza fr pc a crac pac. t c-kg xp g ccra fGGG[]prac cul u a rf a laff cpr a av c a p c f rcpr

lcul a ar capa f g cvall cupl GGG[]prac cau f a lack f car fucal up a . T v p a cc af r crk p rcp a pal la f cDNA cl fr pr fll xp f pr cuur cl. L Dag a 5 av aag purf cr-k p a a ac quc fr p a a cDN c f cDN lrar f p cDNAc ca a k r a la a z A

lat f f a A p a fra cag a ac a ca h . T prc p cta aaa a av rc c ru a fur rpaquc f appral 0 a ac puav rcpr l- lppr a r cpr rpc (8

I a pr f cllular a uprg fal ca arpa quc 9

T a apprac c av u f apparac rcpr 70 A aalu f rac (Gl-GGTrGl-C-Val

TrG-aPr-GC-ValGl-GlGl-ALuNH ) a ccalz a p a rpraula arcpr- acv a ac T raa aau a u c-k pr u a f a ajr aa pra a lcuar g f 77 000 ajr aacv a a pr a appa cua g f 0000 Sprac fa cp Ial aalgu clk p a raacvac ffcv cp aacv crlk pr a uqul f a z uala cca 70 t

appar rac cpr a ac quc fr pr ra a a cDNA c a uu a Th prca ac quc f guala ccla a p a fa f




y .

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730 GARR

986 amino acids with a 21amino-acid sgnal peptide and a sngle transmembrane domain separating the protein into 478 extracelllar and 459 ntracelllar amno acds. The enzyme was shown to possess signifcant identityn a carboxyl domain to the catalytic domans of all protein kinases, thereforemaking it a member of the protein kinase famly.

After the isolaton of the cDNA clone from A puncuaa spermaozoa, acDNA clone from a S purpuraus cDNA library was isolated and seqenced(72). A comparison of the predicted amino acid seqences between the twosea rchin clones is shown in Figre 3. igh conservation of amino acds isobserved, particlarly in the cytoplasmic domain that is homologos toprotein kinases. Greater variability in the amino termnal domain is apparent

and in the carboxyltail of the protein, the S purpuraus seqence completelydevates from that of A puncuaa

The findng of two apparent egg peptide receptor molecles (for speract andresact), whose strctres showed no high degree of simlarity, was nexpected. Since the biochemical responses of the sperm cells to the varospeptdes appear to be simlar or identical, the simplest receptor model woldbe lke that described for the bacterial chemoreceptors ( 13) where theaminoterminal domains are highly variable and the carboxyl domains remain

highly conserved. The conservation of the cytoplasmic domains of the peptidereceptors bt variation of the extracelllar, binding domain cold easilyaccont for the diversity of peptide specificity bt retention of the samebological responses. A possible explanation of the above receptor data woldbe that the 77 ,dalton protein and ganylate cyclase are actally in closeapposition in the membrane and possibly even sbnits of the fnctionalreceptor. Since crosslinking stdes relied on disccinimidyl sberate as thecopling reagent, t remains possible that either protein cold have been

crosslnked dependent on the proimity of the aminotermins of the peptideand a fnctional amino grop in one of the two proteins; the specifictyexpected for the receptor wold still be observed Unpblished data of L Dangott sggest that a speract receptorlike protein exists n A puncuaa;

this evidence is based on the isolation of positive-hybridizing cDNA clonesfrom a testis cDNA ibrary . Since it is known that speract does not bnd to A puncuaa spermatozoa, the homologos proten mst be modfied in theligand-binding domain if it, in fact represents the receptor. Wth cDNA

clones avalable for both apparent receptors t will now be possible todetermine whether or not both molecles are in fact cell srface receptorswhether they form sbnits of a receptor, or whether one of the protens ismerely in close apposition to the other.

Comparion with Somati Cell Reeptor

At abot the same time stdies demonstrated the cross-linking of resact toganylate cyclase of spermatozoa (70), reports appeared sggesting that aial

A n n u . R e v . B i o c

h e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2 . D o w n

l o a d e d f r o m w w w . a n n u a l r e v i e

w s . o r g


7/18/2019 Garber s 1989


S LFLFFM-RDFNPINEDRKIHVGLLDGLGFPLGPLSLQDSNLNGFDVQFTBCDN (00A L LV R BY V L P L I M S N NS SA F QY N M H Y N A

S AVSDGFVGVGPYEGLASNIDYVDENPVSDKSIYPFLRPPSIQVVEAIIQRYELDQVSVENIKFND (201A L F A N EF DS L O D V EQ (201

S ERDYELEEYYAGFDYEDPFSEQRKERIFGDASDLRQFLDEGIDSGDYVIGADLVRSQDYSDDSENQI 301A M EEW PDE D K SG S V N A V l E D 302

SP NPDYARFKNREYRSDNDLEKSVVGAVLKRNWDRFSVDNLDAPFNGELELRADFASYDATLLELDRHDIGEE (402)A P QA E D E M L E RSQA HY A E K D M E K A Q M SQ (403)

S SLLNSSKDFYQFDENGDGVPYLLLIPPGDAKSYPGFNRENFEEALDEDNVLKPNREPPLDMPGFBG (502A N F S AD S R V MP P V A S H S -N P D N D V 499)

S LALYGASPFLGFFYAYELDSLSEVQNSFSSAISVISNAEKIFAGRGVCHV 603)APG G A I GL YY K RESE S L (600

SP NDRAREKDONPFGADRPHSIYAGSQDENDDKLDSSSADLGLHSSEKSHGHLKSSNCDR 704)A M A L (701

S QDYGLNEQDVDGDAAQWSEQEGSMPAGSDYSFALEYSRQEPHENELADIGRVKSGEVPPRPLNA (805)A E E K A EGK PG K S M E A SK V 80)

S NPDLSAAPEDPADRPNAPLQKGLKPNLDARYNNLEELVDERQEQEKKEQLLPPSISQLKGA 906AP E V E A S (903)

S EESFFSDVGFALSSQVVNLNYLFDASNYDYKEGDAYVSGLPLRNGRHAASAHHLLESGFIVPHKPEL 007)A D HS SFSSQPVSH LS FAQSSA SHSSALSS* (986)

ur mprs f the ece mn seqee f he uttve gylte cycse rm uult (AP) S purpurtu SP he .unu DA ce ws iste escre Ref th ce ws he se s a pe se the S. pururtu cA ce he precemn c sequece f S purpurtu re shwn n thse f untut e given y where ffeen

A n n u . R e v . B

i o c h e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2 . D o w n l o a d e d f r o m w w w . a n n u a l r e v i e w s . o r g

b y U

n i v e r s i t y o f C o n n e c t i c u t o n 0 8 / 0 1 / 1 3 . F o r p e r s o

n a l u s e o n l y .

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7 GARBRS

narirec pepdes also cold bind o ganylae cyclase (776. In bohcases he pepdes were shown o civae ganylae cyclase acviy ofmembrane preparaions (7780 or of deergensolbilized enzyme pre

pons 8, 82) hese esls no only sggesed he consevon of cellsrfce ecepo wih enzymaic civiy n gem cells and somic cells bhe conservaion of hs signaling sysem from echinodems hrogh mammls.

The dedced mino acid seqences o mammalian membrane orms ofgnylae cyclase have now been obained (8 84, and a lowmoleclarwegh rial narirec pepide (AN recepor h does no possess ganylae cyclase acivy also has been cloned (85). The sea rchin sperm gnylae cyclases show some ideny o he low-moleclarwegh AN recepor

n he presmed pepide-binding domain whereas he mammlin embranefors re pproximely 33% idencl wh h reepor in he pesmedexracellla domin I is now known ha gnyle cyclase binds AN andherefore seves s ell sfe ecepo (8, 84. Of consderble neeshs been he compson of he membne forms of gnyle cyclse o oneo he sbns o he solble form of ganyle cyclse from bovine lngOne of he sbnis of he bovine lng solble om o gnyle cyclse hsbeen cloned by Koeslng e al (86 A shown n Figre 4, he clone rom S

purpurtus shows a hgh degree of deniy o he bovine lng solble form inhe crboxyl egon he enzyme of S purpurtus n fac esembles hesolble sbni wihin his region more so hn he membrane form rom A

punu (71) ince S purpurus is believed o have evolved mch laerhn A puntut he similriy in he carboxyl regon o ganylae cyclse

S GC 379

PGC 82

L RA

E Df

dKTlo

Tf h

P

f

N

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Q K

�

T

Q

H

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S Q

S CPGC

S GCP

406

R H K R P

PA K R D899I

K G IAL

P F E

4 C S K HAS G EGA H K

9 L SAAS T P Q

N[b

I

L

f

ISFD

I

lv N L L N D L YV V N L L N D L

S C 46PGC 949 i

S R H� �

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S C 511PGC 1000

P C I HlR fl H dlL r

IA

Q

L R N G 0 R�G QUAS TH

H L LS

VK F I

Q D S v Q I rl

-H

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SC

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A S NLA L RH

gure Comparon of a region o the purpurau (PGC) guaylate cycase amo acdeqene wt tat o te obe om o gnle e om ovne lng (SGC) 86).Ide ede e bxe

A n n u .

R e v . B i o c h e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2



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BAI OF FTIIZATIO 733

fm S purpurtus ad ie lug may idicae e cseai f eseesides i geea i species a aose ae S purpuratus is is suped y oseai a e mammalia memae fm f guaylaecycase as displays a ig degee f ami acid ideiy e sluleeyme s i is same ego (8 84)

I addi e memae fms of guayae cyclase coai a egioomologos o e poei kiases (7 83 84) d eefoe ey esemblei pa e cell suface ecepos of e poei yosie kiase family (87).ey als semle is family y ie f pssessig e pediced asmemae dmai A scemaic model f e gaylae cycase family eis so ige 5 Based e seqece aiailiy of e ezymebeee sea uci species i also ca e pediced a e guaylae cyclasefamiy il cde oe memes Aog ese lies i old e ecaleda gaylae cyclase podces cyclic AM a a lo ae elaie o cycicGM i some cases ee aes of cycic AM faio aeappoaced 0% f e aes of cyclic GM fomai (88 eefealug e ae o siig similaiies eee e aceial yeasadeylae cyclases ad e spem guaylae cycase (7 7) i sequecedaa o addioal adeylae ad gayae cyclases ad sbseqe closeexamiaio esiced coseed egios may ecome appae ome daaalei slig i fac as ee eped sugges egios of ideiy eee

e yeas adeyl ae cycase ad e mammalia guaylae cycase 8

RCYC PDGR PCR PGC PGC

r� +

-6

OLG PSMMMBR

g Membrs of th guanylat cyclas aiy. snd ar h ddcd prons hr b meme orm o gye cyce 83) (RCYC, DGF eceor DGFR (136),

-mch ANP co ANCR (85 A. puculaa ye yce (APGC S pupuatu gye cyce (GC h bovn soub orm o guanyacycle oLGC (86 Rego o can ey bewee he membae fo o ganylecyce h he e boxe a he he me




y .

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73 GARBER

Regulai f Reepr AiviGunylte cyclse exsts s phosphopoten n spetozo 89 90) nd

when membnes e peped such tht the enzyme emns n phosphoylted stte the egg peptdes mkedly ctvte the enzyme 77) The ctvton s rnsent howeve nd concdent wth decese n enzyme ctvty loss of phosphte occus 77. In the ntct cell elevtons of cyclc GMPcused by egg peptde ddton lso e tnsent nd s wth membnes pd dephosphorylton of gunylte cyclse concdes wth subseuent deceses n cellul cyclc GMP 89 90). In the bsence of peptde spegunylte cyclse hs been found to contn up to 7 mol phosphte/molenzyme ll on sene esdues 9 9 fte ddton of esct only 3 molphosphte/mol enzyme emn Theefoe the egg peptde ecepto:gunyltecyclse coupled ecton ppes to be desenstzed by dephosphoylton sopposed to desenstzton by phosphorylton s hs been epoed n mnyothe ecepto models (, 4

GAMT ADHSION

deiai f Aive Mlecules

In the se uchn the spetozoon fst encountes n cellul stuctueclled the jelly cot; fte penetton of ths mtel the spe cell ntectswth the vtellne lye po to ts fuson wth the plsm membne o theegg. Fo the gmetes to dhee to ech othe upon collson s dstnctdvntge nd n fct spe cells dhee to the jelly cot of eggs. One of themedtos of ths dheson ppers to be fucose sulfte polyme FSG fofucose sulfte glycoconjugte 0 tht lso nduces n cosome ecton9597. FSG s hghly sulfted polyme nd when dded to suspensons of

ethe lve o ded se uchn spetozo cuses mked gglutnton. seems lkely tht the lge FSG molecule contns multple ecognton stesnd theefoe s ble to bnd to multple spetozo esultng n gglutnton. The hghly chged ntue of FSG howeve leds to eltvely lowspecfcty n tes of mny potens nd cells dheng to the jelly cot

Specesspecc bndng tht demonsttes hgh fdely s obtned t thelevel of the vtellne envelope the stctue of one of the ntectve moleculess now known nd s nmed bndn 98) t s mjo poten of the cosoml

pocess nd hs been cloned nd the mno cd seuence deduced 99)ndn s pesent t n ntcellul ste nd theefoe medtes specesspecc bndng followng the cosome ecton. Its sructue would beexpected to vry between the speces nd some pelmnry dt n suppor ofths suggeston hve been pesented 99 . Bndn s syntheszed s 5 000-dlton pecuso molecule tht s subseuenly pocessed to yeld 24000-don mtue proten n the se uchn S pupuau . The synthess ofthe proten ppers specfc to the tests As ponted out by Go et l 99) the

A n n u . R e v . B i o

c h e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2 . D o w n

l o a d e d f r o m w w w . a n n u a l r e v i e

w s . o r g

b y U n

i v e r s i t y o f C o n n e c t i c u t o n 0 8 / 0

1 / 1 3 . F o r p e r s o n a l u s e o n l y .

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A RA 73

speiiy of he ieraio of bidi wih is reepor appears o represeoe of he mehaisms ha preves gee ow bewee he speies sie i

a leas S purpurau ad Srongylocenrou anccanu i is kow haviable hybrids bewee he speies a be formed. However he bidi of S

purpurau fails o iera wih he reepor of Sfranccanu eggs demosraig a barrier a he level of ferilizaio. The sruure of he bidireepor remais o be deermied alhough i has bee parially haraerized(100 101).

I he mouse he proei prese i he zoa pelluida resposible foradhesio has bee isolaed ad is mRNA has bee loed ( 1 1 1 2 102) . Theproei amed Z3 is a glyoproei is O-liked oligosaharide side

hais appear required for adhesio bu aoher appare propery of Z3he iduio of a arosome reaio is los upo removal of he proeiompoe ( 1 1 12) . The preursor of Z3 has a moleular weigh of 4307wih a 22-amio-aid sigal pepide ha would resul i a sereed proei ofmoleular weigh equal o 43943 (102). Alhough he sruures of Z-3-likemoleules i oher speies are o ye kow he DNA probe from mousehybridizes wih he RNA obaied from ovaries of various oher mammaliaspeies suggesive ha osiderable ideiy eiss despie he relaive

speies-speifiiy of adhesio. The DNA probe from mouse does oross-hybridize however wih geomi DNA from various omammaliaspeies Xenopu laev S purpurau Droopla melanogaer ad Raibow rou) hereby demosraig a leas uder he odiios of srigeyused ha Z-3 represes a mammaliaspeifi proei produ ( 102)Clearly he omparaive sruures of Z-3 will be of grea imporae i heargeig of he aive sies of adhesio as well as hose sies ivolved i heiduio of a arosome reaio.

The sperm reepor for Z-3 has o bee uequivoally ideifiedalhough ree repors sugges galaosylrasferase as he reepor moleule( 1 1 12 103) . I has bee suggesed ha aoher zoa pelluid a proei Z2mediaes seodary adhesio of he gamees (104 105). This proei whihwould bid o a sper reepor afer ompleio of he arosome reaiowould be aalogous o he bidi/egg reepor mehaism repored i he seaurhi. The sruure of Z-2 has o ye bee repored bu daa demosraig ha proeiase ihibiors preve his seodary adhesio sugges ha arypsilike ezyme (possibly arosi) is eiher direly or idirely ivolved

i he adhesio proess (105107).

INDUTION O THE AROSOME REATION

iio of Aiv Molul

hose aimals whose spermaozoa possess a arosome he iduio of aarosome reaio is hough o be esseial for ferilizaio. Alhough here

A n n u . R e v .

B i o c h e m . 1 9 8 9 . 5 8 : 7 1 9 - 7 4 2 . D o

w n l o a d e d f r o m w w w . a n n u a l r e v

i e w s . o r g

b y

U n i v e r s i t y o f C o n n e c t i c u t o n 0

8 / 0 1 / 1 3 . F o r p e r s o n a l u s e o n l y .

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GARBER

were iiial sggesios ha his reacio did o omally poceed i resposeo exacellla sigals here is ow compellig reaso o sgges ha

specic molecles associaed wih he egg omally sigal he spemaozoowhe he ime is appopiae for he reacio. There are also repors offolliclar flid ad varios cosies of he female reprodcive ac or ofmeule baed fm umuu pu el aug e du f aacrosome reacio 082). I he sea rchi, safish, ad mose cosiderabe fma abu ee gag meule is ow kow, aug espem recepors remai o be ideified.

he sea rchi a fcoseslfaerich molecle (SG 0) has beeisolaed ha idces he acrosome reacio i a relaively species-specific

fashio (95) ad i has bee sggesed ha he species specificiy is due hesrcre of he sgarslfae polysaccharides (9). SeGall earz (95)uggeed ealy ude a e ema f p did o desroy heabiliy of he remaiig sgar-slfae o idce a acosome reacio. However, i ms be caioed ha i he face of o coceraio-respose daa, iremais possible ha sicie remaiig molecles wih proei sillaaed eed a ee mleue aualy aued e ame ea.I fa Garbers e al (l3) demosraed ha hey cold o remove alldeecable proei from a preparaio by aOH pae eame e fial

prepaaio of eriched fcose-slfae (a eve ha coicides wih Capake ad he acosome reacio) sill elevaed cyclic AM coceraios,

b he relaie poecy decreased appreciably.Z- fm e mue a za elluda pe dued abe w

respec o is role i adhesio of he gamees also has a proposed role i heidcio of he acrosome reacio ( 2). Ulke e ua wih adhesio, however he poei compoe of he molecle appeas reqired foridcio of he acrosome reacio 2). Now ha cDNA cloes areavailable, i will be possible o defe srcre/aciviy relaioships of he

mleule. e appae dua fu f Z a bee aged ude eassmpio ha he pried maerial is acally chemically pre herefore,here sill exiss he possibiliy ha a poe small polypepide coamiaedhe prepaaios ad accoed for he acrosome reacioidcig properesof Z- his, however, seems likely sice he same polypepide wold beexpeced o associae wih oher zoa pellcida glycoproeis Uder heassmpio ha Z- has a dal fcio he i ca be poslaed ha a leaswo recepors exis o he spemaozoo oe ha recogizes he carbohydraeuue ad medae ade ad oher ha recogizes a proei com

poe of Z ad po bidig rasdces a iracelllar sigal(s) haclmiaes i he acosome reacio

he srface here ae similariies bewee Z ad he SG descrbedabe f e ea u. B mleule due e ame ea ad

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BAI O TILIZATION 737

boh appe o do so by prmary effes o a+ upake. ZP medaes eadheree ad FSG appears o possess he same propery based o s abyo ause spem aggluiaio whe mied wih eher live or dead els ad o

he observed adheso o spem els o he ey oa o sea urhi eggs . ohFSG ad Z3 appear o mediae eir ees o he ia el pror o aarosome reaio, ad boh ehbi relaive speies speiiiy wih respe oher aby o due a aosome reao.

I he safsh he arosome reaio apparey ours i respose o woompoes, a suaed gyoproei, ad seroida sapois ( I hasbee show ha a moeue, mos key a ogopepde, aso a paiipae he arosme reao (6 he papao of seroda sapos heaosome reao of sea urhs or oher speies has o bee repoed,

however, Ymaguh e a (7 have suggesed ha spera a poeae hearosome reaio appy idue by FSG e mehasm of hepoeiaio is o ye udersood bu o surpsig sie may o hebiohemial hages aused by FSG (elevaed iraellular pH, ireasedoeraos o yi AMP, mobiizaio o Ca are aso idued by eegg pepides.

Mechanim of Acion

he siga/asduo pahways o mammaia ad omammalia spemaozoa may be simiar if o ideia. No sudy has ye dissoiaed a a+

u aros he plasma membrae fom he arosome reao, ad presumably a primry eve afer reepor oupao, herefore, s Ca+ mobizao I e o be eee y wa meaism e a+ mbzao avaed, ahough i remas possbe ha he sperm reepor, se, s a ohael. he deao of he appare reepor foowed by he log ofhe RNA for he eepor is ow of paramou porae ie buoiued speuaio wil ikey our prior o his aompishme

Neverheess, preseed here s a brief summary of urre views o hemoeuar bass o he aosome reao. Gozaez-Marez & Darszo(8 have repored ha a rapid hyperpolarzaio, possibly due o K+ eu,ours rasiey i respose o FSG he hyperpoarizaio is olowed by adepolarizaio due pipaly o Ca+ upake ( 9 , 0). I has bee suggesed ha possiby a volagedepede Na+ /H+ ehager is reguaed byFSG due o s ees o membrae poeia (8 a rease iraeua pH s aso a pmay espose o FSG aagimai ( has

suggesed ha a

u represes e primay siga for iduio o aarosome reo. He has he posuaed ha he Na+ /+ APase woud beihbied by he elevaed a+ oeraos folowed by eevaed iraeluar Na+ oeraos ha he suggess woud he resu a eu




y .

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ARBR

f H i t H tipr Hwr t H tipr itis scm wd trsrt t d i d c lwr itrclr

I t s rci cycic MP cctrtis r mrkdly ltd irss t FG, bt FG ds t ctit dyt cycs i b crrtis t prsc r bsc f GTP r GTP lgs (19). ict tis f cyclic AMP c b prdcd by gts tt cs Ct it smtz d sic D r rmil c blck cyclic AMPltis i rss t FG it prs tt cyclic AMP is rgltd byC wr i t s rci dirct rglti f sm dytcycls by C s t b dmstrtd. Tis s t t cs fr tsprmtz f ri tr pci wr rgltd dylt cy

clss b rptd ( 221 24). Rctly , ld t l ( 1 25) rptd ZPidcd ltis f cyclic AMP i ms spmtz, dts ct ls ppr t dpd t prsc f xtrclllr C

It sms likly bsd ts rrts, tt rti srltirprsts cmpt f t crsm rcti Hwr, t r f tisclt mdificti i t rcti rmi t b dtmid. ic dddcyclic AMP r cyclic AMP lg will t idc crm rctibt my ccrt t tim rqird fr crm rcti rtipsprylti i t icit, by tlf, fr ti mplgicl t t

ccr.Wit t rizti tt isit pypsts cd rgt C c

ctrti f my cll it bcm f itrt wtr r t f tmtblits cld mdit t FG r P ct C mbilizti.Dmi & Gabrs (26) rtd tt sitl 4,trisst (IP) wstd i s rci sprmtz i rss t FG wr t lon IP lk lao of l MP aad o qu xallua m f r t Icctrti Trfr IP IP r tr isitl plyppts prbblyd t mdit t C mmt coss t sm mmbr. ic tDmi t (127) dmstrtd mrkd icrss i stidicid ftr FG dditi t srmtz. T stidic cid ws sbsqty sw t is by it f ctiti f splips D, btt l if , f pptiic ci i t cm rcti rmi t bdtmid.

Rct dt f Ed t (8) sggt tt gi cltid rgltprtis a ild i t igtrdcti ptwy i ms spmt

. sm cs frm bt rts d itbrts cti G r

Gik prtis (29, ), d V Dp t () rcty istdd qc DA c fr f t pm gi ctidrglty rtis. T idtificti f ts rotis i smtz s

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I OF FRIIZION 739

eled on the use of toxns nd the blty to ADbosylte potens of thsfmly howeve the possble exstee of unne nuleotde eultoypis i speatooa tat ae issii ADisyai mais

isigad 2. I ms pssis i a iddasm ai ad pi iai f ms spmaza t Ganaloues an blok te petusss tox bto 12 ese obsevatose entn sne they would ple the Z eepto nto the unnenuleotde bnd potenoupled fmly of eeptos a dvese fmly thts be to be udestood at the a e .

As oe ompaes the boheml esponses of spematooa to FS ZP-3o the small e peptdes ny smltes e obvous. In the se of FSnd the e peptdes K+ eux ppes to ou ely tht esults n

hypepolton the hne n membe potentl my n tu eulte + ntpote tht esults n lklnton of the ell nteo. C sobled espose to FS o the e peptdes but te moleulamehnss nvolved eue futhe nvestto. It s lkely that the eultoy ess fo C moblaton ae dffeent fo Z and FSGomped to the e peptdes o tht eeptos fo the lnds exst n speeons (e. peptde eeptos n flellum FS nd ZP-3 eeptos nspe hed. he e peptdes do not ndue n osome eton lthouh

they may potentate the eato used by FSG 1 7 . Slly SG doesnot stmulate spem espton tes e the aosoe eton n ftespaton ates ted to deese 33. FSG Z nd the e peptdesuse elevtons of yl AM howeve only te e peptdes snfntlyelevte yl M onenttons. It s lkely then tht yl GM plys key ole n e es of ato of the e peptdes.

MMBRAN RCOGNITION AND GG ACTIATION

One mete membnes ome nto ott the pobblty of ossspeesfetlto neses dtlly he denuded hmste e fo explen be peneted by the spem ells of mny but not ll mmmln spees 1 1 1 2 21 . heefoe eonton snls at ths level ae ppaentlyhly oseved aoss te spees.

ould nd olleues 13 135 hve povded ntun dt on thetvton of Urechis es by osoml extts of spemtoo. he exttsepodue the eets of ntat spetooa n tes of te e atvton he

ablty to atvate es wth suh extats sould allow te pufton ndstutu detenato of bot te effeto and eepto moees nd tedetenato of wete o not sant onsevton exsts oss thespees by the use of eombnnt DA tenoloy




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